t '•.■»•-

JANUARY, 1973

THE

NAUTILUS

Vol. 87

No. 1

A quarterly

devoted to

malacology and

the interests of

conchologists

ifcgical L?.^or5itory

JAN 2 61973

J

Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker.

Editors; R. Tucker Abbott and Charles B. Wurtz

EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of Mollusks

National Museum of Canada

Ottawa, Ontario, Canada K1A-0M8

Dr. William J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 1 0024

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

Dr. James H. McLean

Los Angeles County Museum of Natural History

900 Exposition Boulevard

Los Angeles, California 90007

Dr. Arthur S. Merrill

Biological Laboratory

National Marine Fisheries Service

Oxford, Maryland 21654

Dr. Donald R. Moore

Division of Marine Geology

School of Marine and Atmospheric Science

1 0 Rickenbacker Causeway

Miami, Florida 33149

Dr. Joseph Rosewater

Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. Gilbert L. Voss

Division of Biology

School of Marine and Atmospheric Science

10 Rickenbacker Causeway

Miami, Florida 33149

EDITORS

Dr. R. Tucker Abbott

Delaware Museum of Natural History

Box 3937, GreenvQle, Delaware 19807

Dr. Charles B. Wurtz

3220 Penn Street

Philadelphia, Pennsylvania 19129

Mrs. Horace B. Baker

Business and Subscription Manager

1 1 Chelten Road

Havertown, Pennsylvania 19083

OFFICE OF PUBLICATION

Delaware Museum of Natural History

Kennett Pike, Route 52

Box 3937, Greenville, Delaware 19807

Second Class Postage paid at Wilmington, Delaware

Subscription Price: $7.00 (see inside back cover)

■^jft^!^P>\ THE

,, / ' NAUTILUS

(^ '■-■'-'.'' ,

io- \ Volume 87, number 1 --January 1973

CONTENTS '^

David L. West

Notes on the Development ofColus stimpsoni {Prosohr^nchia: Buccinidae) 1

T. S. Katsigianis and W. N. Harman

Variation in the Radular Teeth of Helisoma anceps(Menke) 5

Branley A. Branson

Significant Pelecypod Records from Oklahoma 8

James B. Sickel

A New Record of Corbicula manilensis (Philipp'i) in the Southern Atlantic Slope Region of Georgia ..11

David Bickel

Non-marine Mollusks and Two New Species of Pisidium from the Tongue River Formation

(Paleocene), North Dakota 13

John W. Ropes and Arthur S. Merrill

To What Extent Do Surf Clams Move? 19

William K. Emerson

LeoGeorgeHertlein(1898-1972), AnObituaryandFinalBibliography 22

John W. Clark, Jr.

The Problem of the Land Snail Genus Rabdotus in Texas Archeological Sites 24

Lothar Forcart

Noteson Veronicellidae and Athoracophoridae in the Field Museum of Natural History, Chicago ... .25

SHORT PAPERS

James L. Murphy

Cerion from an Archeological Site in Northern Ohio 28

J. P. E. Morrison

New Name for a Texas Hvdrobia 28

Dorothy E. Beetle

Phvsa acuta in Virginia 28

R. Tucker Abbott

Spread of Melanoides tuberculata (Thiaridae) 29

Dorothy E. Beetle

Fresh-water MoUusks from Coastal Virginia 29

Book Reviews

(oOPat Halliday,4; A. Feininger and W. K. Emerson, 12;S. and L.Goodman, 22; K.J. Boss, 29.

Obituary and News 27, 30

i

STATEMENT OF OWNERSHIP. MANAGEMENT AND CIR-

CULATION (Required by) Act of October 23, 1962: Section

4396. Title 39. United States Code, and postal regulation

132-622.

1. Title of publication: THE NAUTILUS.

2. Date of filing, September 25, 1972.

3. Frequency of issue: Quarterly (4 per year).

4. Location of known office of publication: Delaware Museum

of Natural History, Kennett Pike, Box 3937, Greenville, De.

19807.

5. Location of the Headquarters or General Business Offices of

the Publishers: Delaware Museum of Natural History, Ken-

nett Pike, Box 3937, Greenville, De. 19807.

6. Names and addresses of publisher, editor, and managing edi-

tor: Publisher, Mrs. Horace Burrington Baker, 11 Chelten

Rd., Havertown, Pa. 19083. Editor, R. Tucker Abbott, Dela-

ware Museum of Natural History, Box 3937, Greenville, De.

19807. Managingeditor, none.

7. Owner: Mrs. Horace Burrington Baker, 11 Chelten Rd.,

Havertown, Pa. 19803.

8. Known bondholders, mortgages, and other security holders

owning or holding 1 percent or more of total amount of

bonds, mortgages or other securities: none.

9. Extent and Nature of Circulation:

A. Total No. Copies Printed (Net Press Run)

B. Paid Circulation

1. Sales through dealers and carriers,

street vendors and counter sales

2. Mail subscriptions

C. Total Paid Circulation

D. Free Distribution (including samples)

by mail, carrier or other means

E. Total Distribution (Sum of C & D)

F. Office use, left-over, unaccounted and

back start subscription copies

G. Total (Sum of E & F) - should equal net

press run shown in A.

I certify that the statements made by me

and complete,

(signed) R. Tucker Abbott, Editor

Average Single

12 Mos. Issue

1,000 1,000

Singing cockles and mussels . . .

Andreas

Feininger

William K. Emerson

Photographs by Andreas Feininger

Text by WilUam K. Emerson

Drawing from two special collections, a noted photographer

and the curator of invertebrates of the American Museum of

Natural History present the shell world well-drawn. A

panoramic view of the many varieties of shells, not just as

scientific objects but as creations of spectacular beauty.

Just how each shell— from the spiny Spondylus dncalif to the

spirally Conus betuUnus— is shaped for its particular

environment is clearly described and it is the

beauty of these fabulous shapes, structures,

and designs that makes Shells a beauty

of a book. Witness shells as you've never

seen them before in a book that creates

almost a mystic satisfaction

and a sense of awe.

A STUDIO BOOK

208 black-and-white and 32 color illustrations

$27.50

'At your bookseller or use this coupon to order

THE "vIkTnG PRESS

Dept. ATE-N,

625 Madison Avenue

New York, NY. 10022

i

Please send me

1 enclose che

copies of Shells (??' $27.50 ea.

money order for $_

total. (If nol completely satisfied, return the book within 10

days for a full refund Please add sales tax where applicable.)

^ Name -

Address_

'Cily

_State_

_2ip_

11

Vol. 87(1)

THE NAUTILUS

NOTES ON THE DEVELOPMENT OF COLUS STIMPSONl

(PROSOBRANCHIA: BUCCINIDAE)'

David L. West

Marine Science Institute

Northeastern University

Nahant, Mass. 01908

ABSTRACT

The buccinid whelk of New England, Colus stimpsoni (March), feeds on the snails,

Littorina littorea (Linnej and Polinices heros (Say), and occasionally on damaged bi-

valves. In the laboratory, C. stimpsoni lays single capsules on rocks, each containing 4,000

■ 5,500 eggs, but with only 1 to 8 eventually developing into young snails. During five

years of observations, capsule-laying occurred from February to May. Development to a

shell length of 5 to 8 mm takes 5 to 6 months.

Colus stimpsoni (Morch, 1 867) ranges from Maine

to North Carolina along the Atlantic coast, and oc-

curs intertidally in its nortliernmost range. For the

past 5 years, a population of C. stimpsoni has been

maintained at the Marine Science Institute. The orig-

inal population and subsequent additions were col-

lected intertidally at Cobscook Bay State Park,

Edmunds, Maine and at Eastport, Maine. A few ob-

servations on the larval development are herein de-

scribed.

Littorina littorea (L.) serves as a food item in the

field and is used to maintain C. stimpsoni in the

laboratory as recorded by Riser (1969). However, it

will feed as well on Polinices heros (Say). The be-

havior of Colus wliile feeding on P. heros is the same

as for L. littorea (for description see Riser, 1969).

However, there is one important additional condition.

When P. heros is moving about and has its metapo-

dium extended, Colus cannot attack. The predator

cannot gain purchase when the prey's metapodium is

extended over the sheU. C. stimpsoni has also been

observed to feed on damaged bivalves and, on one oc-

casion, upon a molting hermit crab. Smaller snails

(less than 7 cm.) have also been observed to feed on

dead, gaping Mercenaria mercenaria (L.) In the labo-

ratory the smaller snails will feed also on the de-

posited egg capsules of its own species. Crowded con-

ditions in the aquaria may lend to this behavior.

Thorson (1935) suggested that the capsules of Colus

(=Sipho) curtus (Jeffreys) were attacked by various

Natica in East Greenland. However, it is possible that

'Contribution number 18 from the Marine Science

Institute.

the capsules were preyed upon by small Colus.

In the laboratory, C. stimpsoni deposits egg cap-

sules throughout the year with increased deposition

from February to May. Capsule-laying can be stimu-

lated by periods of starvation followed by abundant

food (similarly reported for Thais by D'Asaro, 1966).

Capsules are laid singly and attached to rocks, the

sides of aquaria, and occasionally on the shells of liv-

ing C stimpsoni. Each capsule contains approxi-

mately 4,000 to 5,500 eggs.

In surface view, the capsules (Fig. 1 ) are circular to

oval (10 - 16 mm. diameter) and, in side view, are

subhemispherical (4 - 8 mm. in height). The capsule

operculum (exit hole) is fusiform and measures 4-5

mm. in long axis and 3-4 mm. in short axis. When the

capsule is viewed from the side, the operculum is situ-

ated 1/3 - 2/3 the distance from the base to the apex.

Capsules are whitish hyaline in appearance with a yel-

low central portion containing the eggs. The capsule

surface is slightly undulated with irregular striations.

These egg capsules appear very similar to those of

Colus (=Sipho) islandicus (Gmelin) figured by

Thorson (1935: 14, Fig. 5). In cross section the cap-

sule wall is composed of 3 differentially staining lay-

ers and appears similar to the capsule wall of Ur-

osalpinx cinerea (Say) described by Tamarin and Car-

riker(1968).

Individual eggs measure 1 80-1 90 /n in diameter and

are densely packed with large yolk granules.

The number of young snails which develop in a

capsule varies from 1 to 8 (occasionally none) with an

average of 4. The remaining eggs serve as food for the

developing young. There is apparently no relation be-

tween the size of the capsule and the number of snails

THE NAUTILUS

January 1973

Vol. 87(1)

which develop. The largest capsules do not always

contain the largest numbers of developing young.

The first indication of embryos (which are young

veligers) within a capsule still attached to the sub-

strate is what appears to be an enlarged egg, 15-20

days after capsule deposition. The early veliger (figs.

2 & 3), after washing free from the surrounding eggs

and albumin, measure 700-800 /i in length and

400-500 11 across the head vesicle and foot. The

velum is small with the larval kidneys situated slightly

ventral and near the junction of the velum and

visceral mass.

As the veliger continues to ingest the nurse eggs,

the visceral mass increases in size at a rapid rate. The

head vesicle and foot enlarge slightly, and the velum

expands laterally (figs. 4 & 5).

Development is usually delayed at this stage until

all nurse eggs are ingested. However, this is variable

depending on the number of larvae per capsule and

the size of the capsule. In a large capsule with 4

veligers, development will continue before all nurse

eggs are consumed; whereas, in a small capsule with 4

veligers, all nurse eggs will be consumed before devel-

opment proceeds. The veligers may remain in this

Fig. 1. Egg capsule (i]Xo\ti%%\.\m^%on\.

Fig. 2. Early veliger. ventral view.

Fig. 3. Early veliger, lateral view.

Fig. 4. Late veliger, lateral view.

Vol. 87(1)

THE NAUTILUS

stage for 3 to 5 weeks, depending upon temperature

and the relative number of nurse eggs.

As development continues, whether all nurse eggs

are ingested or not, the velum and head vesicle en-

large, and the teloconch is laid down. The body be-

gins to coil, and the mantle cavity enlarges (figs. 6 &

7). The foot is relatively small, and there is no ap-

parent operculum at this stage. The digestive gland

also begins to differentiate at this stage. The ingested

nurse eggs can be seen through the shell.

As tlie foot develops and the operculum is laid

down, the velum enlarges laterally (fig. 8). The head

vesicle does not increase and appears relatively small.

The tentacles appear as small buds at the junction of

the velum and head esicle and the body cannot be

retracted into the shell at this stage.

Development is completed when 2'/i • 3 whorls are

formed. The protoconch is cast off just before or

shortly after the velum disappears. The apex of the

shell has a slight indentation where the protoconch

was (see fig. 10). All nurse eggs are consumed before

development is completed.

Fig. 5. Late veliger, ventral view.

Fig. 6. Early pediveliger, lateral view.

Fig. 7. Early pediveliger, ventral view.

Fig. 8. Pediveliger, ventral view.

THE NAUTILUS

January 1973

Vol. 87(1)

The young snails remain within the capsule until

the capsule operculum ruptures. In cases where this

has been observed, the snails aid in removing the

operculum by bumping against it as they wander

about within the capsule. Apparently the capsule

operculum is softened by external factors such as bac-

teria and fungi. Young snaOs exit when they discover

the hole in the capsule.

The shell of the emerging snail (figs. 9 & 10) is 5-8

mm. in height and 3-4 mm. wide. The aperture has an

undulating margin and about the first ^h of the body

whorl has fine spiral striae.

Total development time, from capsule deposition

to emergence, takes 5 to 6 months in the laboratory.

Development time apparently depends on temper-

ature as well as the number of young and the number

of nurse eggs available to the embryos. In all cases, all

nurse eggs were ingested before emergence. Usually

Figs. 9 & 10. Shell of emerging young Coins

stimp soni.

the young are of different sizes during development

and at emergence. Tlie development of C. stimpsoni is

similar to that of Buccinum undatum (L.) described

by Portmann (1925). Both C. stimpsoni and B. un-

datum emerge with about 3 whorls. However, the em-

bryos of C. stimpsoni are of unequal size whereas

Portmann (1925) indicates that the young of B. un-

datum are of equal size througliout development and

emergence.

LITERATURE CITED

D'Asaro, C. N. 1966. The egg capsules, embryo-

genesis, and early organogenesis of a common oys-

ter predator. Thais haemastoma floridana (Gastro-

poda: Prosobranchia). Bull Mar. Sci. 16: 884-914.

Morch, O. A. L. 1867. Faunula molluscorum Insu-

larum Faeoroensium. Beretning om de hidtil fra

Faeroerne bekjendte Bloddyr. Vidensk. medd. fra

Naturh. Forn. Kjb. 1867 No. 4-7: 67-111.

Portmann, A. 1925. Der Einfiuss der Nahreier auf die

Larvenentwicklung von Buccinum und Purpura. Z.

Morph. Okol. Tiere 3: 526-541.

Riser, N. W. 1969. Feeding behavior of some New

England marine gastropods. The Nautilus 82:

112-113.

Tamarin, A. and M. R. Carriker. 1968. The egg cap-

sule of the muricid gastropod Urosalpinx cinerea:

an integrated study of the wall by ordinary light,

polarized light, and electron microscrope. Jour. Ul-

trastr. Res. 21: 26-40.

Thorson, G. 1935. Studies on the egg-capsules and de-

velopment of Arctic marine prosobranchs. Medd.

Greenland. 10(K5): 1-71.

BOOK REVIEW

SNAILY VERSE by Pat Halliday. 48 pp., numerous

pen drawings; paperback. 1972. Volturna Press,

Wellington House, Peterhead, Aberdeenshire AB4

7JH, Scotland. 30 pence (about 90c U. S.).

This charming booklet of malacological rhymes

and limericks is the funniest to be published in a

century, and contains a remarkable amount of zoolo-

gical information, as well as several parodies of

English poets laureate. The numerous teleological

etchings add to the humor of the 39 conchological

poems.

Otala, Otala, 0, snail of Morocco!

Please answer me tliis, if you will -

Do you suffer a lot during summer sirocco?

Do you wish that the air was more chill?

etc . . .

R. Tucker Abbott

Delaware Museum of Natural History

Vol. 87(1)

THE NAUTILUS

VARIATION IN THE RADULAR TEETH

OF IIEUSOMA ANCEPS (Menke)

T. S. Katsigianis' and W. N. Harnian^

' Graduate student in Malacology, Biology Dept.,SUNY,Oneonta, New York 13820

^ Assoc. Prof., Aquatic Biology, Biology Dept.,SUNY,Oneonta, New York 13820

ABSTRACT

Variation in the structure of the radular teeth in several populations of Helisoma an-

ceps ( Menke j ( Pulmonata: Planorbidae) have been studied in central New York State.

A ratio was calculated, utilizing measurements obtained from a series of the first later-

al radular teeth from individuals in each population. The data show that interpopulation

variation is greater than intrapopulation variation, and that individual variation is less

than intrapopulation variation.

Interpopulation differences in the number and size of the cusps of the radular teeth of

H. anceps were observed. Two populations studied possess first lateral teeth with simple

entocones, mesocones and ectocones; another exhibits an accessory cusp on the distal

portion of the entocone, while a second shows accessory cusps both distal and proximal

on the entocone.

INTRODUCTION

The existence of intraspecific variation within spe-

cies is an important aspect in the study of freshwater

mollusks. Morphological differences of an ecopheno-

typic or genetic nature often are apparent. Since

populations of freshwater animals are normally sepa-

rated in isolated environments, it is relatively easy for

genetic differences to arise. This phenomenon is fur-

ther enhanced by the hermaphroditic characteristic of

pulmonale gastropods, which makes it possible for an

entire population to be founded by a single individual

having only a fraction of the genetic potential of the

parent population. However, due to the relatively

short duration of most freshwater biotopes, tliis gene-

tic variation does not usually proceed to the species

level (Hubendick, 1962).

It has often been logically assumed that the

morphology of the radulae of the omnivorous fresh-

water pulmonates is not subject to severe selection

pressures in the relatively homogeneous microhabitats

in which these organisms live. Therefore, the radular

teeth of these mollusks have long been considered

conservative characters and have always been an im-

portant part of the descriptions of these species [e.g.,

figure 1 illustrates the first lateral tooth of Helisoma

antrosa (=anceps) (Menke) from Baker's (1928)

description of the species] .

A comparative study of the radular teeth of

Lymnaea peregra (MuUer) (Pulmonata: Lymnaeidae)

(Berrie, 1959) showed that although the structure of

the teeth on the same radula, and within the same

population exhibited variation, this was greatly ex-

ceeded by interpopulation differences. In fact, there

was often statistically significant differences between

populations.

This work was initiated to determine if differences

could be observed between the structure of the radu-

lar teeth in four populations of Helisoma anceps

(Menke) (Pulmonata; Planorbidae) occurring within a

20 mile radius of Oneonta, Otsego Co., New York,

U.S.A.

METHODS

Samples were taken at; 1. WUber Reservoir,

Oneonta, Otsego Co., Susquehanna watershed; 2.

Susquehanna River, Oneonta, Otsego Co., Susque-

hanna watershed; 3. Moe Pond, SUNY at Oneonta Bi-

ological Field Station, Cooperstown, Otsego Co.,

Susquehanna watershed; 4. Delaware River, Walton,

Delaware Co., Delaware watershed (figure 2).

Upon collection, snails of approximately the same

size were preserved in 70% ethanol. The radula of

each snail was removed from the buccal mass by heat-

ing it in a concentrated potassium hydroxide solu-

tion. Each radula was then rinsed in water foUowed

by 90% ethanol and mounted unstained in euparol.

An eyepiece micrometer was used to make measure-

ments of individual teeth. All measurements were tak-

en at 970X with an oil immersion lens on a phase

contrast microscope.

THE NAUTILUS

January 1973

Vol. 87(1)

FIGURE I. First lateral tooth of H. anceps showing measurements.

Ec = ectocone. Me = mesocone. En = entocone (modified from

Baker. 19281.

FIGURE II. A portion of New York State showing the location of

each population ■ Wilber Reservoir, ♦ Susquehanna River, • Moe

Pond, A Delaware River.

We calculated the B/E ratio as shown in figure 1,

where B is the distance between the tips of the ecto-

cone and the entocone, and E is the height of the me-

socone from its junction with the ectocone to its tip.

These measurements were taken on the first lateral

teeth of one transverse row of teeth. Teeth from

three rows, five rows apart, were studied on each

radula. Six individuals were chosen from each popula-

tion, therefore, a total of 36 teeth were studied from

each. AH teeth measured were from the central por-

tion of the radulae. It is in this region that the teeth

are fully developed, yet not severely eroded.

These are the same methods utilized by Berrie

(1959). He calculated the B/E and F/E ratios (figure

1) on the first and sixth lateral teeth of each snail but

concluded that the calculafion of the F/E ratio and

the analysis of measurements on the sixth tooth were

not necessary. The F/E ratio varied independently

from the B/E ratio as did the data collected from the

first and sixth lateral teeth.

Figure III. First lateral radular teeth of H. anceps (drawn to scale,

1940x). A = Delware River, B = Moe Pond.

RESULTS

The means, standard errors, and standard devia-

tions of the calculated B/E ratio are given in Table I.

By adopfing the method used by Berrie (1959),

the interpopulation differences have been estimated.

Tliis analysis utilizes the following ratio.

Mean - Mean

Standard Error + Standard Error

1 2

A value of more than two for this ratio indicates a

significant interpopulation difference while a value of

three or more indicates that the populations are ob-

viously different. Table II summarizes these results.

In all cases a significant difference exists between

each population. Althougli the Delaware River popu-

lation is from an entirely different watershed than the

other groups considered, it is extremely different

only from the Wilber Reservoir population. There is

no correlation between the proximity of each popula-

tion and the degree of interpopulation variation.

Tlie Moe Pond population was selected to examine

the intrapopulation variation and the differences that

\'ol. S-( 1 )

THE NAUTILUS

occur bcivvcoii the iccth on the sumo r;idula. To ac-

comphsh this, the stiiiulaid deviation lor the entire

population was compaiei.1 to the standard deviation

ot" tiie teeth on one ladula of one snail trom that

population (Table 3). The standard deviation is much

greater between the teeth from several members of

the population than from the teeth of one individual,

which parallels Berrie's results with /,. peregra. This

indicates less variation between the radular teeth of

each individual than between the radular teeth of the

members of the population.

DISCUSSION

Althougli there were not as many populations con-

sidered in this work as were studied by Berrie ( 1959).

and although the populations are geographically

much closer together, the results obtained are similar.

The data show that interpopulation variation in the

B/E ratio of the first lateral radular teeth of local

populations of H. anceps is greater than intrapopu-

1; tion variation, and that individual variation is less

than intrapopulation variation.

Interesting interpopulation differences in the num-

ber and size of cusps of the first lateral teeth of //.

anceps have been observed. Figure 3 illustrates teeth

from snails in two of the four populations studied.

Each tooth was drawn with the aid of a Whipple

micrometer in a phase contrast microscope at 1940X.

All teeth are drawn to scale. They should be com-

pared with figure I, showing Baker's ( 1928) typical

//. anceps. Note the three cusps on the entocone of

the tooth representing the Moe Pond population. This

is the only population studied which had teeth that

exhibited this morphology. The size of the marginal

cusps on the entocone vary considerably. In some

cases all three cusps are of equal size, in others the

accessory cusps are much smaller. Other variations

within this population include teeth which entirely

lack the marginal cusps on the entocone and others

which have a small cusp between the mesocone and

ectocone. All of these variations may appear on the

same radula. The population from the Delaware River

exhibits teeth with one marginal cusp on the ento-

cone. However, some individuals in that population

show teeth having only the simple ectocone. inesti-

cone. and entocone more typical of the Wilber Reser-

voir and Susquehanna River populations. The latter

populations never exhibited accessory cusps on the

first lateral teeth of their radulae and were essentially

the same as the tooth illustrated in figure I.

LITERATURE CITED

Baker, F. C. 1928. The freshwater mollusca of Wis-

consin, Pt. I. Gastropoda. Wisconsin Gcol. Nat.

Hist. Suiv. Bui. 70(1): I -.507, pi. 1-28.

Berrie. A D. 1959. Variation in the radula of the

freshwater snail Lyninaca peregra (Mliller) from

northwestern Europe. Arkiv. for Zool. 12(27):

391-404.

Hiibeiidick, B. 1962. Aspects on the diversity of the

fresh-water fauna. Oikos.l3(2): 251-261.

Table I. Means, standard errors and standard deviations of

the B/E ratio for the first lateral tooth.

Table II. Differences between the means of the first

lateral teeth for the calculated B/E ratio.

Table III. The variation between the teeth of the six

Moe Pond specimens as compared to one specimen from

Moe Pond. Means, standard errors and standard deviations

are given.

THE NAUTILUS

January 1973

Vol. 87(1)

SIGNIFICANT PELECYPOD RECORDS FROM OKLAHOMA

Branley A. Branson

Eastern Kentucky University

Richmond. Kentuci<y 40475'

ABSTRACT

34 species of Unionidae and one Sphaerium are reported from the Ouachita Mountai:i

river drainages of southeastern Oklahonw, and all show a relationship with the fauna of

the lower Mississippi Valley, rather than the Ozarkian Assemblage. Construction plans for

the Arkansas River threaten these species.

In a previous paper ( 1966), I indicated that the en-

tire moUuscan fauna of the Oklahoma Ozarks (as well

as that of adjacent Kansas) belonged to van der

Schalie's (1950) so-called Ozarkian assemblage, and

that the fauna was probably most closely related to

that of the Cumberland region. However, there is at

least a small segment of this assemblage which reflects

a lower Mississippi Valley relationship (Branson,

1963; 1970).

The fauna of Ouachita Mountain streams, and of

those tlowing through the Gulf Coastal Plains south

of them, in extreme southeastern Oklahoma is an-

other matter. The relationship here is not Ozarkian.

but is primarily with the lower Mississippi Valley, the

southeastern U. S., and, of course, Texas and Louisi-

ana. Isely (1925) indicated some of this in reporting

Ptychobranchus clintuncnsis. P. phaseolus (merely a

southern form of P. fasciolarc, in my opinion) and

Lampsilis hydiana from several Red River tributaries.

Hubricht (1965) and I (1963) reported Eupera

singleyi from southeastern Oklahoma, a species occur-

ring mainly in northern Florida and adjacent Georgia,

and southern Louisiana and Texas.

Over the years, several interesting pelecypod

records have accumulated in the author's files, rec-

ords which lend some weiglit to the comments made

above. Most of these records resulted from collections

secured while I was on the staff of the University of

Oklahoma Biological Station, Willis, Oklahoma, to

which I am indebted for laboratory facilities and vehi-

cles. I am also grateful to Drs. J P. E. Morrison and

Joseph Rosewater, Division of Mollusks, U. S. Nation-

al Museum, for verifications and identifications. The

entire collection (plus many others) herein discussed

' ConI libulion from Fisheries Management.

Department of Biological Sciences.

reside under accession number 23470 at the U. S. Na-

tional Museum. Records for 20 genera and 34 unionid

species, and one sphaeriid species are presented

below.

COLLECTING STATIONS

In the annotations the following form is used:

3(A), 2(B), etc. The Arabic numeral indicates the

number secured, and the letter in parentheses refers

to the station number as listed below: (A) 17 April

1959, Machire Creek, State Hwy 31 crossing, Haskell

Co.. Oklahoma; (B) 3 May 1959, Illinois River, 5 M

east of TahlequalT, Cherokee Co., Oklahoma: (C) 16

May 1960, Grand Lake (Lake O" the Cherokees),

Delaware Co.. Oklahoma; (D) 11 June 1960, Slate

ford. Poteau River, Leflore Co., Oklalioma; (E) 17

April 1959, Mountain Fork River, just above mouth,

McCurtain Co.. Oklahoma; (F) 22 April 1959. Moun-

tain Fork River, T75, R6E. SIO, McCurtain Co.. Ok-

lahoma; (G) 6 July 1952. Little River, between

Broken Bow and Idabel, U. S. 70 crossing. McCurtain

Co., Oklahoma; (H) 24 April 1955, Kiamichi River,

State Hway 2 crossing, McCurtain Co., Oklahoma; (1)

2 September 1955, Lost Creek, near Wyandotte, Ot-

tawa Co., Oklahoma; (J) 16 March 1955, Lake Carl

Blackwell, Payne Co.. Oklahoma; (K) 7 May 1960,

Honey Creek, Camp Classen. Arbuckle Mountains,

Murray Co., Oklahoma; (L) 6 May 1958, Water fall

Creek, just east of Idabel. State Hwy 21, McCurtain

Co., Oklahoma; (M) 28 June I960, Blue River, at

Connoi-ville, Johnston Co., Oklahoma; (N) 6 June

1960, Muddy Boggy Creek, 1 M north of Jessie

Pontotoc Co., Oklahoma; (O) 28 June I960, Moun-

tain Fork River, 6 M southwest of Smithville, State

Hway 21. McCurtain Co., Oklahoma; (P) 24 July

I960. Yashau Creek, 3 M south of Broken Bow, U. S.

70, McCurtain Co., Oklahoma; (Q) II June 1959,

Pennington Creek, 5 M north of Tishamingo,

Johnston Co.. Oklahoma.

Vol. S7(l)

THE NAUTILUS

UNIONIDAE

1. Strophiius uihhilatus Say. Collections: 1 1(B). A

common species througliout the Ozarks, in lowland,

mudbottomed streams with a relatively low profile.

2. Aiiodonta imbecilis Say. Collections: 4(L). Very

dark, chlorophyll-green with indistinct, broad rays;

locally abundant in vegetated backwaters.

3. Anodonta grandis Say. Collections: 1(C); 1(0).

4. Lasmigona complaiiata Barnes. Collections;

1(1). This species, nowhere exceptionally abundant,

gives way in numbers to the next species in southeast-

ern Oklahoma.

5. Lasmigona costata Rafinesque. Collections:

1(0).

6. Throgoiiia verrucosa Rafinesque. Collections;

3(A); 1(E); 1(F); 4(H). A common species through-

out the watersheds of eastern Oklahoma, but absent

in the far west.

7. Fusconaia ccrina (Conrad). Collections: 25(B);

1(M). The distribution of this interesting little

unionid demands more intensive study. The species

was. and probably still is. abundant in the Little Red

River of Arkansas (Call, 1895), southern Louisiana

and Mississippi (Meek, 1896). and clear streams of

east Texas (Murray and Ray, l%8). In most of the

streams of Oklahoma, F. flava outnumbers F. ceriua,

but the latter is quite abundant in the Illinois River

(Ozarks) and common in Blue River (Red River

System). Probably should be synonomized with F.

Jlava. Blue River is a typical stream with regard to

most of the other drainages in its vicinity. It is clear

and spring-fed. and has several relicts of fishes, such

as Etheostoma microperca, Etheostoma radiosum,

Notropis chrysocephahis, and others. The clam fauna,

several representatives of which are listed here, may

likewise be considered in part to be relict.

8. Quadnda pustulosa Lea. Collections: 3(B); 2(F);

3(0).

9. Quadnda cylindrica Say. Collections: 4(F);

l(M). Another relict form and locally common in

Blue River. The next sizable populations of this clam

are found in Mountain Fork and Little Rivers of

McCurtain County.

10. Quadnila melaneiira ( Rafinesque). Collections;

1(H).

11. Plectomerus trapezoides (Lea). Collections:

1(F). This specimen, representing several collected,

was diagnosed as P. dombeyaiiits by the U. S. Nation-

al Museum, a form relegated to synonomy by Murray

and Ray (1968), whom I follow. Strecker (1931)

found this species to be higlily variable in east Texas,

and the same is surely true of adjacent populations in

Oklahoma. P. intcrniptus (Say) is a synonym.

1 2. Amblema perplicata (Conrad). Collections:

2(F). Merely a "washboard" variation of A. costata.

13. Aiid^lcnw costata (Rafinesque). Collections:

1(E); 1(F); 1(H); 1(0). Very common at all locaHfies

(riffles).

14. Plcurobciiia cordatum (Rafinesque). Col-

lections: 1(F); 2(0), but very common. These spec-

imens are of the P. c. catilus (Conrad) form, but I

consider this, as well as P. coccineitm (Conrad), P.

pyramidatum (Lea) and P. c. solidum (Conrad) to be

ecotypes rather than races or spepies.

I 5. Pleurobema aiitillus (Conrad). Collections:

1(F). Merely a southern variety of P. cordatum

16. Elliptio dilatatus (Rafinesque). Collections:

1(D).

1 7. Uniomems tetralasmus (Say). Collections:

3(F); 12(J). In most lakes and ponds of central and

western Oklahoma.

18. Ptychobranchus clintonensis (Simpson). Col-

lections: 5(F) (Common). Simpson (1914) delimited

the range of this southern form as the Little Red

River of Arkansas and, questionably, west to Okla-

homa. It is probably P. phaseolus (HUdreth).

19. Proptera alata (Say). Collections: 1(B); 1(C).

20. Proptera (Leptodeaj ohiensis (Rafinesque).

Collections: 6(C). Questionably distinct from P.

laevissima ( Lea ).

21. Proptera (Leptodeaj fragilis (Rafinesque). Col-

lections: 1(F) (Common).

22. Proptera (Leptodeaj amphichaema (Frierson).

Collections: 1(F). An interesting species, common in

Texas (Murray and Ray, 1968; Strecker, 1931).

23. Proptera ( Proptera j purpcrata (Lamarck). Col-

lections: 7(E); 5(F).

24. Tntncilla pcriniculata (Rafinesque). Collec-

tions; 2(F). I cannot believe this form is distinct from

T. donaciformis ( Lea ).

25. Caninculina pan'a (Barnes). Collections: 1(H);

4(J); 2(L). Common in small creeks and ponds. See

remarks below.

26. Caninculina rc.Yffs«;5/s ( Lea). Collections: l(P)

(but common). Altliougli Murray and Ray (1968)

consider this form a subspecies of C. pan'a, and

Strecker (1931) opinioned that typical L. pan'a was

not known from Texas, I have found both forms to-

gether in northeastern Texas, and Meek (1896) re-

10

THE NAUTILUS

January 1973

Vol. 87 (i:

ported C texasensis to dominate the Caninculina

populations in portions of the abandoned channel of

the St. Francis River near Greenway, Arkansas; both

species were present.

27. Lampsilis anodontoides Lea. Collections: 3(A).

28. Lampsilis liiteola Lamarck. Collections: 2(C);

1(E); 2(K); 6(M); 1(0) (very common).

29. Lampsilis hydianus (Lea). Collections: 2(E);

3(F) (common); 1(0). Tliis species, still common in

the St. Francis and Saline Rivers of Arkansas (Call,

1895), Texas tributaries of the Red River (Strecker,

1931), is characteristic of the lower Mississippi Drain-

age (Louisiana, Texas, Arkansas and neigliboring

states) (Coker, 1919).

30. Lampsilis (Villosa) lienosa (Conrad). Collec-

tions: 2(E); 4(F) (common). This species, with a type

locality in southern Alabama (Clench and Turner,

1956), has a rather wide distribution in the Ohio and

Mississippi basins of Kentucky, Red and Saline rivers

of Texas (Strecker, 1931) (higlily variable), southern

Arkansas (Meek, 1896), and Oklahoma. The habitat is

invariably in mud.

31. Lampsilis cardiiim satuma (Lea). Collections:

1(F) (common). Probably a synonym of Lampsilis

ventricosa (Barnes).

32. Ligumia subrostrata (Say). Collections: 4(F);

1(H).

33. Obovaria jacksoniana Frierson. Collections:

5(F); 1(0) (common in both localities). A species

characteristic of the Pearl and Yalabusha rivers of

Mississippi.

34. Actinonais carinata (Barnes). Collections:

1(C); 4(E); 1(G); 1(0). Specimens from southeastern

Oklahoma are almost invariably referred to A. c.

gibba (Simpson).

SPHAERIIDAE

35. Sphaerium partumeium (Say). Collections:

4(K); 26(M); 2(N); 3(Q). Generally distributed in the

eastern two-thirds of the state.

Discussion

As Meek (18%) indicated in his appraisal of some

pelecypod faunules in southern Arkansas, many of

the species listed herein have a decidedly southern re-

lationship when compared with populations in north-

eastern Oklahoma. Regardless of their systematic dis-

tinctness as full-species, the following nominate forms

indicate southern affinity: Pleurobema antillus,

Ptychobranchus clintonense, Proptera amphichaema.

Proptera purpitrata, Carnnculina texasensis, Lampsilis

hydianus, L. (villosa) lienosa, Actinonais carinata

gibba, and Obovaria jacksoniana.

In view of the extensive habitat modification such

as dam constructions and the opening of the seaway

in the main Arkansas River, it is imperative that much

field work be accomplished in the Ouachita drainages,

particularly in the Poteau, Mountain Fork and Little

Rivers, before it is too late.

LITERATURE CITED

Branson, B. A. 1963. New Mollusk records from Ok-

lahoma and their Zoogeographical Significance.

Trans. Kansas Acad. Sci. 66: 501-512.

Branson, B. A. 1966. A partial Biological Survey of

the Spring River Drainage in Kansas, Oklahoma

and Missouri. Part I, Collecting sites, basic limnolo-

gical data, and mollusks. Proc. Kansas Acad. Sci.

69: 242-293.

Branson, B. A. 1970. Glebula in Oklahoma. Sterkiana

36:25.

Call, R. E. 1895. A study of the Unionidae of Arkan-

sas, with Incidental Reference to their Distribution

in the Mississippi Valley, trans. Acad. Sci. St.

Louis 8: 1-65.

Clench. W. J. and R. D. Turner. 1956. Freshwater

Mollusks of Alabama, Georgia and Florida, from

the Escambia to the Suwannee River. Bull. Fla.

State Mus. 1:97-239.

Coker, R. E. 1919. Freshwater Mussel Industries of

the United States. Bull. Bur. Fish. 36:1 1-89.

Hubricht, L. 1965. Eupera singlegi in Oklahoma.

Nautilus 78:106.

Isely, F. B. 1925. The Freshwater Mussel Fauna of

Eastern Oklahoma. Proc. Okla. Acac. Sci.

4:43-118.

Meek, S. E. 1 896. A List of Fishes and Mollusks Col-

lected in Arkansas and Indian Territory in 1894.

Bull. U. S. Fish. Comm. 1895:341-349.

Murray, H. D. and E. C. Ray. 1968. Checklist of

Freshwater and Land Mollusks of Texas. Sterkiana

30:25-42.

Simpson, C. T. 1914. A Descriptive Catalogue of the

Naiades or Peariy Freshwater Mussels. Bryant

Walker, Detroit, Mich. 1 540 p.

Strecker, J. K. 1931. The Distribution of the Naiades,

or pearly Freshwater Mussels, of Texas. Baylor

Univ. Mus. Sp. Bull. 2:1-71.

Van der Schalie, H. and A. 1950. The Mussels of the

Mississippi River. Amer. Midi. Nat. 44: 448-466.

Vol. .S7(l)

THE NAUTILUS

II

A NEW RECORD OF CORlilCULA MANILENSIS (PHILIPPI) IN THE

SOUTHERN ATLANTIC SLOPE REGION OF GEORGIA

James B. Sickel

Emory University

Department of Biology

Atlanta, Georgia 30322

ABSTRACT

Basal (III previous extensive surveys, it appears that the freshwater clam, Corbicula

manilensis (Philippi), was introduced into the Altamaha River system of Georgia in 1968,

initially into the Ocmulgec River from which it has spread downstream into the Altamaha

River, being found there for the first time in the summer of 1971. This is a first record of

this Asian species occurring in the Southern Atlantic Slope region.

The range of Corbicula manilensis (Philippi) con-

tinues to expand and has finally reached the Southern

Atlantic Slope region, an extension of its range from

the Gulf of Mexico drainage. As of last year, Sinclair

(1971) excluded the Atlantic Slope region from the

known range of Corbicula. In 1968 the author

(Sickel, 1969) made extensive collections of bivalves

in all habitat types from the Altamaha River, Appling

County, Georgia, between River Mile 113 and 118.

Ten species of unionids were found but no Corbicula.

Recent collections have shown Corbicula is now in

the Altamaha River system, both in the Ocmulgee

River and the Altamaha River itself.

On October 15, 1971, while collecting in the Alta-

maha system, the author found one Corbicula mani-

lensis shell on a sandbar in the Ocmulgee River three

miles from its confluence with the Oconee River at

which point the Altamaha begins. Shell annual rings

are not absolute determinants of age since they are

frequently indistinct, but based on what appear to be

annual rings, i.e., variations in the width of concentric

ridges, and the size, 10.0 x 8.6 x 6.3 mm., the shell is

estimated to be two years old and to have died in the

spring of 1971.

While collecting in the Flint River, Crawford

County, on November 4, 1971, the author found an

abundance of Corbicula manilensis. The Flint River is

a part of the Apalachicola River system in the Gulf

drainage and is not connected with the Altamaha

system, but tributaries of each reach within a mile of

the other. The Flint River population was in coarse

sand along the upstream end of most sandbars where

the sandy bottom was relatively stable. Using a view-

ing box in shallow water and viewing a 1 sq. ft. area.

10 to 15 adult Corbicula could be seen in any square

foot area chosen within the favorable habitat. Along

the middle and downstream reaches of the sandbars

fewer Corbicula were found, and scattered among

them were the following unionids: Lampsilis ano-

dontoides floridensis (Lea), Lampsilis binominatus

(Simp.), and Quincuncina infucata {Comad). Wliere the

Corbicula were most dense there were no unionids,

even though the habitat appeared suitable. This indi-

cates some form of competition, and it is unlikely to

be simply spatial competition since the size of Cor-

bicula and its density did not appear to be great

enougli to exclude the much larger unionids.

The adult Corbicula in the Flint River were still

expelling veliger larvae on the date collected. This ex-

tended breeding season is probably common in south-

ern rivers and means that control measures taken by

industry will have to be extended correspondingly.

In the Altamaha River, Appling County, at one of

the same sites where the author collected unionids in

1968, five juvenile Corbicula were found on Decem-

ber 15, 1971. Using a modified Peterson dredge, sam-

ples were taken on sandy and muddy bottom. The

Corbicula were found only in coarse sand near the

bank where the sand grades into mud. Six grab sam-

ples taken at one site produced five Corbicula which

ranged in length from 6.0 mm. to 7.8 mm. Each still

retained a byssal thread and had the characteristic

purple rays of the juveniles. They were obviously in

their first year of development.

The first report of Corbicula in the Altamaha came

from Drs. Donald C. Scott and James Schindler, Uni-

versity of Georgia (personal communication), who

found several young specimens while sampling in

i:

THE NAUTILUS

January 1 973

Vol. 87(1)

August, 1971, near the same area where the author

collected.

Corbicula nianileiisis is not yet abundant in the

Altamalia River. No adults over 2 years old have been

found. Many sites have been examined in the Alta-

maha and lower Ocmulgee in which Corbicula has not

been found.

The author proposes that Corbicula was intro-

duced into the Ocmulgee River by overland transfer

from the Flint River in 1968 or 1969. This transfer

could have occurred by fishermen who frequent both

rivers and use mussels for bait or accidentally in the

sand dumped into boats, possibly from anchors or

nets. One fertile adult could produce enougli off-

spring to establish a breeding population from which

veligers would be carried downstream to populate the

entire Altamalia to the Atlantic. It will be only a mat-

ter of time before Corbicula moves northward to in-

habit the remainder of the Southern Atlantic Slope

region.

Appreciation is expressed to John G. Adams and

the Georgia Power Company for the boat, collecting

equipment, and facilities so graciously provided for

the Altamaha River collections.

LITERATURE CITED

Sickcl, J. B. 1969. A survey of the mussel populations

(Unionidae) and Protozoa of the Altamaha River

with reference to their use in monitoring environ-

mental changes. MS Thesis. Emory University,

Atlanta.

Sinclair, R. M. l^?!. Annotated bibliography on the

exotic bivalve Corbicula in North America,

1900-1971. Sterkiana 43:11-18.

BOOK REVIEW

SHELLS. Photography by Andreas Feininger, text by

William K. Emerson. A Studio Book, the Vikmg

Press, New York. 295 pp. (including the plates),

175 pis. (including 32 in color), 7 numbered and

194 unnumbered figs. 1972. $27.50.

This is a large format (12 3/8" x 10") book of

photographs of marine moUusk shells, chosen by a

distinguished photographer "not from the point of

view of the scientist but from that of the engineer in-

trigued by structure and the interdependence of func-

tion and form, and from the point of view of the

artist who is fascinated by shape and design." It is

said to be "a shell-appreciation book" differing from

other shell books which "are too much preoccupied

with facts."

Most of the photographs are indeed spectacular.

They are grouped around chapters with the following

lieadings: Forms of the Sea; an Infinite Variety of

Shapes; Carrier Shells; the Structure of Sliells; the

Spiral; the Color of Shells; the Surface Texture of

Shells; Knobs and Spines, and Ornamental Design.

Magnifications vary greatly, and the shells are orient-

ed in all sorts of positions, resting im sand, gravel,

boulders, or glass bespattered with water, sometimes

with a background of sky or cloud. Not a single shell

is shown alive or in its natural habitat.

There is an elementary Introduction to malacology

and the text following it parallels fairly well the se-

quence of photographs, diverting in places to such

subjects as fluorescence. There is no first-hand in-

formation about the biology of mollusks. The plate

captions consist solely of the scientific name and a

reference number relating it to the systematically ar-

ranged catalog at the end. Only by referring to the

latter would the novice learn that the Haliotis shells

shown on pis. 1 19 and 1 20 are polished. Most of the

shells in the catalog are refigured at natural size and

in more conventional orientations. There are also a

glossary, bibliography and index.

The Foreward unwisely states that the text is

"scientifically accurate down to the last detail." The

phyletic successions of marine bivalves and snails

(figs. 2 and 3) are grossly misleading. Two entirely

different species on pi. 101 each are identified as

Comis maldivus Hwass. Various scientific names are

misattributed to authors, and various adjectival speci-

fic names disagree in gender with the generic names.

Vermicularia is wrongly included in the Vernietidae.

How well does this book achieve its objective of

being "a complement to all other kinds of shell

books"? It differs little in inieiii from The Shell; Five

Hundred Million Years of Inspired Design by Stix.

Stix and Abbott (l')68). Both of these are modern

photographic counterparts of the type of shell book

begun by Buonanni in 1084: Rccrcario Mentis et

Oculi. Thus there is nothing unique about the

Feininger-Emcrson book; the photographs are its

iiKist redeeming feature.

Robert Robertson

Academy of Natural Sciences of Philadelphia

Vol. 87(1)

THE NAUTILUS

13

NON-MARINE MOLLUSKS AND TWO NEW SPECIES OF PISIDIUM FROM THE TONGUE

RIVER FORMATION (PALEOCENE), NORTH DAKOTA.

David Bickel

Science Division

Minot State College

Minot, North Dakota 58701

ABSTRACT

Fifteen taxa of Paleocene freshwater Mollusca were recovered from relatively soft

carbonate sediments of the Tongue River Fommtion (Paleocene: Fort Union Group). The

beds, exposed at two sites in southern Ward County, North Dakota, apparently accumu-

lated in a small hard-water lake. The faunule is discussed and illustrated. Unlike the few

previously described species of Pisidium from early Tertiary rocks, preservation of Pisi-

dium wardensis and Pisidium russelli n. spp. is good enough to permit detailed description

of their hinge morphology.

INTRODUCTION

The Late Cretaceous and early Tertiary deposits in

the North Dakota portion of the Williston Basin

record a conformable transition from marine to

brackish water to non-marine sediments. The Fort

Union Group includes Paleocene rocks in North

Dakota and occurs in the western half of the state

and on an outlier in northeastern North Dakota.

These strata in North Dakota include from base to

top the Tullock. Ludlow, Cannonball, Tongue River,

and Sentinel Butte Formations. All are of alluvial

origin except for the Cannonball Formation which

apparently accumulated in an early Tertiary relic or

rejuvenation of the Cretaceous seaway that crossed

the midcontinent region. The Tongue River Forma-

tion was deposited over Tullock, Ludlow, and Can-

nonball sediments by a system of streams flowing

from a western source with low gradients and veloci-

ties (Royce, 1970). Tongue River sediments are most-

ly poorly consolidated yellowish sUts and clays with

lesser amounts of sand, lignite coal, and limestone.

Thick units that formed in backswamp and floodplain

environments characterize the formation. Thick

deposits of glacial drift cover pre-Pleistocene rocks

east and north of the Missouri River and bedrock out-

crops are small and generally limited to the larger

stream valleys, shallow road cuts, and strip mines.

Royce (1970) summarizes the sedimentary history of

the Tongue River and Sentinel Butte Formations.

A higlily fossiliferous exposure of the Tongue

River Formation (Paleocene) was brought to my at-

tention in January, 1971, by Mr. Fred Ballentyne of

Unit Paleocene: Tongue River Formation

4 Buff to light gray limestone, friable, blocky;

lower half coarse grained near base, finer

textured toward top; mollusks abundant in

Umonitic band near middle.

3 Dark gray calcareous shale, soft, strongly fissOe

2 Light gray to buff limestone, blocky; moderately

hard at base; moderately friable above.

1 Brown to gray lignitic clay, weakly calcareous;

base not exposed.

Thickness (Feet)

11.3

1.0

6.2

0.3

TOTAL 18.8 ft.

THE NAUTILUS

Jamuirv I'^V.^

Vol. 87(1)

Sawyer, North Dakota. Most of the fossils were col-

lected that month. The faunule from this locality is

notable for the number of ta.xa present and the occur-

rence of new species of Pisidiiim. The outcrop is

located about 8 miles south of Sawyer. Ward County.

North Dakota m NW'/a. sec. 10, T15:N. R81W. The

exposure is an isolated man-made cut on the west

wall of a shallow coulee at about 1 800 ft. above sea

level. A stratigraphic section is described below:

Bedrock is completely covered on both sides ot

the outcrop; however, more typical Tongue River

clastic sediments are exposed at the same elevation in

an abandoned strip mine about 100 yds. to the west.

Carbonate sediments with similar lithology and strati-

graphy are poorly exposed at the same elevation

about 1 mi. to the east in SW'/i, NWii. sec. 20 of the

same township, but any lateral transition of this se-

quence into silt-clay facies that outcrop nearby is also

covered.

The stratigraphic position of these beds in the

Tongue River Formation cannot be accurately fixed

at present. Lemke (1960. p. 30, 3'i) places the

Tongue River-Cannonball contact at an elevation of

1540 ft. along the Souris River 8 to 9 mi. to the

northeast. Allowing for a sliglit northeast dip in this

area, these exposures are probably 200-250 t~t. above

the base of the Tongue River Formation.

The deposit apparently accumulated in a shallow

tloodplain depression or series of depressions that

held a permanent hard water lake. The lake formed

over clastic and organic backswamp sediments and

maintained a permanent water level long enougli for

diverse moUuscan and ostracode assemblages to be-

come established. Pisidiwn, small amnicolids, and

ostracodes suggest a lacustrine environment. Vivi-

parids are prominent elements in the Fort Union

fauna and these snails probably inhabited slower mov-

ing streams as well as standing water in backswamp

areas.

SYSTEMATIC PALEONTOLOGY

Family Uniunidae

Plesielliptio priscus (Meek and Hayden, 1 856)

Fig. 16

Uiiio priscus Meek and Hayden, 1856, Proc. Acad.

Nat. Sci. Philadelphia 8; 117.

Unionid mussels are common and generally occur

with both valves connected, aligned, and closed. Pres-

ervation in place is often excellent except for frac-

tures induced by compaction, but specimens crumble

and exfoliate during extraction even v<.'ith careful field

and laboratory techniques. One specimen is intact

enough to show both the double looped beak sculp-

ture o\' PIcsiclllptli) and the proportions off. priscus.

Its measurements are: length 78.0 mm., heiglit 45.0

mm., width 21 mm. Three valve fragments have the

concentric looped ornamentation o\ Plcsiclliprid but

are too incomplete for specific determination. About

30 other large fragments of exfoliated hinge areas

cannot be identified to genus.

Family Corbulidae

Bicorbula niuctriturniis (Meek and Hayden. 1856)

Fig. 1 5

Corhula niactritcnuis Meek and Hayden. 1856. Proc.

Acad. Nat. Sci. Philadelphia 8; 117.

Bicorbula mactriforniis is the most common bi-

valve encountered. The numerous specimens ex-

amined range in length from a few millimeters to a

maximum of 23 mm. Outlines vary from sub-ovate to

sub-trigonal with most valves appearing somewhat tri-

angular. Sub-trigonal valves seem to have greater

heights in proportion to length than do ovate speci-

mens. The genus is inequivalved. Dentition is well pre-

served and all intact specimens are easily distin-

guished from other bivalves.

Family Pisidiidae

Hinge tooth terminology in the following descrip-

tions is that of Herrington (1962. p. 1 1 ) with addi-

tion of the abbreviations RV and LV for right and

left valves.

PisiJiuni wardcnsis new species

Figs. 9. 10

Dcscriptidn: Shell small, inflated, oulline ovate;

w;ills moderately thick; beaks prominent, sliglitly to

distinctly raised, sub-central toward posterior; pos-

terior margin rounded to slightly truncate; anterior

margin rounded, narrower than posterior margin; dor-

sal margin broadly angular at umbo, forms slight

rounded angle with posterior margin, smoothly

curved into anterior margin; ventral margin convex;

striae fine, evenly-spaced; growth pauses shallow in-

dentations or steps; hinge narrow; lateral teeth long,

covering about four-fifths of shell length; inner ante-

rior lateral tooth Al (RV) pointed, triangular; outer

anterior lateral AlII (K\) linear, narrow, near distal

hinge margin; anterior lateral All (LV) straight, tri-

angular, cusp distal; posterit)r lateral teeth PI. PIM.

(RV) with short cusps; posterior lateral tooth PIl

(LV) toward proximal hinge margin, cusp prominent;

\ol. 87(1)

THE NAUTILUS

15

FIG. 1. Lioplacodes nebrascensis 'pxo&\KVA(White)X I: FIG. 2. Lioplacodcs mariana YeiiX I:FIG. 3. Lioplacodes

limnaeformis (Meek ami Haydenj, a slender specimen X 1.5: FIG. 4. Vivipaius raynoldsanus Meek and Hayden,

specimen slightly compressed into plane oj illustration X I: FIG. 5. Campeloma nebiasceiisis whitei Russell XI;

FIG. 6. Campeloma nebrascensis (Meek and Harden) X I: FIG. 7. Vivipaius peculiaris (Meek and Haydenj X I:

FIG. 8. Gyraulus cf. G. militaris (White) X 10: FIG. 9. Pisidium wardensis /). sp. interior of Holotype (USNM

180367) X 10: FIG. 10. Pisidium wardensis n. sp., exterior of Holotype X 10: FIG. 11. Pisidium russelli «. sp..

interior of Holotype (USNM 180370) X 10: FIG. 12. Pisidium russelii n. sp.. exterior ofHohitypc X 10: FIG. 13.

Hydrobia warrenana (Meek and Hayden) X 4: FIG. 14. Hydrobia anthonyi (Meek and Hayden) X 4: FIG. 15.

Bicorbula mactriformis (Meek and Harden), interior view X 2: FIG. 16. Plesielliptio priscus (Meek and Hayden) X

'A.

16

THE NAUTILUS

January 1973

Vol. 87(1)

cardinal tooth C3 (RV) distinctly curved, enlarged

posteriorally; cardinal teeth C2, C4 (LV) slender,

rouglily parallel; C2 curved, shorter than C4; C4

slightly curved or distinctly curved on anterior half.

Tvpes and measurements: Types are deposited in

the United States National Museum of Natural His-

tory. Holotype (USNM 180367) length 2.0 mm.,

height 1.8 mm.; Paratype (USNM 180368) length 1.9

mm., heiglit 1.8 mm.; Paratype (USNM 180369)

length 1.6 mm., heiglit 1.5 mm.

Type locality: About 8 mi. south of Sawyer.

North Dakota, NW'/j, NW>/4, sec. 19, T152N, R81W.

Paleocene; Tongue I^ver Formation. Collected Jan.

1971.

Remarks: The 40 specimens examined vary in out-

line and the degree to wliich beaks project above the

hinge line. Smaller specimens tend to appear quite in-

flated. Pisklium wardensis is distinguished from other

Late Cretaceous and early Tertiary pisidid clams by

its inflated ovate shell, broad and rather prominent

beaks, and small size. The species is named for Ward

County, North Dakota.

Pisidium russelli new species

Figs. 11, 12

Description: Shell moderately small, moderately

inflated, outline ovate; beaks prominent, raised above

hinge, situated about one third of shell length from

posterior edge; anterior and posterior valve margins

rounded; dorsal margin sliglitly convex, rounded pos-

teriorally, forms rounded angle with anterior margin;

ventral margin convex; striae fine, evenly spaced;

hinge narrow; lateral teeth long, covering about

three-fourths of sheU length; anterior inner lateral

tooth Al (RV) long, cusp toward distal end; anterior

outer lateral AIII generally parallel to AL prominent;

anterior lateral All (LV) on inner margin of hinge,

straight, cusp rounded and distal; posterior lateral

teeth PL PHI (RV) slender, low, parallel, depression

for opposing tooth long, narrow; posterior lateral

tooth PU (LV) slender, low cusp rounded; cardinal

tooth C3 (RV) distinctly curved, enlarged poster-

iorally; cardinal teeth C2, C4, (LV) parallel, very

sliglitly curved, C2 sliglitly shorter than C4.

Types and measurements: Types are deposited in

the United States National Museum of Natural His-

tory. Holotype (USNM 180370) length 2.85 mm.,

heiglit 2.4 mm.; Paratype (USNM 180371) length

1.75 mm., height 1.55 mm.; Paratype (180372)

length 1.8 mm., height 1.5 mm.

Type locality: About 8 mi. south of Sawyer, Ward

County, North Dakota, NW'/*, NWi4, sec. 19. T152N.

R81W. Paleocene: Tongue River Formation. Col-

lected Jan. 1971.

Remarks: Pisidium russelli resembles P. wardensis

in outline and hinge structure. It is distinguished from

that species by its less inflated shell, sliglitly angular

dorsal-anterior margin, more posteriorally situated

beaks, and parallel cardinal teeth on the left valve.

The name honors Dr. Loris S. Russell.

Family Viviparidae

Vivipanis peculiaris (Meek and Hayden. 1856)

Fig. 7

Paludina peculiaris Meek and Hayden. 1856, Proc.

Acad. Nat. Sci. Philadelphia 8: 122.

The species is characterized by having a thin

trocliiform shell, about 5Vi flattened and slightly con-

vex whoris, fine growth Hues, and very indistinct spi-

ral ridges. The body wliori is angular at its base and

the aperture is ovate to sub-quadrate. Three incom-

plete specimens were found at this site. Two speci-

mens measure: heiglit 18.6 mm. (juvenile whods miss-

ing) and 18.2 mm.; width 16.5 mm. and 13.4 mm.;

aperture heiglit 1 1 mm. and 10 mm.

Vivipanis planolatcrc Russell from the Paleocene

of Alberta seems to be a junior synonym of V. peculi-

aris. The relarionship of V. peculiaris to the similar

Cretaceous species. V. ro/irat// (Meek and Hayden). is

uncertain, and the name V. conradi, has priority if

the two shells are conspecific.

Vivipanis raynoldsanus Meek and Hayden. 1861

Fig. 4

Meek and Hayden, 1861, Proc. Acad. Nat. Sci. Piula-

delphia 13: 446.

Large adult specimens are crushed but several

body whorl fragments suggest maximum shell widths

of over 25 mm. The largest intact shells have heiglits

of about 30 mm. and widths of about 20 mm. Four

of eleven specimens have somewhat flattened whorls,

a rounded angle where the side and base of the body

whori meet, and whods with sliglit shoulders. Such

specimens have an almost turbonate appearance.

Meek (1876, p. 584) mentions specimens of V. ray-

noldsanus with slightly flattened and shouldered

whorls.

On the basis of crushed and fragmcntaiy speci-

mens of V. peculiaris and V. raynoldsanus. 1 incor-

rectly reported V. trochiformis (Meek and Hayden)

and V. retusus (Meek and Hayden) from this locality

in an abstract (Bickel and Hall, 1971).

Vol. 87(1)

THE NAUTILUS

17

Canipdoina iicbrasccnsis (Meek and Hayden, 1 856)

Fig. 6

Buliimis ncbrascensis Meek and Hayden. I 856. Proc.

Acad. Nat. Sci. Philadelphia 8: 118.

Two specimens from this site have the convex

whorls, intlated body whorl, slight shoulder, and

raised spiral lines of the typical form of C. ncbrascen-

sis. Meek (1876. p. 586) states that on some speci-

mens "the lower whorl shows faint traces of an ob-

scure revolving ridge just below the suture." This

characteristic occurs commonly on specimens 1 have

seen from other localities in the Williston Basin and

Delimata (1969, p. 29) notes it on material from the

Little Missouri Badlands. Shoulders, when present,

are less prominent than on C. ncbrascensis whitci

Russell. The two specimens measure: heiglit 27.0

mm. and 16.6 mm.; width 18.0 mm. and 12.8 mm.;

aperture heiglit 13.3 mm. and 9.0 mm.

Campcloma ncbrascensis whitci Russell, 1931

Fig. 5

RusseU, 1931, Trans. Roy. Soc. Canada, ser. 3, 25

(sec. 4): 12, pL 2, fig. 1.

This shell is the most common of the larger vivi-

parids encountered with 17 fairly intact specimens

being recovered. Many shells are crushed by sediment

compaction and others were destroyed during col-

lection. Campcloma ncbrascensis whitci differs from

typical C. ncbrascensis in having a prominent

shoulder at the top of the penultimate and body

whorl and a broad, shallow depression or sinus just

below the shoulder. Five specimens lack raised spiral

ornamentation. This form occurs with typical C.

nebrascensis and is easily distinguished from it, sug-

gesting that C. ncbrascensis whitci may be a distinct

species.

Lioplacodes Umnaeformis (Meek and Hayden, 1856)

Fig. 3

Bulimus Umnaeformis Meek and Hayden. 1856, Proc.

Acad. Nat. Sci. Philadelphia 8: 118.

Specimens are fairly common at the exposure and

generally well preserved. Most specimens have a

slightly convex spire but occasionally sheUs (4 of 50

specimens) have relatively straight-sided spires. Three

specimens have the following measurements; heiglit

17.0 mra, 15.5 mm., 12.2 mm.; width 7.9 mm., 7.5

mm., 5.5 mm.; aperture height 7.2 mm., 6.8 mm., 5.8

mm.

Lioplacodes mariana Yen, 1946

Fig. 2

Yen, 1946, Anier. Jour. Sci.. 244: 44-45, pi. 1, figs.

5a- 5c.

Five specimens are referred to this species which is

distinguished from L. nebrascensis by having a slender

spire, less intlated body whorl, more loosely coiled

whorls, and a smaller aperture. The upper two-thirds

of each whorl is sliglitly convex and the lower third is

strongly convex where it meets the lower suture. Two

specimens measure: height 27.5 mm. and 26.5 mm.;

width 14.0 mm. and 12.7 mm.; aperture height 11.0

mm. and 1 0.0 mm.

Lioplacodes nebrascensis producta ( Wliite, 1 883)

Fig. 1

Campcloma producta White, 1883, U. S. Natl. Mus.

Proc, 3: 97, pi. 3, figs. 7-9.

Over 100 specimens were examined and they vary

in spire heiglit, convexity of whorls, and degree of

ornamentation. The original description of L. nc-

brascensis (Meek and Hayden, 1856, p. 124) depicts

the variability of this species, and Russell (1931) con-

siders L. ncbrascensis producta and typical L. ne-

brascensis end members in the range of variation in

one species. The trinomial is used here as a form

name. Many of the specimens are ornamented with

fine spiral raised lines that are occasionally developed

enough to make juvenile and intermediate whorls ap-

pear liglitly carinate. Liglitly carinate shells resemble

L. tenuicarinata (Meek and Hayden) but lack the

strongly keeled and convex whorls of that species.

About one-fifth of the material has the tall, slender

spire and whorl morphology of L. mariana but retains

the fuUer body whorl and larger aperture of L. nc-

brascensis producta.

Family Planorbidae

Gyraulus cf. G. militaris (White, 1 880)

Fig. 8

Planorbis militaris White. 1880, Proc. U. S. Natl. Mus.

3: 159.

Two specimens of small planorbids, both crushed

along the axis of coiling, resemble G. militaris. The

illustrated specimen shows the proportions of G. mili-

taris and has the deep umbilicus of that species. Shell

widths of the two specimens are 1.9 mm. and 1.8

mm.

THE NAUTILUS

January 1973

Vol. 87 ( 1 )

Family Amnicolidae

Hydrobia anthonyi (Meek and Hayden, 1 856 )

Fig. 14

Melania anthonyi Meek and Hayden, 1856, Proc.

Acad. Nat. Sci. Philadelphia 8: 124.

On large specimens, penultimate and body whorls

are somewhat more convex than the immature

whorls. Intermediate sized shells tend to have sliglitly

convex whorls and well impressed sutures. Smaller

specimens generally have subrounded apertures, while

apertures on larger individuals tend to be more elon-

gate. Persistant and very fine spiral striae are present

on adult whorls but are absent or extremely fine on

juvenile whorls. Fifty-seven specimens were exam-

ined. One large and one intermediate-sized specimen

measure: heiglit 6.3 mm., 3.4 mm.; width 33 mm.,

1.8 mm.; aperture heiglrt 2.8 mm., I.l mm.

Hydrobia warrenana (Meek and Hayden, 1 857)

Fig. 13

Melania warrenana Meek and Hayden. 1857, Proc.

Acad. Nat. Sci. Philadelpliia 9: 137.

Hydrobia warrenana is distinguished by its conical

spire, 6 to 7 sliglitly convex whorls, and a body whorl

that forms a rounded angle with the base. This angle

tends to be sharply defined on immature shells but

more rounded on large specimens. The aperture is

subquadrate. Two large specimens measure; length

5.6 mm., 4.8 mm.; width 2.6 mm., 2.2 mm.; aperture

heiglit 2.0 mm., 1.7 mm. Most of the 30 specimens

have lengths of less than 3.0 mm.

Family Physidae

Pliysa sp.

One fragmented and apparently immature speci-

men of this genus was found. The apex and aperture

periphery are missing and specimen length is 7.0 mm.

LITERATURE CITED

Bickel, D., and Hall, D. H. 1971. Paleocenc fresh-

water Mollusca from southern Ward County,

North Dakota (abstr.). North Dakota Acad. Sci

Proc.pt. 1,25: 3.

Delimata, J. J. 1969. Fort Union (Paleocenc) mol-

lusks from southern Golden Valley and south-

eastern Billings Counties, North Dakota. MS

Thesis. Univ. North Dakota. 73 p.. 6 pis. (unpubl..

typewritten).

Herrington, H. B. 1962. A revision of the Sphaeriidae

of North America (Mollusca: Pelecypoda). Mus.

Zool. Univ. Michigan Misc. Publ. 118, 74 p., 7 pis.

Lemke. R. W. 1960. Geology of the Souris River area.

North Dakota. U. S. Geol. Surv. Prof. Pap. 325,

138 p.

Meek. F. B. 1876. A report on the invertebrate Creta-

ceous and Tertiary fossils of the Upper Missouri

country. U. S. Geol. Geogr. Surv. Terr. (Hayden

Survey) 9: 629 p., 45 pis.

Meek, F. B. and Hayden, F. V. 1856. Descriptions of

new species of Acephala and Gastropoda . . . Proc.

Acad. Nat. Sci. Philadelphia 8: 11 1-125.

Meek, F. B., and Hayden, F. V. 1857. Description of

new species and genera of fossils, collected by Dr.

F. V. Hayden in Nebraska Territory . . . Proc.

Acad. Nat. Sci. Philadelphia 9: 117-148.

Meek, F. B., and Hayden. F. V. 1861. Descriptions of

new Lower Silurian (Primordial). Jurassic. Creta-

ceous, and Tertiary fossils, collected in Nebraska .

. . Proc. Acad. Nat. Sci. Philadelphia 13: 415-447.

Royce, C. F., Jr. 1970. A sedinicntologic analysis of

the Tongue River-Sentinel Butte intei-val (Paleo-

cenc) of the Williston Basin, western North

Dakota. Sedimentary Geol. 4: 19-80.

Russell, L. S. 1931. Mollusca from the Upper Creta-

ceous and Lower Tertiary of Alberta. Roy. Soc.

Canada Trans. Ser. 3, 25 (sec. 4): 9-19, 2 pis.

White, C. A. 1880. Descriptions of new invertebrate

fossils from the Mesozoic and Cenozoic rocks of

Arkansas. Wyoming, Colorado, and Utali. Proc. U.

S.Natl. Mus. 3: 157-162.

Wliite, C. A. 1883. New molluscan forms from the

Laramie and Green River Groups, with discussion

of some associated forms heretofore known. Proc.

U. S. Natl. Mus. 5: 94-99, pi. 3-4.

Yen. T.-C. 1946. Paleocenc freshwater molhisks from

Sheridan County. Wyoming. Amer. Jour. Sci. 244:

41-48. 1 pi.

Vol. 87(1)

THE NAUTILUS

19

TO WHAT EXTENT DO SURF CLAMS MOVE?

John W. Ropes and Arthur S. Merrill

National Marine Fisheries Service

Middle Atlantic Coastal Fisheries Center

Oxford, Maryland 21654

ABSTRACT

Locomotion by surf clams, Spisula solidissima, includes rapid burrowing, crawling, and

leaping from the bottom. Observations of these activities led to the suspicion that some

movement over the sea floor may occur. We found that the usual response of surf clams

removed from a burrow is rapid re-entry into the substrate. Occasional leaping, when

seen, was usually by recently disturbed, small and very young specimens. They propelled

themselves very short distances and immediately burrowed upon landing. Unusual circum-

stances seemed necessary to elicit the leaping response and it was considered a relatively

ineffective means of locomotion

Surf clams {Spisula solidissima (Dillwyn)), like

many other bivalves, are considered to be benthic, in-

faunal, and sedentary. They are normally found

buried and expose only their siphon tips at the sea

bottom. In an account of a trip in the deep sub-

mersible Alvin. Edwards and Emery (1968) made a

brief statement about seeing a 6-inch surf clam "leap-

ing" across the bottom. Leaping, a behavior that

seems to contradict the usual sluggish reactions of

surf clams, may be a hitherto unsuspected means of

escaping hazards which threaten survival. It may also

be an exceptional behavior to which an observer

miglit attach too great a significance. The purpose of

this paper is to review the literature on the movement

of surf clams and report on our observations of its

activities.

Three general locomotor activities have been re-

ported for surf clams: gliding, crawling, and burrow-

ing (Belding, 1910; Ropes and Merrill, 1966; Ropes,

1967). Gliding, the result of a clam forcefully pushing

itself off of a substrate by a sudden straiglitening of

the bent, extended foot, is termed "leaping" by

Ansell (1969). The clam glides with the posterior part

of its shell directed like a keel (Fig. 1). Muscular acti-

vities of the foot or adductor muscles that miglit be

used for swimming movements have not been seen

during the gliding act. After a leap, a surf clam glides

for only a few seconds and then falls to the bottom

because of negative buoyancy.

Our observations of gliding surf clams were inci-

dental to other experiments. The action was very in-

frequent. For example, more than 30,000 small

clams, hand dug between 1964 and 1967 from the

beach at Wallops Island, Virginia, immediately rebur-

rowed if left on the sand surface. This action hap-

pened whether the clams were in or out of the water.

Only two small ones, 25 to 30 mm. in length, were

seen gliding in the water. Upon landing on the sand

surface, they began to reborrow.

Stimuli for the gliding or leaping response in Mac-

tra corallina, a close relative of S. solidissima in the

family Mactridae, have been reported by Ansell

(1969) to be: (1) close proximity or contact with a

predator, (2) failure of the foot to penetrate the sub-

stratum, and (3) exposure to air. He considered the

latter two stimuli to be of minimal ecological signi-

ficance and explained that, to increase the chance of

contact and obtain a reaction from predators, the

clams were confined in small aquaria. No mention

FIG. 1. A small (about 25 mm. long) surf clam leap-

ing.

20

THE NAUTILUS

January 1973

Vol. 87(1)

was made of the possibility of a clam closing its shells

or burrowing deeper in the substratum to avoid

predator contact. This latter activity was described by

Pratt and Campbell ( 1956) as a behavior of northern

quahogs IMercenaria mcrccnaria) confined in boxes

with predaceous starfish.

Surf clams have adaptations for living in a burrow

and divers have always found them there. Burial may

be well below the sand surface, because the siphons

can be extended nearly equal to the length of the

shell. Numerous papillae fringing the siphonal open-

ings (Morse. 1919) may be adaptations to prevent

sand from being drawn into the body cavity by the

respiratory and food-gathering water currents. The

clean shells of most clams taken by dredges or thrown

onto beaches by storms indicate that they had been

covered completely. A report by Ropes and Merrill

(1966) on SCUBA observation of a bed of surf clams

near Cape Cod. Massachusetts, stated that the clams

were completely buried in the bottom. Visible siphon

holes were the only evidence that surf clams were

present in the bottom. Another SCUBA diver (Bul-

loch. 1965) described a surf clam bed about 4 miles

offshore from Point Pleasant, New Jersey. He saw

surf clams with the posterior portion of their shells

protruding from the sandy bottom. All seemed to be

oriented in the direction of a strong current flow

which had apparently removed sand from around the

shells. In spite of this, no clams were seen on the

surface or moving about on the surface. On rare occa-

sions, we have seen quantities of commercially cauglit

clams with algae growing on a portion of the shell

nearest the siphon. The localized growth of algae im-

plies that these clams have been embedded for some

time in the bottom with the posterior end slightly

above the surface. It is important to note that none

of the divers reported surf clams out of the bottom.

The diver who reported on the surf clams near Cape

Cod dislodged some and they promptly reburied

themselves (Ropes and Merrill, 1966). A clam of com-

mercial si/e ( 1 25 mm. long) took about 2 minutes to

burrow into the bottom. Subsequent to these obser-

vations, the diver has collected surf clams for us at

monthly intervals during 2 years. Althougli we have

asked him to observe and photograph movements, he

has been unable to obtain any evidence of crawling,

gliding, or leaping.

The movement of surf clams out of their burrows

has not been clearly demonstrated as a usual loco-

motor activity. Surf clams have been observed to ele-

FIG. 2. Surf clams raised our uj the cxpascJ inur-

tidal substratum at Wallops Island. Virginia.

vate themselves out of the bottom substratum on ex-

posed beaches during periods of low tide (Ropes.

1967) (Fig. 2). Exposure of the drained bar to solar

heat may have initiated the clam movement out of

the bottom. Apparently the effort to dig deeper in

the more compact, drier substrate moved the clam

upward. However, our attempts to cause surf clams to

exhume in laboratory experiments, by draining water

from the bottom of the aquarium and exposing the

surface of the sand to radiated heat, were unsuccess-

ful.

Storms appear to be very effective in moving surf

clams from an area; large numbers of surf clams are

found washed upon the beach after severe winter

storms (Yancey and Welch, 1968). Bottom currents,

produced during storms, dislodge clams from the bot-

tom and many are unable to burrow quickly enough

to avoid being thrown further up on the beach. That

many do reburrow is evident, since the surf zone at

low tide is one of the best areas to collect small surf

clams.

Armstrong (1965) studied the burrowing limita-

tions of 10 species of clams. He found that the clams

showed no movement other than rotational or bur-

rowing, and that none were able to exhume or elevate

in their burrows under their own power. Clams

planted 4 to 7 inches below the tips of their extended

siphons died. Our experiments with the juvenile and

adult surf clams produced similar results.

We have conducted several other experiments to

determine burrowing movements of surf clams. Bur-

rowing ability was considered essential to survival of

clams marked and replanted for growth studies

Vol. 87(1)

THE NAUTILUS

21

(Rt)pes and Mcnill. 1^70). Clams. 50 to 75 mm. long.

placed in an aquarium, dug into the bottom soon

after initiating burrowing. Descriptions and photo-

graphs of this action were reported by Ropes and

Merrill (1966). However, small clams averaging 25

mra also showed some gliding movements, suggesting

that small surf clams may be more active than larger

ones. Gliding by these small clams was apparently

caused by the disturbance of other clams burrowing

nearby. The clams burrowed upon landing on the

sand surface. Burrowing was the only consistent re-

sponse obser\'ed.

Clams we marked and planted off Wildwood, New

Jersey, were recovered in the same area one year

later. Several hundred clams were recovered up to

two years after being marked and planted in shallow

water near Chincoteague Inlet, Virginia (Ropes and

Merrill, 1970). Recovery of clams was always within

the release area, indicating a lack of movement.

Heavy, hydraulic jet dredges used by commercial fish-

ermen are not sufficiently disturbing to cause clams

actively to "flee" an area, because the fishermen re-

peatedly dredge througli a dense patch or "hot spot"

for periods of several days or sometimes months and

still take clams in commercial numbers. The staff of

the BCF Exploratory Fishing and Gear Research

Base, Gloucester, Massachusetts, have obsei-ved surf

clam beds and the action of a commercial-type hy-

draulic jet dredge ' by underwater television and

cinematography. At no time were surf clams seen on

the surface of the sand bottom, nor were any seen

gliding through the water in escape movements' . The

many field and laboratory observations demonstrate

that surf clams rarely move other than to reburrow

when disturbed or removed from their burrows.

' Personal communication by Phillip S. Parker,

Fishery Biologist, formerly with the BCF Explora-

tory Fishing and Gear Research Base, Gloucester,

Massachusetts.

We wish to stress that, from our observations, the

movement of surf clams can be considered to be in-

frequent and exceptional. All of the movements we

have observed have occurred after the clams were dis-

turbed by removal from the substratum, and the

clams invariably reburrowed. Voluntary movements

of surf clams out of the bottom have never been re-

ported nor have we been able to induce this activity

in laboratory experiments. The abihty of most pele-

cypod mollusks to move througli the water is so rare

that notes are published reporting such unusual

phenomena (Baker and Merrill, 1965). Surf clams,

especially the larger adults of commercial size, are,

for all intents and purposes, sedentary bottom dwel-

lers.

The above observations were used to prepare a

justification for adding the surf clam to the List of

Creatures of the Continental Shelf An amended

regulation to the law included surf clams and was

published in the Federal Register on June 23, 1971.

LITERATURE CITED

Ansell. Alan D. 1969. Leaping movements in the

Bivalvia. Proc. Malacol. Soc. Lond. 38: 387-399.

Armstrong. Lee R. 1965. Burrowing limitations in

Pelecypoda. The Veliger 7 (3): 195-200.

Baker, Emmett B. and Arthur S. Merrill. 1965. An

observation of Laevicardium mortoni actually

swimming. The NautUus 78 (3): 104.

Belding, David L. 1910. The growth and habits of the

sea clam (Mactra solidissima). Rep. Comm. Fish.

Game Mass. 1909, Publ. Doc. 25: 26-41.

Bulloch, David K. 1965. Field observations - 1964.

Underwater Natur. 3 (1): 27-28.

Edwards. R. L. and K. 0. Emery. 1968. The view

from a storied sub - the Alvin off Norfolk, Va.

Commer. Fish. Rev. 30 (8-9): 48-55.

Morse, Edward S. 1919. Observations on living

lamellibranchs of New England. Proc. Boston Soc.

Natur. Hist. 35: 139-196.

Pratt, David M. and Donald A. Campbell. 1956.

Environmental factors affecting growth in Venus

mercenaria. Limnol. Oceanogr. 1: 2-17.

Ropes, John W. 1967. The locomotion and behavior

of surf clams, Spisula solidissima. Proc. Nat. Shell-

fish. Ass. 57: 4. (Abstract).

Ropes, John W. and Arthur S. Merrill. 1966. The bur-

rowing activities of the surf clam. Underwater

Natur. 3(4): 11-17.

Ropes. John W. and Arthur S. Merrill. 1970. Marking

surf dams. Proc. Nat. Shellfish. Ass. 60: 99-106.

Yancey, Robert M. and Walter R. Welch. 1968. The

Atlantic coast surf clam - with a partial biblio-

graphy. U. S. Fish WUdl. Serv. Circ. 288, 14 pp.

THE NAUTILUS

Januarv I 973

Vol. 87(1)

LEO GEORGE HERTLEIN (1898 1972). AN

OBITUARY AM) FINAL HII5LI0GR APHY

William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

Central Park West at 79th. N. Y.. N. Y. 10024

Dr. Leo George Hertlein. Curator Emeritus of

Invertebrate Paleontology, California Aeadeiny of

Seiences. San Francisco, died on January 15, 1972

after a brief illness. His deatli came as a shock to his

many friends, for he appeared to be enjoying robust

health and looked much younger than his years. He

was actively working on several research projects at

the Academy until the time of his illness. He liad

noted on his 1971 Christmas card that he was enjoy-

ing his retirement and was now able to devote most

of his time tt) the completion of a long-term study on

the geology and paleontology of the marine Pliocene

of San Diego, California. The third part of this

treatise, a monograph on the pelecypods of the San

Diego formation, was in press at the time of his death

and has now been published (see review. The

Nautilus. 86 (2-4), This major contribution, together

with more than 150 scientific publications, testifies

to his ability and devotion as a scientist who held the

universal esteem of his colleagues.

On the occasion of his retirement, the October,

1970. number of The Nautilus was subtitled the "Leo

George Hertlein Honor Issue" by his friends who con-

tributed a biographic profile, a list of taxa proposed

in his honor, a bibliography of his works, a list of

names proposed b\ him. and original papers on

MoIIusca dedicated to him.

A gentleman and a scholar. Leo was a quiet,

friendly man who was always willing to assist others,

be they amateurs, students, or professional col-

leagues. He will be remembered by his many friends

as a kind man who combined characters of high moral

and ethical strength with a gentleness and warmth of

aftection thai wa^ uiuque. Leo Hertlein was trul> a

Prince of a Man,

I have appended l)i. ilerlleni's bibliography and a

list ol new laxa pro|iosed by him, fi)r the period of

1970 to 1972.

Bibliography of Leo G. Hertlein

for the period of 1970-1972

Papers preceded b\ an asterisk (*) contain new

taxa.

1970a, *A new species of fossil Kclktia (Mollusca:

Gastropoda) Irom the Loniita Marl, late Cenozoic

of San Pedro, California, Contributions in Science.

Los Angeles Co. Mus.. no. 190, p. 1-8, fig, 1-3,

1970b. Review of Carrington, A, J. and B. F.

Kensley. 1969. Pleistocene Mollusks from the

Namaqualand Coast. Veliger, 12: 381.

1970c. Review of Figueiras. Alfredo and Omar E.

Sicardi, 1968, Catalogo de los Moluscos Marinhos

del Uruguay. Veliger, 12: 381-382.

1970d. Review of Work, Robert C, 1969. System-

atics. Ecology, and Distribution of the Mollusks of

Los Roques, Venezuela. Veliger, 12: 382.

1970e. Review of Barnard. K. H.. 1969. Contri-

butions to the knowledge of South African Mol-

lusca. Part IV: Veliger, 12: 382.

1970f, Review of Ludbrook, Nellie H., 1969, The

genus Miltha in the Australian Cainozoic: Veliger,

13: 114.

1970g. Review of Kensley, Brian and Mary-Louise

Penrith. 1970, New records of Mytilidae from the

northern South West African coast: Veliger, 13:

205.

1970h, (v\'ith Victor A. Zullo) Catalogue of specimens

in the type collection of the Department of Geo-

logy, California Academy of Sciences. Cephalo-

pi)da: Occ. Papers Calil'ornia Acad, Sci.. no. 82,

130 pp.

1 97 la. Review of Kosuge. Sadao. 19(i9, Fossil Mol-

lusks of Oaliu. Hawaiian Islands: Veliger. 13: 301.

1971b. Review of Fischer-Piette. L,, M. Kempf, and

A. M. Testud, 1969, Nouvelles donnees sur les

Veneridae (mollusques, lamellibranches) du Bresil:

Veliger, 13: 302,

1971c. Review of Darragh. Thomas A., l970,Cata-

logue of Australian Tertiary Mollusca (except

chitons): Veliger, 13: 302.

197 Id. The researcher and the library: Spec. Libr.

Assoc, San Francisco Bay Reg, Chap, Bull.. 41

(4): 7-8,

|97 1e. Review of Osorio, Cecelia and Nibaldo

Bahamonde, 1970. Lista preliminar de lameli-

branquios ile Chile; Veliger, 13: 373.

1 97 If. Review of Ageitos de Castellanos, Zulma J.,

1970, Catalogo de los moluscos marinos

Bonaerenses: Veliger, 13: 374.

Vol. 87 ( 1 )

THE NAUTILUS

23

1972a. *Description of a New Species oi Chlamys

(Mollusca: Pelecypoda) From the Galapagos Is-

lands; Proc. California Acad. Sci., ser. 4, 39 (1):

1-6, 5 figs.

1972b. (with Victor A. ZuUo): Catalog of Specimens

in the Type Collection of the Department of Geo-

logy, California Academy of Sciences. Echino-

dermata: Occ. Papers California Acad. Sci., no. 91,

26 pp.

1972c. *Pliocene fossils from Baltra (South Seymour

Island), Galapagos Islands: Proc. California Acad.

Sci., ser. 4, 39 (3): 25-46, 39 figs.

1972d. *(and Grant, U. S., IV). The geology and

paleontology of the marine Pliocene of San Diego,

California, Pt. 2b, Paleontology (Pelecypoda): San

Diego Soc. Nat. Hist. Mem., 2: 135-411, text figs.

7-13, pis. 25-57.

Names Proposed by Leo G. Hertlein

From 1970 To 1972

The format follows that used in the previous list-

ing for the period of 1925 to 1970 (Nautilus 84 (2):

pp. 52-69). All the new taxa in the present list are

mollusks and are referable to the class Bivalvia

(Pelecypoda), except Kelletia kanakoffi, which is a

gastropod.

abbotti Hertlein & Grant, Chlamys abieds 1972d: p.

196, pi. 34, fig. 5;pl. 36, figs. 1,6.

allisoni Hertlein & Grant, Chione (Chione) 1972d: p.

273, pL 51, figs. 11-16.

ashleyi Hertlein & Grant, Semele 1972d: p. 299, pi.

48, figs. 3, 4, 6, 9, 10.

axinola Hertlein & Grant, 1972d: p. 159. Type spe-

cies by original designation: Axinaea (? septentri-

onalis, war.) siibobsoleta Carpenter, 1864.

balboana Hertlein & Grant, Nucula (Ennucula)

1972d: p. 145, pi. 27, figs. 1-4.

baltrana Hertlein, Diplodonta subquadrata 1972c: p.

33, figs. 8, 11.

diegoana Hertlein & Grant, Aligena 1972d: p. 235, pi.

44. figs. l,6;pL45, figs. 6, 7, 10, 11, 13.

diegoana Hertlein & Grant, Dosina (Dosina) pon-

derosa 1972d: p. 265, pi. 47, figs. 2, 3, 6, 8, 10;

pi. 49, fig. 8, pL 51, fig. 14.

ellisi Hertlein & Grant, Chlamys (Chlamys) hastata

1972d: p. 190, pi. 31, figs. 2, 3; pi. 34, fig. 6.

frankiana Hertlein & Grant, Borina (Temblorinaj

1972d: p. 238, text fig. 11.

incantata Hertlein, Chlamys (Chlamys) 1972a: p. 2,

figs. 1-5.

imsella Hertlein & Grant 1972d: p. 279. Type species

by original designation: Venus lamellifera Conrad,

1837.

kanakoffi Hertlein & Grant, Chione (Securella)

1972d: p. 274, pL 49, figs. 1, 3, 4-6; pL 51. figs.

4-7, 10.

kanakoffi Hertlein, Kelletia 1970a: p. 1, figs. 1-3.

kanakoffi Hertlein & Grant, Thracia 1972d: p. 338,

pi. 42, figs. 11, 13-15.

orcutti Hertlein & Grant, Lima (Limaria) 1972d: p.

215, pi. 35, fig. 11; pL 36, figs. 2-5; pi. 57, fig. 10.

stephensae Hertlein & Grant, Psephidia 1972d: p.

280, pL44, figs. 21,22, 24, 26.

stembergi Hertlein & Grant, Mytilus (Crenomytilus)

coalingensis 1972d: p. 163, pL 41, figs. 10, 14.

BOOK REVIEW

ART FROM SHELLS. By Stuart and Leni Goodman.

210 pp., 393 photographs, 21 in color. Crown

Publ. Inc., 419 Park Ave. South, N. Y., N. Y.

10016. Paperback, S3.95; cloth, hardback, $7.95.

This is probably the most extensive and most prac-

tical book on the subject of using shells in jewelry,

shell pictures, collages, figurines, and shell sculptures.

It is profusely illustrated with simple pictorial steps

on how to clean, mount and use shells in various art

forms. There are short sections on collecting and

cleaning shells, as well as a current list of supphers.

R. Tucker Abbott

Delaware Museum of Natural History

24

THE NAUTILUS

January 1973

Vol. 87(1)

THE PROBLEM OF THE LAND SNAIL GENUS

RABDOTUS IN TEXAS ARCHEOLOGICAL SITES

John W. Clark, Jr.

Balcones Research Center

Rt. 4, Box 189, Austin, Texas 78756

Vast numbers of land snaUs, principally Rabdotus

dealbatus (Say) in central Texas and R. schiedeanus

(Pfr.) in south and southwest Texas (Pratt 1969:

47-48), are found in prehistoric archeological sites.

These snails often occur in lenses of thousands of

adult shells associated with cultural debris in burned

rock middens in central Texas and in rockshelters in

southwest Texas.

The area in central Texas in which this pheno-

menon occurs lies within the Edwards Plateau region,

especially to the east and south edges of the Balcones

Escarpment where limestone forms the bedrock and

numerous streams dissect the plateau. This area falls

within the Balconian Biotic Province described by

Blair (1950; 12-13).

In southwest Texas, the area of snail-archeological

association is quite restricted. It is centered on the

Rio Grande River in the vicinity of the mouths of the

Pecos and Devil's Rivers. This area falls on the bound-

aries of Blair's Chihuahuan and Balconian Biotic Pro-

vinces (Blair 1950: 105-108). It is characterized by a

limestone plateau dissected by deep canyons. Both

areas are within the Great Plains Province which is

characterized by decreasing annual rainfall from east

to west (Carr 1967: 3-4) so that the central Texas

area receives between 22 and 30 inches of rain per

year while the southwest area receives between 13

and 16 inches per year. Rabdotus shells are also

found in some archeological sites in south Texas as-

sociated with Blair's (1950: 102-105) Tamaulipan

Biotic Province.

The vast numbers of these snails in central Texas

burned rock middens was noted as early as 1932

(Pearce) and 1933 in southwest Texas (Martin).

Numerous hypotheses have been formulated since

then attempting to explain this phenomenon. The

four principal hypotheses are:

(1) These snails were intentionally collected as a

source of protein in the diet (Martin 1933 and Suhm

1957),

(2) They represent a natural population of snails

attracted by decomposing midden refuse,

(3) They were carried to the sites inadvertently on

vegetation collected by the Indians, and

(4) They were collected for their shells which were

used as adornment (Martin 1933). Rabdotus shells

appear to have accumulated or been collected in the

two regions over quite a long period of time, from

about 8,000 B.C. at Arenosa Shelter at the mouth of

the Pecos River to about 1.000 B.C. in central Texas.

Unfortunately, much of the speculation has been

based on little empirical evidence. There is no good

ethnographic account of Indians in either region eat-

ing snails and the evidence for the use of shells for

decoration is scant. No empirical evidence for the

remaining hypotheses has been obtained through

archeological techniques. One of the principal prob-

lems concerning the presence of Rabdotus is

whether they were principally collected as a food

source. Providing they are preserved, the presence of

radulae of these snails in coprolites (dessicated human

feces) could be a major factor in assigning hypothesis

number one as the principal cause of the snaO lenses

in the sites. Coprolites are present in a collection

from a rockshelter in southwest Texas. Analysis of

coprolites from occupation lenses in which numerous

snail shells were found should provide some im-

portant data on the snail collection at this site and,

by extension, at other sites.

Tliis short paper is principally a plea for informa-

tion concerning the validity of the hypotheses pre-

sented herein and an appeal for co-operation of

malacologists in the analysis of coproUte samples for

snail radulae.

LITERATURE CITED

Blair, Frank W. 1950. The Biotic Provinces of Texas.

The Texas Journal of Science. 2(1).

Carr, John T., Jr. 1967. The Climate and Physio-

graphy of Texas. Texas Water Development Board,

Report, 53.

Martin, George C. 1933. Archaeological Exploration

of the ShuirJa Caves. Tlie Witte Memorial Muse-

um, Bulletin 3.

Pearce, J. E. 1932. The Present Status of Texas

Archeology. Bulletin of the Texas Archeological

and Paleontological Society 4.

Pratt, William D. 1969. Jiabdotus in Texas, Prelimi-

nary Revision of the Land Snail Genus. The

American Malacological Union, Inc., Annual

Reports for 1969: 47-48.

Suhm, Dee Ann. 1957. Excavations at the Smith

Rockshelter, Travis County, Texas. The Texas

Journal of Science, 9(1).

Vol. 87(1)

THE NAUTILUS

25

NOTES ON VERONICELLIDAE AND ATHORACOPHORIDAE

IN FIELD MUSEUM OF NATURAL HISTORY, CHICAGO.

Lothar Forcart

Naturhistorisches Museum, Basel

Switzerland

Dr. A. Solem entrusted me with the identification

of slugs in the collections of Field Museum of Natural

History, Chicago. These materials expand our know-

ledge on the distribution and taxonomy of several

veronicellid and athoracophorid species. The follow-

ing abbreviations indicate the repository of cited

materials:

FMNH - Field Museum of Natural History, Chicago.

MG - Zoologisches Institut und Museum, Gottingen.

NMB - Naturhistorisches Museum, Basel.

ZMB - Zoologisches Museum, Berlin.

VERONICELLIDAE

Laevicaulis alte (Ferussac, 1821)

Material examined: New Caledonia, E. coast 5 km.

N. of Theim (2 spec, FMNH 144278, coU. Jan. 1962

by A. & B. Solem); W. coast 8 km. W. Bourail (6

spec, FMNH 159224, 159237, coU. Oct. 1967 by L.

Price).

Remarks: Grimpe & Hoffman (1925, p. 366) pre-

viously recorded L. alte from New Caledonia. A sum-

mary of its complex synonymy is given in Forcart

(1969, pp. 148-149).

Veronicella (Tenacipesj tenax Baker, 1931

Veronicella (Tenacipesj tenax Baker, 1931,

Nautilus, 44(3): 131, pL 8, figs. 3-5. (Ensenada

de San Vicente, Pinar del Rio, Cuba).

Material examined: West Indies: Puerto Rico, foot

of stone cliff (2 spec, FMNH 157272, coU. Feb.

1968 by M. Moran). Dominica (5 spec, FMNH

157135, coll. Feb. 1968).

Remarks: All specimens are juvenile, the largest

being only 46.5 mm. long, as compared to the 96-105

mm. holotype and paratypes described by Baker.

They agree with the original description in the notum

having small papillae, in the sole being considerably

shorter than the notum and completely surrounded

by the hyponota, the posterior end being rounded

and crossed by numerous fine, transverse grooves.

The anus is as in the subgenus Leidyula, only partly

covered by the foot. The vagina is without accessory

bursa. The penis and the stimulatory organ

(dart-gland in Baker 1931) are as figured by Baker

(1931, pi. 8, figs. 4, 5) for the holotype.

All these specimens differ from the types in lack-

ing whitish spots on the notum. The three specimens

from Dominica have dark lateral bands on the notum.

?Vaginulus (Latipesj occidentalis (GuUding, 1825)

Onchidium occidentale Guilding, 1825, Trans.

Linn. Soc, London, 14: 323, pi. 9, figs. 9-10.

(St. Vincent, Windward Islands).

Vaginulus (Latipesj occidentalis (Guilding), Baker

1925, Proc Acad. Nat. Sci. PhUadelpliia, 77:

174-177, pi. 5, figs. 18-20.'

Material examined: Almirante, Bocas del Toro,

Panama. (5 spec, FMNH 86536, coU. Mar. 1959 by

H. S. Dybas).

Remarks: The examined specimens agree with

those Baker (1925) described from Panama. No

type-specimens of V. occidentalis exist, and 1 have

not been able to examine specimens from St.

Vincent. Onchidium occidentale is a nomen dubium

and it is doubtful whether the specimens from

Panama are the same species. Proposal of a new name

for the Panama specimens should wait until the iden-

tity of the St. Vincent specimens has been deter-

mined.

Vaginulus (Angustipesj buergeri (Simroth, 1914)

Vaginula burgeri Simroth, 1914, Mem. neu-

chateloise Sci. nat., 5: 328, pi. 14, figs.

124-126. - Santo Domingo. Lectotype and 2

lectoparatypes MG (see Thome, 1970: 73-76,

figs. 1-7).

Cylindrocaulis occidentalis Hoffmann 1925, Jena.

Z. Naturw., 61 ('/-): 144-147 pardm, pi. 5, fig.

45, 2, d, F.

Material examined: Salilia, Dominica. (3 spec,

FMNH 157147).

Remarks: The detailed redescription of V. buergeri

by Thome (1970, pp. 73-76, figs. 1-7) makes it evi-

dent that Baker (1925, 180-182, pi. 6, figs. 28-30)

described another species under this name.

The bursa of the receptaculum seminis is elongate,

sessile (Thome, 1970, p. 88, fig. 5), and has no dif-

ferenfiated stalk as described by Baker (1925, p. 181,

pi. 6, fig. 28). The stimulatory organ has only 9

tubules, of wluch some are bifurcated (Thome, 1970,

p. 175), and not 26, as menfioned by Baker (1925, p.

26

THE NAUTILUS

January 1973

Vol. 87(1)

181). The verge is short, lanceolate, as figured by

Hoffman (1925, pi. 5, fig. 45, d, 2, F) and Thome

(1970, p. 87, fig. 7), and has no spiral ridges, as des-

cribed by Baker (1925, p. 1 81, pi. 6, figs. 29, 30).

Vaginulus (Sarasinulaj plebeius Fischer, 1 868

Material examined: Saint Martin, Leeward islands

(1 spec, FMNH 173062, coll. Oct. 1947 by Oden

Meeker); Saba Island (1 spec, FMNH 157155, coll.

Jan. 1968).

New Caledonia, 19 km north of Boulouparis (2

spec, FMNH 159240, coll. Oct. 1967 by L. Price)

and 8 km west of Bourail, on pierced rock (3 spec,

FMNH 159224, coll. Oct. 1967 by L. Price).

Marianas, Saipan (1 spec, FMNH 173065, coll.

Feb. 1945); Guam, Ritidian Point (1 spec, FMNH

173063, coU Aug. 1945 by H. S. Dybas).

Philippine Islands, Leyte, San Jose (1 spec, FMNH

173064, coll. Dec. 1944 by E. Ray).

Moluccas, Ambon, Wasi (25 spec, FMNH 172984,

172986, 172991, 172992, 172993, coll. 1964, Sept.

1965 by A. M. Wegner).

Remarks: Except for New Caledonia, V. plebeius

was not reported previously from these islands. It is

probably of West Indian origin, and has been acci-

dently introduced to Pacific and Mascarene Islands

(Baker, 1931, p. 135). A full synonymy is given by

Forcart(1969: 150).

Sempenda sclmeideri (Simroth, 1895)

Vagimda sclmeideri Simroth, 1895, SB. Naturf.

Ges. Leipzig, 19/21: 7. - Tandjong, Kuba, East

Sumatra.

Material examined: Tebing Tingii, Pedis, Malaya ( 1

juv. spec, FMNH 156725, colL Feb. 1966 by A.

Berry) under moist leaves at foot of hill.

Sempenda celebensis Hoffmann, 1941

Vaginula melotomus, Bollinger 1918, Rev. Suisse

Zool., 26 (9): 319, not Vagimda melotomus?.

Sarasin & F. Sarasin, 1899.

Semperula idae, Grimpe & Hoffmann 1925, Z.

wiss. Zool., 124 (1): 33-36 partim, fig. 14 (p.

14); Hoffman 1925 Jena Z.Naturw., 61 ('/.):

186-189, 255-256 partim, pL 6, fig. 45, k, 2.

Not Vaginula idae Semper, 1 885.

Vaginula (Semperula) idae, Forcart 1935, Verh.

Naturf. Ges. Basel, 46, p. 54, not Vaginula idae

Semper, 1885.

Semperula wallacei var. celebensis Hoffmann,

1941, Zool. Anz., 136(11/12); 234. New name

for Vaginula melotomus Bollinger, 1918, not P.

Sarasin & F. Sarasin. 1899. Type locality (re-

stricted): South Celebes, Lamontjong. Lecto-

type NMB 2419-b, coll. P. & F. Sarasin

1902/03. 5 lectoparatypes, NMB 2419-a from

Central Celebes, Gimpu, coll. P. & F. Sarasin

1902/03.

Material examined: Western Samoa, Upolu,

Solaua-Sauniato road junction, coconut-cocoa planta-

tion (1 spec, FMNH 152677, coll. Nov. 1965, by A.

Solem & L. Price). Moluccas, Ambon, Wasi (5 spec,

FMNH 172987, 172988, 172991. coll. 1964 by A.M.

Wegner).

On the Veronicellidae from Ambon

Van Benthcm Jutting (1953. p. 300) listed from

the Moluccan Island of Ambon Laevicaulis alte

(Ferussac), Filicaulis bleekeri (Keferstein), and

Sempenda maculata (Templeton). These citations are

erroneous. Martens (1867. p. 176) described the first

Veronicellid known from Ambon as Vaginulus has-

selti. mentioning it (p. 68) from Bangkok, Thailand,

and (pp. 176-177) from Sumatra, Borneo, Ambon,

and Banda. Hoffmann (1925: 136-137), revising

syntypes of V. liasselti (ZMB). found those from

Bangkok and Sumatra identical with Filicaulis

bleekeri (Keferstein. 1865), but those from Ambon

and Banda with Sempenda maculata (Templeton,

1858).

Vagimda bocagei Simroth. 1893 was described as

from Ternate, Ambon, and from Banda. Grimpe &

Hoffmann (1925: 29-30), revising 14 specimens in

the ZMB labeled V. bocagei. probably the syntypes of

V. bocagei, stated that 9 specimens from Ambon are

identical with Sempenda maculata, and 5 specimens

from Banda with Laevicaulis alte (Fer.). Laevicaulis

alte is consequently not known from Ambon, and the

citafions of Ambon as locality of I. alte in Grimpe &

Hoffmann (1925, p. 28) and Hoffmann (1925: 227)

are erroneous.

Tlie revisions of the specimens from Ambon pub-

lished as Vaginulus liasselti and Vaginula bocagei re-

sulted that these determinations arc misinterpreta-

fions for Sempenda maculata, hitherto the sole spe-

cies known from Ambon. Semperula celebensis

Hoffmann and Vaginulus (Sarasinulaj plebeius

Fischer are added to the fauna of Ambon by the new

material.

Vol. 87(1)

THE NAUTILUS

27

ATHORACOPHORIDAE

Aneita (Aiicita) simrothi Grimpe & Hoffmann, 1925

Aiicira simrothi Grimpe & Hoffmann, 1925, in

Sarasin & Roux, Nova Caledonia, 3 (3): 421,

figs. 15e, 16g, 17d, 18e, 20, pi. 5, figs. 5 a, b,

14 a. b. Type locality: New Caledonia, Mt.

Ignambi near Oubatche, 700-1300 m alt.

Syntypes NMB 3448 a, b, coll. April & May,

1911 by F. Sarasin & J. Roux.

Material examined: 6.4 km East of Ouegou, North-

east New Caledonia, at ± 130 m alt. (6 spec, FMNH

159245, coll. Oct. 1967 by L. Price). The biggest

specimen has a length of only 53.8 mm. as compared

with the 85 mm. length of the largest syntype.

LITERATURE CITED

Baker, H. Burrington. 1925. North American Veroni-

ceilidae. Proc. Acad. Nat. Sci. Philadelphia, 77:

157-184, pis. 3-6.

Baker, H. Burrington. 1931. Notes on West Indian

Veronicellidae. Nautilus: 44(4): 131-137, pl. 8.

Benthem Jutting, W. S. S. van. 1953. Annotedlist of

the non-marine Mollusca of the Moluccan Islands

Ambon, Haruku, Saparua and Nusu Laut. Treubia,

22(2): 275-318.

Bollinger, G. 1918. Land-Mollusken von Celebes. Rev.

Suisse ZooL, 26 (9): 309-340, pl. 1 1.

Forcart, L. 1935. Revision der Vaginuliden aus

Celebes in der Sammlung des Easier Natur-

historischen Museums. Verb. Naturf. Ges. Basel,

46: 50-55.

Forcart, L. 1969. Veronicellid land slugs from the

New Hebrides, with description of Semperula

solemi new species. Fieldiana Zool., 51 (12):

147-156.

Grimpe, G. & Hoffmann, H. 1925; Versuch einer Re-

vision der indischen, indo- und polynesischen

Vaginuliden. Z. wiss. ZooL, 124(1): 1-50.

Grimpe, G. & Hoffmann, H. 1925a. Die Nackt-

schnecken von Neu-Caledonien, den

Loyalty-lnseln und den Neuen Hebriden. Sarasin,

F. & Roux, J., Nova Caledonia, A. Zool., 3 (3):

339-476, pis. 5-6.

Hoffmann, H. 1925. Die Vaginuliden. Jena. Z.

Naturw., 61(1/2): 1-374, pis. 1-11.

Hoffmann, H. 1941. Ueber einige Vaginuliden auf

Grund bisher fur verschollen gehaltener

Typstucke. Zool. Anz., 136(1/2): 229-242.

Martens, Ed. 1867. Die Landschnecken. Die Preus-

sische Expedition nach Ost-Asien, Zool., 2: 1-Xll,

1-447, pis. 1-22.

Thome, J. W. 1970. Redescricao dos tipos de Veroni-

cellidae neotropicais: III. Especies depostadas no

"11. Zoologisches Institut und Museum der Univer-

sitiit" de Gbttingen, Alemania. Iheringia Zool., 38:

73-88.

OBITUARY

Gilbert Ranson (1899-1972)

Gilbert Ranson, distinguished French zoologist

and oyster expert died in Paris August 19, 1972, at

the age of 72. He received the equivalent of B. S. in

natural sciences at the Sorbonne in 1923, and was a

student of Louis Joubin. He received his Ph. D. in

1927. his thesis being on the physiology of aquatic

animals. He published well over 200 articles on dia-

toms, medusae and especially the taxonomy of

oysters. In 1951, he authored the book, Les

Huitres—biologie--culture. Dr. Ranson became

sub-director of the Malacological Laboratory of the

National Natural History Museum in Paris in 1938.

He was born in France on September 4, 1899.

St. Croix Shell Booklet

A revised edition of "A Supplementary Listing of

New Shells (Illustrated) to be added to the Check List

of the Marine Shells of St. Croix" by G. W.

Nowell-Usticke was published on Dec. 20, 1971. It is

privately printed and distributed by the author, 1

North St., Christiansted, St. Croix, Virgin Islands

00820, and is available for $3.69 post fiee, or S4.00

by airmail. It has 32 pp., an index, 6 pis., many new

species and subspecies, many of which are synonyms

and/or homonyms.

THE NAUTILUS

January 1973

Vol. 87(1)

SHORT PAPERS

NEW NAME FOR A TEXAN HYDROBIA

J. P. E. Morrison

Division of Mollusks

National Museum of Natural History

Washington. D. C. 20560

CERION FROM AN ARCHEOLOGICAL SITE

IN NORTHERN OHIO

James L. Murphy

Sears Library

Case Western Reserve University

Cleveland, Ohio 44106

A single fragmentary specimen of the terrestial gas-

tropod Cerion incanum (Binney) has been found in

collections from a Late Woodland or Late Prehistoric

village site in Lorain County, Ohio. The Eiden Site

hes on a bluff top in the vUlage of Sheffield, near the

confluence of French Creek and the Black River, two

mOes south of Lake Erie. The collections were made

by A. A. Bungart over ten years ago and are now

being analyzed for the Lorain County Park Board by

Dr. D. H. McKenzie and Dr. J. E. Blank, Cleveland

State University, Dr. 0. C. Shane, Kent State Univer-

sity, and the author.

McKenzie and Blank estimate that the major occu-

pation of the site lasted for approximately 50 years,

sometime between 1300 and 1450 A.D. Minor Early

Woodland and Late Prehistoric components also

occur at the site, however. Nonetheless, the over-

whelming majority of the materials recovered from

the site appear to represent a Late Woodland com-

ponent, and it is very probable that the Cerion shell

found in the general midden represents the Late

Woodland component.

Other shell artifacts associated with the Late

Woodland component include marine shell disc beads,

conch columella beads, and Marginella beads. The

occurrence of such a variety of worked marine shell

in a Late Woodland component is unusual and in it-

self lends support to the contention that the Cerion

shell belongs to the Late Woodland occupation of the

Eiden Site and is a Late Woodland trade item from

the Florida Keys.

Dr. R. Tucker Abbott kindly identified tlie Eiden

shell for me. The rather prominent vertical riblets of

the two last whorls suggest the form saccharimeta,

which Pilsbry (1946: 164) has found to intergrade

with typical C. incanum

The species named Odostomia barret ti Morrison

(Proc. Biol. Soc. Wash. 78: 220, fig. 4, 1965) is pre-

occupied by Odostomia barreti L. Morlet (Journ. de

Conchyl. 33: 314, 1885), as James X. Corgan of

Austin Peay State University, Tennessee, has so kind-

ly pointed out.

On July 25, 1968, with the guidance and helpful

assistance of Mrs. Hollis Q. Boone of Houston, Texas,

a considerable number of these tiny snails were col-

lected from a narrow stretch of intertidal sand on the

west side of Galveston Bay. Examination of the strik-

ingly pink, living animals proved them to belong to

the genus Hydrobia, as reported {in litt.) by Dr.

Robert Robertson of the Academy of Natural

Sciences of Philadelphia.

I wish to rename this brackish water species

Hydrobia booneae in honor of Mrs. Boone, who first

discovered them in numbers in Texas waters. At

present Hydrobia booneae is known to be Uving in

waters of reduced salinity between the northeast

corner of Heron Bay, Mississippi (the southwest

corner of that state) and the west side of Galveston

Bay, Texas.

•

PHYSA ACUTA IN VIRGINIA

Dorothy E. Beetle

Peninsula Nature and Science Center

Newport News, Virginia 23601

Physa acuta Draparnaud, an introduced European

fresh water snail, has been found at several localities

in coastal Virginia. It is well-established at two locali-

ties in the City of Hampton: along the shores of Big

Bethel Reservoir and in a small pond receiving water

pumped out of Rice's Fossil Pit, a well-known marine

Miocene deposit. In the City of Newport News, /%^sa

acuta occurs in Lake Maury. It is in York County at

Wormley Pond in the Colonial National Historical

Park. Another locality is on the North Carolina bor-

der of the City of Virginia Beach, Rt. 165, in a small

pond behind the beach at Currituck Sound.

Pond bottoms consist of sandy silt, and the snaUs

were Ibund along the shore in dead leaves. Specimens

were kindly identified by Dr. Wilham J. Clench.

Vol. 87(1)

THE NAUTILUS

29

SPREAD 0¥ MELANOIDES TUBERCULATA

R. Tucker Abbott

Delaware Museum of Natural History

Greenville, Del. 19803

The freshwater Asian thiarid, Melanoides tuber-

culata (Miiller) is now spreading rapidly southward

from United States into Central America. Mrs. Dot

Myhre, of Stuart, Florida, has sent us specimens col-

lected in the vicinity of Vera Cruz, Mexico, where the

shells are sold by the bagful for making novelties. Dr.

Byron N. Chaniotis, of the Walter Reed Army Insti-

tute of Research, Canal Zone, collected live speci-

mens in Panama in 1971. The snaU, an intermediate

host in China for the bile fluke, Clonorchis sinensis,

arrived in Texas and Florida prior to 1963, and soon

thereafter in Puerto Rico. A distributional map show-

ing the range of the species from North Africa to the

East Indies prior to 1927 was published by Pilsbry

and Bequaert, 1927, BuU. Amer. Mus. Nat. Hist., vol.

53. p. 253.

FRESH-WATER MOLLUSKS FROM

COASTAL VIRGINIA

Dorothy E. Beetle

Peninsula Nature and Science Center

Newport News, Virginia 23601

The following moUusks were collected from small

fresh water creeks and ponds in the independent

Cities of Hampton, Newport News and Virginia Beach

and York County: Anodonta cataracta Say, Pisidium

casertanum Poll, Sphaerium occidentale Prime, S.

partumeium Say, Campeloma decisa Say, Littorid-

inops tenuipes Cooper, Amnicola limosa Say , Pomati-

opsis lapidaria Say, Physa acuta Draparnaud, P.

heterostropha Say, Fossaria humilis Say, F. obrussa

exigua Lea, Pseudosuccinea columella Say, Helisotna

anceps Menke, H. trivolvis Say, Planorbula armigera

Say, P. wheatleyi Lea, Gyraulus parvus Say, Menetus

brogniartianus Lea, M. dilatatus Gould, M. dilatatus

buchanensis Lea, Laevepex fuscus C. B. Adams. Mate-

rial is in the collection of the author.

MONOGRAPH REVIEW

THE GENUS SEMELE IN THE WESTERN

ATLANTIC (Semelidae; Bivalvia) by Kenneth J.

Boss. Johnsonia, Harvard University, Cambridge,

Mass., Vol. 5, no. 49, pp. 1-32. May 25, 1972.

The taxonomic treatment of the six known hying

species of Western Atlantic Semelidae is sufficiently

good to enable workers to identify their unknown

material. The illustrations, descriptions, remarks and

locality records are up to the usual standard of pre-

vious Johnsonia numbers.

Tliere are, however, a startling number of typo-

grapliical errors, spelling mistakes, examples of incor-

rect grammar, and mis-uses of words - at least 97 in

only 32 pages. Part of this can be excused because of

an improved format of the journal, and a change in

printer. Readers should note, however, that the cor-

rect spelling for the common semele is Semele bel-

lastriata (Conrad), not bellestriata, and that the

gender of Semele is feminine (lenticularis, not len-

ticulare, and solida, not solidum). The geological

terms, particularly those such as Group and Forma-

tion, seem not to be employed correctly in several

places. Some biological terms are mis-used, such as

allometric, when allopatric was meant. Scientific

names have been hyphenized, contrary to the

"Rules", such as media-americana on page 11. How-

ever, in the synonymy, they should have been re-

tained, instead of eliminated, as on page 9. The

author, and editor, of this monograph has the sym-

pathy of fellow editors and malacologjsts, but we

shall all look forward to a more painstaking effort in

future numbers. As Dr. Boss recently stated himself

in reviewing another bivalve paper (see Jour. Paleon-

tology, Vol. 45, no. 3, p. 558), "In the final analysis,

the editorial offices ... are responsible for publishing

a manuscript which was never critically reviewed in a

format which was never carefully proofed, and the

short-comings are now the affliction of science ~ to

the embarrassment of all."

R. Tucker Abbott

du Pont Chair of Malacology

Delaware Museum of Natural History

30

THE NAUTILUS

January 1973

Vol. 87(1)

A. M. U. Meeting

INDEX TO THE NAUTILUS

The Thirty-Eighth Annual Meeting of The Amer-

ican Malacologjcal Union will be held in Newark and

Greenville, Delaware, from June 25 through 28,

1973. Lodging and the main meetings will be held on

the campus of the University of Delaware. Early regis-

tration will be possible on Sunday, June 24. One day

of meetings will be held at the Delaware Museum of

Natural History, near Greenville, about 6 miles from

Newark. The current President is Dr. Dee Dundee,

Dept. Biology, Louisiana State Univ., New Orleans,

La. 70150. Further details will be published in the

AMU Newsletter and the April issue of The Nautilus.

Tlie extensive 279-page Author Index to The

Nautilus, vols. 1-75, compiled and published by

Aurele La Rocque in 1963 is still available (unbound)

for the extremely low price of $2.00 postpaid. A con-

tinuation, covering vols. 76-85, was recently pub-

lished in no. 49 of Sterkiana, 1972. Reprints of tliis

very useful addition are also avaUable for 25 cents.

Wlien ordering both of these indispensable indices, we

suggest an additional 50 cents be added to cover

postage. Write: Dr. Aurele La Rocque, 125 South

Oval Dr., Ohio State Univ., Columbus, Ohio 43210.

The first truly comprehensive

and authoritative survey of shells

MORE THAN 250 MAGNIFICENT ILLUSTRATIONS, 100 IN FULL COLOR

In this one colorful volume the entire

field of shells is covered. Expert

guidance takes the reader from

man's early use of shells to today's

fascinating pursuit of oceanic

treasures. Written by one of the

world's leading authorities, it

provides a wealth of information and

lore about shells (their biology;

their influence on art, medicine,

religion, and archeology; their

importance in commerce), a complete

guide for the collector, and much

more. The brilliant photographs, more

than 100 of them in full color, make

this important and useful book a

delight to the eye as well as

informative and engrossing reading

8%"x 11 'A" A Rutledge Book.

KINGDOM OF

THE SEASHELL

by R TUCKER ABBOTT.

du Pont Chair of Malacology.

Delaware Museum of Natural History

$14 95. now at your bool^store, or

CROWN PUBLISHERS

419 Park Ave South. New York, NY. 10016

INFORMATION FOR SUBSCRIBERS

The annual subscription rate for The Nautilus is

$7.00 for individuals and $12.00 for institutions (do-

mestic or foreign). Subscriptions may be commenced

in January when a new volume begins. Send check or

money order to "The Nautilus" to Mrs. Horace B.

Baker, Business Manager, 1 1 Chelten Road, Haver-

town, Pa. 19083.

Back issues from volume 72 to date are obtainable

from the Business Manager. Volumes 1 through 71 (if

available) may be obtained in reprint or original form

from Kraus Periodicals, Inc., 16 East 46th Street,

New York, N.Y. 10017.

Advertising rates may be obtained from the

Business Manager or Editor.

CONTRIBUTORS

Manuscripts: Authors are requested to follow the

recommendations of the Style Manual for Biological

Journals, which may be purchased from the American

Institute of Biological Sciences, 2000 "P" Street,

N.W. Washington, D.C. 20036. Manuscripts should be

typewritten and doublespaced; original and one copy

are required, to facilitate reviews. Tables, numbered

in arable, should be on separate pages, with the title

at the top. Legends to photographs should be typed

on separate sheets. Explanatory terms and symbols

within a drawing should be neatly printed, or they

may be pencilled in on a translucent overlay, so that

the printer may set them in 8 pt. type. There is a

charge of 50 cents per word for this extra service. All

authors or their institutions will be charged 50 cents

per line of tabular material and taxonomic keys. The

publishers reserve the right, seldom exercised, to

charge $32 per printed page.

An abstract should accompany each paper.

Reprints and covers are available at cost to

authors. When proof is returned to authors,

information about ordering reprints will be given.

They are obtained from the Economy Printing Co.,

Inc., R. D. 3, Box 169, Easton, Maryland 21601.

APRIL, 1973

THE

NAUTILUS

Vol. 87

No. 2

A quarterly

devoted to

malacology and

the interests of

conchologists

V

Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker.

Editors: R. Tucker Abbott and Charles B. Wurtz

EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of Mollusks

National Museum of Canada

Ottawa, Ontario, Canada KIA-0M8

Dr. William J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 1 0024

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

Dr. James H. McLean

Los Angeles County Museum of Natural History

900 Exposition Boulevard

Los Angeles, Cahfornia 90007

Dr. Arthur S. Merrill

Biological Laboratory

National Marine Fisheries Service

Oxford, Maryland 21654

Dr. Donald R. Moore

Division of Marine Geology

School of Marine and Atmospheric Science

10 Rickenbacker Causeway

Miami, Florida 33149

Dr. Joseph Rosewater

Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. Gilbert L. Voss

Division of Biology

School of Marine and Atmospheric Science

10 Rickenbacker Causeway

Miami, Florida 33149

EDITORS

Dr. R. Tucker Abbott

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807

Dr. Charles B. Wurtz

3220 Penn Street

PhUadelphia, Pennsylvania 19129

Mrs. Horace B. Baker

Business and Subscription Manager

1 1 Chelten Road

Havertown, Pennsylvania 1 9083

OFFICE OF PUBLICATION

Delaware Museum of Natural History

Kennett Pike, Route 52

Box 3937, GreenviUe, Delaware 19807

Second Class Postage paid at Wilmington, Delaware

Subscription Price: $7.00 (see inside baci< cover)

THE

NAUTILUS

Volume 87, number 2 -- April 1973

CONTENTS

Thomas R. Waller

The Habits and Habitats of Some Bermudian Marine MoUusks 31

Richard Edward Young

Evidence for Spawning by Gonatus sp. (Cephalopoda: Teuthoidea) in the High Arctic Ocean 53

S. L. H. Fuller and C. E. Powell, Jr.

Range Extensions of Corbicula maiiilensis (Philippi) in the Atlantic Drainage of the United States . .59

Margaret C. Teskey

Emarginula dentigera Heilprin, 1 889, a Littoral Mollusk 60

Book Reviews

(of) S. Peter Dance, 52; A. Myra Keen, 58

News; AMU Meeting in June 1973 59, 60

The first truly comprehensive

and authoritative survey of shells

MORE THAN 250 MAGNIFICENT ILLUSTRATIONS, 100 IN FULL COLOR

In this one colorful volume the entire

field of shells is covered. Expert

guidance takes the reader from

man's early use of shells to today's

fascinating pursuit of oceanic

treasures Written by one of the

world's leadmg authorities, it

provides a wealth of information and

lore about shells (their biology:

their influence on art. medicine,

religion, and archeology: their

importance in commerce), a complete

guide for the collector, and much

more The brilliant photographs, more

than 100 of them in full color, make

this important and useful book a

delight to the eye as well as

informative and engrossing reading

8%" X 11 'A", A Rutledge Book,

KINGDOM OF

THE SEASHELL

$14 95, now at your bookstore Of

ogy, CROWN PUBLiSHERS

al History 419 Park Ave Soutri, New York, N Y 10016

Ul

. »»»»»»\»»»»»w%%»»»»»»»»*»*»*v»»»»»»»»»»w»»*v»\»»»w»»»»*»»*^

ii^ne ibneu Lamnet

Successor to:

Richard E. Petit

and

John Q. Burch

OLIVE SHELLS

OF

THE WORLD

SPECIMEN SHELLS

NATURAL HISTORY

BOOKS

ROWLAND f ZEIGLER

HUMBERT CPORRECA

International Directory of Conchologists - $3.00

A current worldwide list of 2,000 people interested

in mollusks and in exchanging shells. Gives addresses

and interests. Available only from The Shell Cabinet.

Olive Shells of the World by Zeigler and Porreca -

$12.95

13 full-page color plates illustrating over 150 species

and named varieties. The only current treatment of

the popular family. Clothbound. Available from your

book store or directly from us.

Other Recently Published Shell Books Available

Post Office Box 29, Falls Church. Virginia 22046

Sea Shells of

Tropical West America

Marine Mollusks from Baja California to Peru

Second Edition. A. Myra Keen with the assistance of James H.

McLean. This revised and expanded classic now treats all molluscan

species recorded from the Panamic province and is illustrated with

over 4,000 halftones and 85 color photographs. The geographic range

has been extended to northwestern Peru. The coverage now extends to

microscopic and deepwater species and to the shell-less mollusks.

For most of the 3,325 species there are detailed descriptions,

synonymies, range data, notes, and one or more illustrations. Included

are anatomical drawings, identifying keys, introduction and

systematics, rejected species, maps and place-name list, glossary,

bibliography, and index. 1,120 pages. $29.50

Stanford University Press

Vol. 87 (2)

THE NAUTILUS

31

THE HABITS AND HABITATS OF SOME BERMUDIAN MARINE MOLLUSKS'

Thomas R. Waller

Department of Paleobiology

Smithsonian Institution

Washington, D. C. 20560

ABSTRACT

During a Jiving sun'ey of the living habits of scallops ( Pectinidae j in bermudian

waters, 160 species of shelled mollusks were collected at 40 diving stations. The

shallow-water bias in previous bermudian collecting is indicated by the fact that 17 of

these species have not previously been reported from Bermuda and 15 of these new

records are from depths greater than 30 m. Among the new records are five species of

the bivalve family Pectinidae, including Argopecten noronliensis (E. A. Smith),

previously known only from off Brazil but now known to be widely but sparsely

distributed in the tropical western Atlantic from Brazil to Bermuda.

Habitat preferences and living habits are described for several species, including a

discussion of species in Harrington Sound relative to the ecological zones described by

Neumann (1965). Twenty-one species inadequately figured in the literature are

illustrated using optical photographs and scanning electron micrographs.

INTRODUCTION

Despite the fact that Bermuda has been a haven

for malacologists since Angelo Heilprin and A. E.

Verrill first visited the islands in the late 1800's, the

habitats of Bermudian moUuscs are still poorly

known. Many of the older collections, althougli rich

in numbers of taxa, are poor in habitat data and

commonly list only "Bermuda" as the sole record

of occurrence for a species. The purpose of tliis

paper is to provide data on the habits and habitats

of species collected through diving and to illustrate

certain poorly known or previously unillustrated

species.^

During a period of two weeks in late May, 1971,

marine moUuscan communities were surveyed by

the author and two diving assistants using SCUBA

equipment. Althougli the primary purpose of the

survey was to gather data on the living habits and

habitats of scallops known from Bermudian waters,

collections of other shelled mollusks were made at

most of the diving stations. At most stations col-

lecting was by visual spotting and hand picking. In

the interest of conservation, many living individuals

were examined underwater and returned to their

habitat, with later identification assisted by dead

shells of the same species found at the site. At the

stations deeper than 30 m (Stations 21, 24, and

39), in the deepest portions of Harrington Sound

(Stations 1 and 6), and on the bottom of Wal-

singham Pond (Station 11), short non-decom-

pression time limits or turbid conditions prevented

visual collecting, and sediment samples in both net

and plastic bags were taken instead. These sediment

samples were retained in running salt water and

later examined in the laboratory for living material.

The resulting coUections contain about 160

species of shelled mollusks (Table 1) distributed

among 40 diving stations (Figs. 1, 2, and Table 2).

Surprisingly, 17 of these species have not been

previously recorded from Bermudian waters.^ Be-

cause all but two of these new records are from the

three diving stations deeper than 30 m (Stations 21,

24, and 39), it is clear that much of our present

'Contribution No. 546, Bermuda Biological Station

for Research, St. George's West, Bermuda.

^With regard to new records, 1 am indebted to Mr.

Russell Jenson, Assistant Curator, Department of

Mollusks, Delaware Museum of Natural History. It

is through his help and his extensive records,

compiled through a painstaking search of the liter-

ature on Bermuda, that I can with reasonable

confidence refer to species as having been

previously unreported.

32 THE NAUTILUS

April 1973

Vol. 87 (2)

knowledge of the Bermudian marine fauna is based

on shore and shallow-water coUections. Although

the West Indian character of the fauna has long

been known, it is probable that more intensive

collecting by means of diving (deeper than 45 m)

and other methods not avaUable to early collectors

will continue to turn up a wealth of species pre-

viously thougjit to be restricted to the West Indies

proper or to the Atlantic coast of South America.

The stations listed below are plotted in Figures 1

and 2. Groupings of stations according to general

environment or ecological zone are shown in Table

2.

Station 1. DevO's Hole, Harrington Sound.

Depth, 25 m; bottom temperature, about 20° C;

barren mud bottom with mounds, trails, fine ex-

crement, and small shells. May 19, 1971.

Station 2. Devil's Hole, Harrington Sound, on a

line extending southeastward from Station 1. Depth,

17 to 9 m; fine sand bottom with abundant

Oculina, Area, and Pododesmus. May 19, 1971.

Station 3. Devil's Hole, Harrington Sound, on a

line extending southeastward from Station 1. Depth,

9 to 6 m; coarse sandy bottom with little vegetation

and abundant Area zebra. May 19, 1971.

Station 4. Rocky shore on east side of Devil's

Hole, Harrington Sound. Depth, 0 to 3 m. May 19,

1971.

Station 5. Between Trunk Island and Small Is-

land, Harrington Sound. Depth, 3 to 4 m; sandy

FIG. I . Map of the Bermuda Islands showing loeations of diving stations.

Vol. 87 (2)

THE NAUTILUS

33

bottom with scattered patches of grass. May 19,

1971.

Station 6. Shark Hole, Harrington Sound, along

traverse extending north northwestward from a

depth of 21 m. Depth, 21 to 18 m; bottom

temperature, 19° C; barren mud bottom witli

mounds, trails, scattered shells, and Oculina frag-

ments. May 20, 1971.

Station 7. At 32° 20.2' N, 64° 42.5' W, near

Shark Hole, Harrington Sound. Depth, 18 to 11 m;

fine sand bottom with abundant Oculina. May 20,

1971.

Station 8. Near Station 7, Harrington Sound.

Depth, 8 to 1 1 m. May 20, 1971. No collection was

made at this station.

Station 9. Midway between Abbot's Cliff and

Trunk Island, Harrington Sound. Shoal area with

large areas of barren, sandy bottom covered with

the shells of Area. May 20, 1971. No collection

made.

Station 10. Southwest side of Trunk Island,

Harrington Sound. Depth, 3 to 6 m; sandy bottom

with patches of grass. May 20, 1971.

Station 11. Walsingham Pond, Bermuda Island.

Depth, 1.5 to 6 m; barren, dark, soupy mud bottom

with film of organic matter and small shells. May

21, 1971.

Station 12. Seaward side of rocks at mouth of

Whalebone Bay, western end of St. George's Island.

Depth, 4.5 m; water temperature, 22°C; coarse sand

bottom with abundant hard coral and Gorgonacea.

May 22, 1971.

Station 13. Western end of Church Bay, Har-

rington Sound. Depth 3 to 6 m. May 22, 1971.

Station 14. Hungry Bay, on south shore of

Bermuda Island. Depth, 1 m; coarse sand bottom

FIG. 2. Map of Harrington Sound, Bermuda, showing locations of diving stations with respect to the zonation

of Neumann (1965).

34 THE NAUTILUS

April 1973

Vol. 87 (2)

with broad-bladed marine grass and the alga, Pedina.

May 24, 1971.

Station 15. Outside mouth of Hungry Bay, on

south shore of Bermuda Island. Depth, 4.5 to 8 m;

water temperature, 22° C; bottom of coarse sand

and bare rock with potholes. May 24, 1971.

Station 16. Off small bay between Tobacco Bay

and Coot Pond, northern St. George's Island. Depth,

6 to 8 m; water temperature 22 to 23° C; coarse

sand bottom with large and small coral-covered

rocks. May 24, 1971.

Station 17. Buildings Bay, near eastern tip of St.

George's Island. Depth, 0.5 to 3.5 m; very coarse,

ripple-marked sand bottom with some large rocks

and abundant green to lavender leafy algae. May 24,

1971.

Station 18. At 32° 22.4' N, 64° 39.2' W, off

mouth of pass between Paget Island and St. Davids

Island. Depth, 6 to 7 m; water temperature, 21.5°

C; coarse, ripple-marked sand with patches of coral

and leafy algae. May 25, 1971.

Station 19. Near Station 18, but further from

shore. Depth, about 6 m. May 25, 1971. No

collection.

Station 20. Higgs Cut, between Higgs Island and

Paget Island at eastern end of St. George's Harbor.

Depth, 0.5 to 4.5 m (within small boat channel and

surrounding area) and 6 m (deep hole at western

end of cut); bottom fine sand and grassy in shallow

water, barren sand at bottom of channel. May 25,

1971.

Station 21. At 32° 30.6' N, 64° 38.2' W, north-

east of "North East" marker buoy. Depth, 37 m;

bottom water temperature, 20.5° C; small coral

knolls rising a few feet from the bottom and

separated by patches of very coarse, ripple-marked

sand. May 26, I97I.

AH specimens recorded from this station were

picked from bulk samples taken from the sand

patches.

Station 22. Anchor chain of "North East"

marker buoy, 32° 30.2' N, 64° 38.5' W. Depth, 6

m. May 26, 1971. No collection.

Station 23. Top of large patch reef between St.

George's Island and "North East" marker buoy

(Station 22). Exact position not plotted. Depth, 18

m. May 26, 1 97 1.

Station 24. At 32° 18.2' N, 64° 40.8' W, off

south shore of Tucker's Town. Depth, 34 m; bot-

tom temperature, 20.5° C; higli coral patches with

scattered small areas of coarse sand. May 27, 1971.

Station 25. At 32° 19.6' N, 64° 41.4" W, inside

breaker zone off south shore of Tucker's Town.

Depth, 6 to 8 m; patch reefs surrounded by coarse,

ripple-marked sand. May 27, 1971.

Station 26. At 32° 22.6' N, 64° 39.0' W. off

entrance to St. George's Harbor. Depth, 6 to 18 m;

coral platform rising 12 m above a medium to fine

sand bottom May 27, 1971.

Station 27. At 32° 23.9' N, 64° 40.9' W, north

northeast of St. Catherines Point, St. George's Is-

land. Depth. 14 m; water temperature, 22° C: silty

mud bottom with large colonies of Ociilina. May

28, 1971.

Station 28. At 32° 24.8' N. 64° 41.2' W, north-

east of St. Catherines Point, St. George's Island.

Depth, 4.5 to 8 m; patch reef with prolific, diverse

coral. May 28, 1971.

Station 29. South side of Ferry Point, western

tip of St. George's Island, from rocky shore to a

depth of 4.5 m in the channel between Ferry Point

and Coney Island. May 29. 1971.

Station 30. At 32° 21.6' N, 64° 42.7' W, in

shallow area in mouth of Ferry Reach southwest of

Ferry Point. Depth, 1.5 to 3.5 m; Ociilina patch

reef and adjacent sandy bottom. May 29, 1971.

Station 31. North half of Ely's Harbor, Somerset

Island. Depth, 1.5 to 6 m. May 30, 1971.

Station 32. Off mouth of Ely's Harbor, Somerset

Island. Depth, 2.5 m; patch reef on sandy bottom.

May 30, 1971. No collection.

Station 33. South half of Ely's Harbor, Somerset

Island. Depth, 1.5 to 6 m. May 30, 1971.

Station 34a. In channel between Wilsons Island

and Bermuda Island, Port Royal Bay. Depth, 1.5 to

4 m; fine silty bottom with mounds, worm tubes,

and few holothurians. May 30, 1971. No collection.

Station 34b. East side of Wilsons Island, Port

Royal Bay. Depth. I to 4.5 m; sandy, trash-covered

bottom at base of steep, rocky drop-off. May 30,

1971.

Station 35. North end of Grace Island, Great

Sound. Depth, 4.5 m; barren, fine sand bottom at

base of rocky drop-off. May 30. 1 97 1. No

collection.

Station 36. Between Hunts Island and Perots

Island, Port Royal Bay. Depth, 3.5 to 5.5 m; firm,

clean sand bottom with grass in shallow areas. May

30, 1971. No collection.

Vol. 87 (2)

THE NAUTILUS

35

Station 37. Transect across channel between Burt

Island and Darrell Island, Great Sound. Depth, 1 to

4 m; firm, fine sand with patches of grass. May 30,

1971.

Station 38. Channel between Coney Island and

Bermuda Island at northeastern tip of Bermuda

Island. Depth, 0.5 to 4.5 m. May 31, 1971.

Station 39. At 32° 18.1' N, 64° 40.2' W, off

south shore of Tucker's Town. Depth, 51 m;

rubbly, coarse, foraminiferal sand with abundant

dark red algal nodules. June 1, 1971.

NOTES ON HABITATS

Harrington Sound

Siibthermocline Zone. Neumann (1965) has pro-

vided an excellent account of the bathymetry,

hydrography, and ecological zonation of Harrington

Sound. As described by Neumann (p. 1003), the

SubthermocUne Zone (Fig. 2) has an abrupt upper

boundary at a depth of 17 m (55 feet), at which

the extensive, essentially barren mud bottom of this

zone meets an area of abundant branching, bushy

coral (Oculinaj, characterized by Neumann as the

Oculina Zone. This zonal boundary is the result of

seasonal water stratification and coincides with the

deepest position of a thermocline in summer.

During the summer there is little mixing of water

beneath the thermocline, resulting in cooler, ox-

ygen-poor water below. Neumann described the

subthermocline bottom as a smooth and barren

surface marked only by the scattered holes of

burrowing organisms, possibly crustaceans, and

having patches of a thin purplish algal or bacterial

mat. He reported the valves of the small bivalve

Transennella conradina (but see below) as common

on the mud surface, with a few live specimens, and

added that this species is the only significant cal-

careous form.

Our observations confirm Neumann's. At Station

1, near Devil's Hole, we found the upper boundary

of the Subthermochne Zone to be sharp and to he

precisely at 17 m. Although underwater visibility

was less than 1 m, we observed that the mud

bottom was marked with numerous mounds, traUs,

small dead moUusk shells, and a Utter of fine

excrement. The dominant bivalve is Gouldia cerina

(see Notes on Taxa), not Transennella as reported

by Neumann, and all specimens were dead. In

addition we recovered a single valve of a young

Pitar fulminata as well as numerous dead specimens

of a tiny diastomid gastropod (Fig. 26), Finella

dubia (Orbigny).

The traverse comprising Station 6 (Fig. 2) ap-

proached the upper boundary of Neumann's Sub-

thermocline Zone, which here is within a depth

interval of 17 to 18 m. We observed that the actual

thermocline was very close to this boundary, with

water temperature rising from 19° C below the

thermocline to 21 above. The more varied fauna

than that collected from the same zone at Station 1

is most hkely due to the fact that we were near the

upper boundary of the Subthermocline Zone at

Station 6. Most of the shells collected are rather

old, the exceptions being the abundant Gouldia and

less abundant Pitar, the same bivalves represented at

Station 1. The valves of Argopecten gibbus collected

here are of particular interest and are discussed

under Notes on Taxa.

As pointed out by Stanley (1970, p. 83), suspen-

sion feeders (such as Gouldia and Pitar) living on

soupy muds face the problem of being clogged by

loose, fine-grained sediment as they sink into the

substrate. Most inhabitants of this type of bottom

remain small, thus keeping their surface-volume

ratio large and maximizing support from the sub-

stratum per unit of animal weight. Gouldia appears

to be an opportunistic species that colonizes the

Subthermocline Zone at times when conditions are

most favorable, probably during the time of max-

imum water mixing in the winter. The species then

flourishes until conditions become lethal, probably

during maximum summer stagnation.

Oculina Zone. Stations 2 and 7 in Harrington

Sound are within the Oculina Zone of Neumann

(1965, p. 1001), an area between 9 and 17 m (30

and 55 feet) in depth characterized by a fine sand

bottom and abundant growth of the branching

coral, Oculina valenciennesi. As described by Neu-

mann, the coral supports itself on the soft mud by

forming bush-like growths on its own debris or on

clusters of Area zebra. Area zebra is by far the most

common mollusk in this zone and lives byssally

attached in clusters on or near the bottom. Other

common bivalves (Table 1) are Pododesmus rudis

(attached to Oculina), Chama congregata (attached

to any hard object), and Lithophaga bisculcata

(boring into Oculina). Neumann described

Vermicularia as abundant in the Oculina Zone, and

Gould (1969, p. 432) has also noted the common

association of Vermicularia and Oculina. Neumann's

36 THE NAUTILUS

April 1973

ol. 87 (2)

report of Anomia simplex as common in this zone

could not be confirmed, and it is probable that he

was referring to Pododesmus, which was absent

from his list.

Shallow Sandy Zone. Stations 3, 5, 8-10, and 13

lie within the Shallow Sandy Zone of Neumann

(1965, p. 1001), an area extending in depth from 9

m to the surface (Fig. 2). As described by Neu-

mann, the sandy floor is covered by marine grasses,

sponges, and algae in the well-lit regions down to

about 4.5 m. Below 4.5 m grass is less abundant

and there are more open sandy areas, frequently

with large clusters oi Area zebra.

The western end of Church Bay, Harrington

Sound (Station 13, Fig. 2), within the harbor of a

private estate, has a silty sand bottom with little

vegetation, common Cassiopeia (the bottom-dwelling

jellyfish), and common Anadara notabilis. About 30

m. out from shore the bottom is more typical of

the Shallow, Sandy Zone, with patches of marine

grass, coarser, barren sand, abundant Area zebra,

and many living Argopecten gibbus, Pecten ziczac,

and Macrocallista maculata. Over sandy bottoms

near the rocky shore of Church Bay, both Pinctada

and Anomia are common on the abundant bottles

and other trash that litter the bottom.

Walsingham Pond

Twenty years ago De Laubenfels (1950, p. 194),

in his colorful style, described the rock walls of

Walsingham Pond as a sponge metropolis and noted

that the bottom is largely mud-covered, but so deep

that the water is clear. A more detailed summary of

conditions in the pond was provided by Walton

(1969), who studied the bathymelr), sediments, and

water chemistry. Tlie maximum depth is about 6 m

and the bottom througliout the deeper portions of

the pond consists of soupy, pelleted mud covered

by a mat of organic matter 5 to 10 cm. thick. The

most abundant moUusks in this soupy, mud layer

are, according to Walton (op. cit., p. 44), Batillaria,

Vermicularia, and Gouldia.^ Beneath this layer is a

thicker layer of brown pelleted mud with more

abundant shells interpreted by V.alton to represent a

time when the pond was less restricted than at

present. Gould (1968) studied the moUuscan fauna

of Walsingham Pond, listing 9 species of bivalves

and 15 species of gastropods. Although he did not

distinguish living from dead or fossil material, he

stressed the similarity of the overall assemblage to

other mangrove-mud environments in Bermuda.

Our collecting concentrated on the mud bottom

of the southwest arm of the pond. Contrary to the

pristine quality of water implied by De Laubenfels'

description, liglit penetration to the bottom of the

pond is now very low, and it was necessary to use

liglits in order to see small objects on the bottom.

Underwater visibility was poor and would rapidly

drop to zero if we allowed our trail of disturbed

mud to overtake us. As noted by Gould (1968), the

bottom harbors a profusion of the bottom-dweUing

jeUyfish, Cassiopeia. Below the organic tllm and

soupy layer is an accumulation of small shells.

^I infer that Walton's "Batillaria minima" is actually

Cerithium variabile and his Gouldia is actually

small Ctena orbiculata.

FIGS. 3-15. Bermuda Marine MoUusks. All specimens in optical photographs are coated with ammonium

chloride and those in scanning electron micrographs coated with gold unless indicated otherwise.

FIGS. 3, 4. Matching left and right valves of Aequipecten acanthodes (Dallj, syntype, USNM 333637, Santa

Lucia, Cuba, height 31.5 mm,

FIGS. 5-7. Right valve uncoated, matching left valve coated, and sculptural detail of left valve f.x 9),

Argopecten noronhensis (E. A. Smith), USNM 501857, Barbados, height 15.0 mm.

FIG. 8. Fragment of left valve of Argopecten noronhensis (E. A. Smith), USNM 708130, Bermuda Sta. 39,

height of fragment 8.0 mm

FIGS. 9-11. Chlamys multisquamata (Danker), matching left and right valves of specimen from off Palm

Beach, Florida, in collection of F. M. Bayer, height 46.8 mm, and fragment of valve, USNM 708131, Bermuda

Sta. 39, height of fragtnent 7.0 mm.

FIGS. 12, 13. Structural detail of outer surface (x 17) and inner surface of right valve of Neopycnodonte

cochlear (PoH), USNM 708132, Bermuda Sta. 39, height 12.5 mm.

FIGS. 14, 15. Matching left and right valves of Limaria hyalina (Verrill and Bush), USNM 708133, Bermuda

Sta. 39, height 4.1 mm.

Vol. 87 (2)

THE NAUTILUS

37

,|ii'

% m

A

r4'.uW,

14

Bermuda Marine Mollusks

(explanation on opposite page)

38 THE NAUTILUS

April 1973

Vol. 87 (2)

primarily Vermicularia, Cerithium, and Ctena

(Table 1). Live individuals of these species were

brought up in samples, but their living positions in

the substrate were not observed. Seven species of

bivalves and 26 species of gastropods were re-

covered, with the most abundant species essentially

the same as those hsted by Gould (op. cit., p. 3).

However, our samples, obtained by visual collecting

and digging in the muddy bottom by hand, con-

tained no Gouldia cerina, listed as "very common"

by Gould, and numerous Anachis catenata, not

recovered by Gould. Many of the less common

species in the pond (see Table 1, Station 11)

undoubtedly come from the fossil shelly layer men-

tioned above.

Although many of the species Usted may have

dropped off the rock walls of the pond, many also

probably hve on the organic debris and algae coat-

ing the muddy bottom, as suggested by a com-

parison of the list of these species with the Ust of

molluscan-algal associations of Warmke and

Almodovar (1963). The abundance of Ctena

orbiculata in the pond may reflect stress conditions.

Jackson (1970, p. 586) found that lucinids are

tolerant of stagnant conditions involving the pres-

ence of hydrogen sulfide and, unlike most other

bivalves, are able to feed on microbial populations

present on plant detritus.

Other Localities

Hungry Bay. Hungry Bay (Station 14), on the

south shore of Bermuda Island, is very shallow, less

than 1 m. deep at low tide, with a bottom of coarse

sand vvdth broad-bladed marine grass and coral frag-

ments. The small alga, Pedina, is also abundant, and

it is in this shallow area that we found many small

(5 mm) Chione. At the entrance to the bay and out

to a depth of about 8 m (Station 15), the max-

imum depth of this dive, the bottom is rock with a

thin algal covering broken here and there by the

tootli marks of large parrot fish. Pot holes up to 1

m. deep and some deep channels have been scoured

out of the rock, and the turbulence of the area has

left few rocks small enough to overturn. Under the

few that could be overturned, Barbatia domingensis

was common. Malleus candeanus was seen far be-

FIGS. 16, 17. Right valve and hinge (x 15) of Erycina sp. aff. E. linella Dall, USNM 708134. Bermuda Sta.

24, length 6.2 mm

FIG. 18. Right valve of Carditopsis smitlii (Dall), scanning electron micrograph, USNM 708135, Bermuda Sta.

11, height 1.7 mm.

FIG. 19. Prodissoconch o/ Carditopsis smithi (Dall), scanning electron micrograph, USNM 708136, Bermuda

Sta. 11, length 0.39 mm.

FIG. 20. Alvania didyma (Watson), scanning electron micrograph, USNM 708137, Bermuda Sta. 21, height 2.1

mm

FIG. 21. Alvania platycephala Dautzenberg and Fischer, scanning electron micrograph, USNM 708138,

Bermuda Sta. 11, height 2.4 mm,

FIG. 22. Caecum delicatulum Verrill and Bush, scanning electron micrograph, USNM 708139, Bermuda Sta.

11, height 2.9 mm

FIG. 23. Cerithiopsis cyntliia? Bartsch, USNM 708140, Bermuda Sta. 24, height 5.0 mm.

FIG. 24. Triphora mirabilis (C. B. Adams), USNM 708141, Bermuda Sta. 24, height 9.0 mm.

FIG. 25. Pusia variata? (Reeve), USNM 708142, Bermuda Sta. 21, height 18 mm.

FIG. 26. Finella dubia (Orbigny), USNM 708143, Bermuda Sta. 1, height 3.3 mm.

FIG. 27. Aesopus stearnsi Tryon, USNM 708144, Bermuda Sta. 21, height 4.9 mm.

FIG. 28. Thala foveata (Sowerby), USNM 708145, Bermuda Sta. 24, height 4.4 mm.

FIG. 29. Pyrgocythara candidissima (C. B. Adams), USNM 708146, Bermuda Sta. 39, height 4. 7 mm.

FIG. 30. Odostomia didyma Verrill and Bush, scanning electron micrograph, USNM 708147, Bermuda Sta. 11,

height 1. 8 mm.

FIG. 31. Mitrolumna bipUcata (Dall), USNM 708148, Bermuda Sta. 39, height 3.9 mm.

FIG. 32. Odostomia nioba Dall and Bartsch, scanning electron micrograph, USNM 708149, Bermuda Sta, 21,

height 2.5 mm.

FIG. 33. Turbonilla heilprini Bush, scanning electron micrograph, USNM 708150, Bermuda Sta. 11, height 2.8

mm

Vol. 87 (2)

THE NAUTILUS

39

16

17

^^^i''^^ 18

^*5

iiv

Bermuda Marine Mollusks

(explanation on opposite page)

40 THE NAUTILUS

April 1973

Vol. 87 (2)

neath the overhangs of pot holes and channels,

whereas Chama was common atop the bare rock

surface.

Ely's Harbor. Almost the entire area of Ely's

Harbor was surveyed for the presence of scallops by

towing divers behind the boat, but no living spec-

imens were found. In the north half of the harbor

(Station 31), the bottom in the center is soft,

barren mud or silt with some coral-covered rocks.

Peripheral areas are sandy and grassy with scattered

clusters oi Area zebra.

North Lagoon. The bottom features and fauna at

Station 27, north of St. George's Island, are very

similar to those in the Oculina Zone of Harrington

Sound (Neumann, 1965; see preceding notes) with

the dominant mollusks being Area zebra, Po-

dodesmus rudis, Vermieularia, Spondylus, and

Chania. Many dead Chlamys imbricata, not found in

Harrington Sound, were present at this station, but

no living individuals could be found after an in-

tensive search.

Outer Platfonru The deepest areas sampled lie

well outside of the breaker zone: Station 21 (depth,

37 m). Station 24 (depth, 34 m) and Station 39

(depth, 51 m). The bottom at these stations consists

of patches of barren coarse carbonate sand, gen-

erally ripple marked, with intervening patches of

coral and algae. Tlie sand at Station 39 consists

almost entirely of the tests of the foraniinifer,

Amphistegiim gibbosa. and the bottom is strewn

with maroon algal nodules. Bulk samples of the

sand yielded a profusion of small shells, particularly

gastropods such as Euchelus giittarosea, Zebina

browniana, various cerithiopsids, Triphora mirabilis.

Trivia quadripimctata. various columbellids and

margineUids, and Williamia krcbsiL Small bivalves

are also common, particularly Area imbrieata,

Barbatia domingensis. fragmented Comptopallium

antillanim, Spondylus, Lima lima, Chama sarda,

Trigonioeardia media, and Gouldia eerina. The

bivalve Ventrieolaria rugatina, previously unknown

from Bermuda, increases in both size and abundance

from Station 24 to the deepest station (Station 39).

In addition to the small mollusks, the tiny

brachiopod, Argyrotheea benmidana (Dall, 1911), is

common in the sand at Stations 24 and 39. Pre-

sumably these specimens, like the many byssate and

cemented bivalves that are also found in the sed-

iment, have become detached from living sites

among and beneath small corals.

The fact that 1 5 out of 1 7 new records listed in

Table 1 are from the three outer-platform stations is

a striking indication of how httle is known about

the Bermudian molluscan fauna from these depths.

TABLE L List of speeies of shell-bearing Mollusca eollected in Bermuda, with references to

illustrations and eollecting stations. The order of families follows that of Moore (1969) for the

bivalves and Taylor and Sohl (1962) for the gastropods. Taxa are alphabetically arranged within

each family. Station numbers in bold-face type indicate living specimens; other specimens are

represented by shells only. Species not previously known from Bermuda are preceded by an

asterisk (*). Specimens that document the list are deposited in the collections of the

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian

Institution, Washington, D. C.

Species

BIVALVIA

Anadara notabilis (Roding, 1798)

Area imbricata Bruguiere, 1789

Area Zebra (Swainson, 1 833)

Barbatia (Area) domingensis (Lamarck, 1819)

*Barhatia (Fugleria) tenera? C. B. Adams, 1845

Barbatia sp.

Arcopsis adamsi (E. A. Smith, 1888)

Brachidontes sp.

Lithophaga bisulcata (Orbigny, 1 842)

Illustration

Stations

Warmke and Abbott 3, 5, 10, 13

Warmke and Abbott 12, 18, 21, 24, 27, 39

Warmke and Abbott 2, 3, 5, 6, 7, 9, 10, 13, 27

29, 31,33, 34b, 35, 39

Warmke and Abbott 11, 12, 15, 18, 21, 24, 27, 39

Warmke and Abbott 39

24

Warmke and Abbott 6, 24

Warmke and Abbott 2, 7

Vol. 87 (2)

THE NAUTILUS

41

Species

Modiolus amcricamis Leach, 1815

Musculus lateralis (Say, 1822)

Pinna carnea Gnielin, 1791

Pinctada imhricata Roding, 1 798

Pteria colymbus (Roding, 1798)

Isognomon radiatus (Anton, 1839)

Isognomon sp.

Malleus (Malvufundus) candeanus (Orbigny, 1842)

Aequipecten acanthodes (Dall, 1925)

Argopecten gibbus (Linndiews,, 1758)

*Argopecten noronhensis (E. A. Smith, 1885)

*Chlamys benedicti Verrill and Bush, 1897

Chlamys imbricata (Gmelin. 1791 )

*Chlamys multisquamata (Dunker, 1864)

*Comptopallium antillanim (Recluz, 1 853)

*Pecten chazaliei Dautzenberg, 1900

Pecten ziczac (Linnaeus, 1758)

Spondylus americanus Hermann, 1781

Anomia simplex Orbigny, 1842

Pododesmus rudis (Broderip, 1834)

L/ma L(wa (Linnaeus, 1758)

*Limaria hyalina (Verrill and Bush, 1 898)

*Neopycnodonte cochlear (Poli, 1795)

Lop/;a /o //a (Linnaeus, 1758)

Anodontia schrammi (Crosse, 1876)

Codakia orbicularis (Linnaeus, 1758)

Ctena orbiculata (Montagu, 1808)

Parvilucina costata (Orbigny, 1842)

Erycina sp. aff. E. linella Dall, 1 899

Lasaea adansoni (GmtWn, 1791)

Carditopsis smithi (DaW, 1896)

Charna congregata Conrad, 1833

Chama macerophylla Gmelin, 1791

*Chama sarda Reeve, 1 847

Chama sp.

Pseudochama radians (Lamarck, 1819)

Crassinella lunulata (Conrad, 1 834)

Laevicardium laevigatum (Linnaeus, 1758)

Laevicardium sp.

Papyridea semisulcata (Gray, 1 825)

Trigoniocardia media (Linnaeus, 1758)

Ervilia concentrica (Holmes, 1 860)

Florimetis intastriata (Say, 1 826)

Macoma tenta Say, 1834

42 THE NAUTILUS

April 1973

Vol. 87 (2)

Species

Strigilla mirabilis (Philippi, 1841)

Tellina (Acon'his) guitldii Hanley. 1846

Tellina (Aiignlus) paramera Boss, 1964

Tellina (Laciolina) laevigata Linnaeus. 1758

Tellina (Laciolina) magna Spengler, 1 798

Tellina (Scissulaj similis Sowerby, 1 806

Tellina (Tellina) radiata Linnaeus, 1758

Tellina (Tellinella) listen Roding. 1798

Semelina proficua (Pulteney. 1799)

Tagelus divisus (Spengler, 1794)

Coralliophaga coralliophaga (Ginelin, 1791)

Macrocallista maculata (Linnaeus. 1758)

Chione cancellata (Linnaeus, 1767)

Gouldia cerina (C. B. Adams, 1845)

Pitar fulminata {Menke, 1828)

*Ventricolaria ntgatina (Heilprin. 1887)

*Corbula (varicorbula) sp.

Gastrochaena Mans (Gmelin, 1791)

GASTROPODA

Diodora dysoni (Reeve. 1850)

Diodora sp.

Emarginula pwnila (A. Adams 1851)

Acmaea pustulata (Helbling, 1779)

Lepeta? sp.

Euchelus giittarosea Dall. 1 889

Synaptocochlea picta (Orbigny, 1 842)

Astraea phoebia Roding, 1798

Truncatella caribaeensis Reeve, 1 842

Alvania didyma (Watson, 1886)

Alvania platycephala Dautz. and Fischer. 1896

Cingiila? sp.

*Micwdochiis jloridanus Rehder, 1943

Rissoina biyerea (Montagu. 1803)

Rissoina sp. aff. /?. /jn'e/rfl (Montagu. 1803)

Rissoina cancellata Philippi. 1 847

Rissoina fischeri Desjardin, 1949

Zebina bwwniana (Orbigny. 1842)

Vitrinella species A

Vitrinella species B

Vitrinella species C

*Spirolaxis exquisita (Dall and Simpson. 1901)

Vermicularia knorrii (Deshayes, 1843)

Vermicularia spirata Philippi, 1 836

Siliquaria sqiwmata Blainville, 1827

Caecum delicatulwn Verrill and Bush, 1900

Caecum plicatum Carpenter. 1 858

Caecum sp.

Modulus modulus (Linnaeus. 1758)

Illustration

Boss (1969)

Boss (1966)

Boss (1968)

Warmke and Abbott

Boss (1966)

Warmke and Abbott

Abbott

Warmke and Abbott

Our Fig. 20

Our Fig. 21

Warmke and Abbott

Gould (1969)

Gould (1966)

Our Fig. 22

Warmke and Abbott

Stations

26

16. 24

24

5. 10. 24. 25

10. 37

24

25

10, 37. 38

16

11

24

3

27

5, 13

11. 14

1. 2. 6. 16,

1. 6. 16

21. 24, 39

39

M. 24, 39

Warmke and Abbott 24

17

5

24

24. 39

24

21, 24. 39

24

24. 39

11

21. 24

11

24

11

24

39

11

11. 24

21, 24. 39

11

24. 39

39

24

24. 27, 39

11

24

II

11, 24.39

11

11, 16, 39

Vol. 87 (2)

Species

Finella dubia (Orbigny, 1 842)

Alaba incerta (Orbigny, 1842)

Cerithium litteratum (Born, 1778)

Cerithium variabile (C. B. Adams, 1845)

Cerithiopsis ara Dall and Bartsch, 1911

Cerithiopsis cynthia? Bartsch, 1911

Cerithiopsis nigiilosum (C. B. Adams, 1 850)

Cerithiopsis vicola Dall and Bartsch, 1911

Cerithiopsis species B

Cerithiopsis species C

Cerithiopsis species D

Triphora mirabilis (C. B. Adams, 1850)

Triphora species B

Triphora species C

Triphora species D

Triphora species E

Bemiudaclis bermudensis (Dall and Bartsch, 1911)

Strombus costatus Gmelin, 1791

Trivia quadripunctata (Gray, 1 827)

Cypraea cinerea Gmelin, 1791

*Cypraea spurca acicularis Gmelin, 1791

Cypraea sp. (Bulla stage)

Natica sp.

*Aspella elizabethae McGinty, 1940

Murex (Favartia) cellulosus Conrad, 1 846

*Pusia variata? (Reeve, 1845)

Aesopus stearnsi Tryon, 1 883

Anachis catenata (Sowerby, 1 844)

Columbella mercatoria (Linnaeus, 1758)

Mitrella lunata (Say, 1 826)

*Nassarina monilifera (Sowerby, 1844)

Colubraria swifti {Tryon, 1881)

Colubraria testacea Morch, 1877

Nassarius albus (Say, 1 826)

Thala foveata (Sowerby, 1 874)

Latirus brevicaudatus (Reeve, 1847)

Hyalina avena (Kiener, 1 834)

Hyalina lactea (Kiener, 1841)

Persicula lavalleeana (Orbigny, 1 842)

Persicula? species A

Persicula? species B

Mitra nodulosa (Gmelin, 1791)

Glyphoturris quadrata rugirima (Dall, 1889)

Mitrolumna biplicata (Dall, 1 889)

Pyrgocythara candidissima (C. B. Adams, 1 845)

Conus bermudensis Clench, 1942

Odostomia didyma Verrill and Bush, 1900

Odostomia nioba Dall and Bartsch, 1911

Turbonilla heilprini Bush, 1 899

43

44 THE NAUTILUS

April 1973

Vol. 87 (2)

Species

Turbonilla pupoides (Orbigny, 1842)

Bulla striata (Bruguiere, 1792)

Atys sp.

Hammoea succinea (Conrad, 1846)

Haminoea sp.

Retitsa candei Orbigny, 1841

WilliamJa krehsii (Morch, 1 877)

Laemodonta cubensis (Pfeiffer, 1854)

Pedipes mirabilis (Mulilfeldt, 1816)

Pleurobranclnts sp.

SCAPHOPODA

Cadulus qiiadridentatus {Dan, 1881)

POLYPLACOPHORA

Radsiella mgidata (Sowerby, 1832)

Clench (1964)

39

Warmke and Abbott 24

Keen

39

NOTES ON TAXA AND THEIR HABITS

Bivalvia

Arcidae

The similarity of living habits of Area imbricata

and Area zebra have been emphasized by Stanley

(1970, p. 122), who also stated that both prefer

shallow subtidal habitats. Our collections indicate,

however, that in Bermuda Area imbricata is more

common in open marine situations than in pro-

tected sounds (see Table 1). Tlie only co-occurrence

of living specimens was in the North Lagoon at

Station 27, where conditions similar to those in the

Oeuliiia Zone of Harrington Sound exist on the

open platform.

Area zebra is perhaps the most conspicuous

mollusk in the shallow inshore waters of Bermuda.

It is especially abundant in Harrington Sound,

where it can be found at depths from about 1.5 m

to 17 m within the Shallow Sandy Zone and

Oculina Zone of Neumann (1965). Secreting a

strong, pedestal-like byssus. Area zebra attaches to

the top or sides of any hard object, most commonly

to shells, rocks, or other debris on the sandy

botton^ Subsequently, other individuals attach to

one another, resulting in clusters or knots which

themselves form a substrate for epifauna. Each

individual Area raises itself on its byssus while

filtering but quickly lowers itself when sensing a

passing shadow. Area zebra does not colonize the

rocky shore, nor does the species attach to corals

far above the bottom. At Station 29, Area zebra

clusters have formed on the old concrete pilings of

the abandoned causeway, but these clusters are

subtidal and close to the bottom.

In contrast to its inshore abundance, the paucity

or absence of Area zebra from the outer shallow

waters of Bermuda is striking. It is usually un-

common in smaU bays with free access to the open

sea and is not common in the shallow waters within

the breaker zone of the south shore. The only

outside area at which Area zebra was found in

abundance is Station 27 in an area of abundant

Oeidina on a silty mud bottom, very similar to the

Oeiilina Zone of Harrington Sound.

In a study of Barbatia dommgensis on the Ber-

muda Platform, Bretsky (1967) found that this

species is most abundant in turbulent environments

and least abundant in protected bays and sounds.

Its most common site of byssal attachment is

beneath corals, less commonly beneath rocks. The

strong representation of B. domingensis in our col-

lections from the open platform confirms Bretsky's

observations, and in view of these habitat pre-

ferences only the occurrence of the species in

Walsingham Pond (Station 1 1 ) is unexpected. Tlie

pond specimens are all immature and possibly in-

dicate that larvae forced into the pond through

crevices are unable to reach maturity.

Vol. 87 (2)

THE NAUTILUS

45

Pteriidae, Isognomonidae, and Malleidac

Among the Pteriacean bivalves (Pteria, Pinctada,

Isognomon. and Malleus), only Pteria colymbits lives

off the bottom, attached by a strong, clasping

byssus to the tlexible, waving arms of gorgonaceans.

At all stations where living individuals were ob-

served Pinctada radiata attaches to shells or rocks

on the sandy bottom, generally with the plane of

commissure nearly horizontal or somewhat inclined

to the substrate. Wliereas Pteria was obsei-ved only

in open-marine areas, Pinctada is common in bays

and sounds.

Isognomon radiatus was observed living only at

two open-marine stations (Stations 1 5 and 25),

where individuals were byssally attached between

large rocks. According to Stanley (1970, p. 136),

nestling beneath or between rocks with weak byssal

attachment is the usual living habit of the species.

Orientation relative to the attachment surface is

variable, although generally the distal portions of

the shell are erect and project into the water rather

than lie flat against the attachment surface.

Malleus candeanus is a rare species throughout

the West Indies and Florida but occurs in abun-

dance at Bermuda. The taxonomy, distribution and

living habits of the species have been described by

Boss and Moore (1967), and Yonge (1968) has

given a detailed account of the functional morph-

ology of the very similar Pacific species, M. regula.

Although Boss and Moore did not observe living

specimens, they concluded that Malleus candeanus

lives attached to hard substrates, particularly in the

crevices of coral or coralline rock. Our underwater

observations confirm this habit. At all localities

observed, specimens were found tightly byssally

attached to the sides and undersurfaces of large

rocks, particularly where there are large crevices or

small caves between rocks, or beneath overhanging

ledges of coral. At Station 26, Malleus was par-

ticularly common on the high; steep side of a large

patch reef. All localities were in open marine areas

generally deeper than 6 meters and with con-

siderable current or surge flowing between the

rocks.

The elongate, distal portion of the shell of

Malleus projects from the attachment surface. This

portion of the shell is thin and composed entirely

of fragile prismatic calcite. Although it is frequently

fractured during attacks by nibbling predators,

Yonge (1968, p. 387) has pointed out that

prismatic calcite can be rapidly regenerated by the

highly letractible mantle, which remains uninjured.

Pectinidae

Both Comptopallium antillarum (Recluz, 1853)

and Pecten cbazaliei Dautzenberg, 1900, have not

been previously reported from Bermuda. Shells and

fragments of both were recovered from sediment

samples taken on dives deeper than 30 m. (Stations

24 and 39). Comptopallium antillarum is a common

species in the Caribbean and tropical Western At-

lantic, where hving specimens have been found at

depths ranging from 3 m. to 70 m. and where, in

shallow waters, they are commonly attached among

Halimeda algae. The previously known geographic

range of the species extends from off Colon, Pan-

ama and Barbados northward to Great Abaco, Ba-

hamas. Pecten chazaliei is known to live at depths

ranging from 18 to 230 m. in an area ranging from

off Brazil, through the Caribbean and eastern Gulf

of Mexico, and up the North American coast as far

north as Cape Fear, North Carolina (depth, 73 m.).

Aequipecten acanthodes (Dall, 1925) was orig-

inally described as being "less inflated, wider, and

with less prominent spines" than Aequipecten

muscosus (Wood, 1828) [= Pecten exasperatus

Sowerby, 1843]. Althougli the type locality of

Dall's species was never given, the syntypes (USNM

333367) are from "Santa Lucia, northwest Cuba,

depth 2 to 4 fathoms". Tlie Bermuda specimens

listed in Table 1 bear a closer resemblance to the

Cuban forms than to A muscosus from the Florida

coast. The larger of the two syntypes of A.

acanthodes is illustrated in Figures 3 and 4.

The occurrence of Argopecten gibbus within a

neady land-locked sound is unusual. Along the

North American coast, the species is found only in

open marine waters generally within a depth range

of 9 to 365 meters (Waller, 1969, p. 37), whereas

bays and sounds harbor populations of Argopecten

irradians and Argopecten nucleus.

As observed by Neuman (1965, pp. 999, 987),

Harrington Sound "does not exhibit extremes in

either dilution or evaporation", and regardless of

the almost complete isolation of the sound, "tidal

exchange througli the narrow inlet is sufficient to

maintain near-normal oceanic temperature and sa-

linity." In addition there is some exchange of water

througli caves and the porous limestone surrounding

the sound. As emphasized by De Laubenfels (1950.

46 THE NAUTILUS

AprU 1973

Vol. 87 (2)

p. 1 59), the small amount of land area surrounding

Harrington Sound in comparison to the large size of

the sound itself, coupled with the porous bedrock,

permits httle land runoff and allows for greater

constancy of oceanic conditions. In contrast, sounds

located along the North American coast commonly

exhibit extremes of both salinity and temperature,

favoring species adapted to variation of these con-

ditions.

In Harrington Sound, Argopecten gibbus rests on

the bottom with its right valve on or sliglitly within

the sediment. Tlie majority of the largest individuals

lie free, without byssal attachment, although many

mature individuals attach a weak byssus to dead

sheOs and coral fragments. The species is sluggish

and rarely begins to swim when approached by a

diver or when handled. Wlien swimming begins, it is

an irregular motion with rapid adductions of the

valves and short flights of no more than a meter.

Like all free-living scallops, A. gibbus has an over-

turning reaction when placed with its left valve

downward. Generally the animal rests motionless for

a time varying from a half minute to three minutes.

Then the valves gape broadly, the vela become rigid,

and with one or two claps the scallop rotates on

either its liinge axis (10 out of 18 trials) or on an

axis perpendicular to the liinge and parallel to the

plane of commissure. Generally the smaller in-

dividuals (those of about 25 or 30 mm in height in

our test sample) begin the righting reaction sooner

than larger individuals (those 50 to 60 mm in

height).

Two valves and numerous fragments of

Argopecten gibbus were found in the Sub-

thermochne Zone near Shark Hole, Harrington

Sound (Station 6, Fig. 2). Tlie larger of the two

nearly complete valves is a typical representative of

the species, with a thick shell and moderate con-

vexity. It is indistinguishable from individuals living

in the Shallow Sandy Zone, and because its auricles

are chipped as thougli bitten, the shell may have

been dropped or dragged into this zone by a

predator.

Tlie smaller of the two valves is strikingly dif-

ferent from any of the specimens found elsewhere

around the Bermuda Islands. Its shell is exceedingly

thin with very low curvature and radial plicae of

low amplitude. Its disk flanks are very low and, on

the posterior side, there is no sharp demarcation

between the disk flank and posterior auricle, which

lacks radial costae. Dark, brownish red pigment

produces mottling on the valve exterior and a more

uniform dark hue on the interior of the valve in

areas outside of the pallial Une. Sculptural details,

such as the fine, concentric lamellae on the exterior

and ridged cardinal crura on the interior, and the

aragonitic myostraca are well-enough preserved to

suggest that the thinness of the shell is not entirely

due to solution. Most of the fragments recovered

exhibit the same thinness and pigmentation as the

smaller of the two whole valves, and according to

G. R. Clark, II (personal communication) such

specimens are common from drill cores of sediments

within the Subthermocline Zone. The thin shells are

within the range of variation exliibited by

Argopecten gibbus and most closely resemble fossil

specimens from early Pleistocene deposits in

southern Florida (see Waller, 1969, pp. 65, 66).

The thin specimens may be the offspring of

shallow-water populations which have entered the

Subthermocline Zone as larvae during maximum

overturn of the water. Subsequent grov^^h may then

have been modified by the unusual conditions pre-

sent in the subthermocline environment. Al-

ternatively, the subthermocline Argopecten may in

fact be fossils introduced into Harrington Sound

some thousands of years ago at a time when

populations throughout the Sound were of this

morphology. These hypotheses can be tested, the

former through culturing of Argopecten gibbus

within the Subthermocline Zone, the latter through

radiocarbon dating of shells and observation of

Bermuda fossils both from within and outside of

Harrington Sound.

Argopecten noronhensis (E. A. Smith, 1885) was

originally described from the coast of Brazil, where

it was dredged by the H. M. S. Challenger from a

depth of 46 m off Fernando Noronlia. Since

Smith's original description, there have apparently

been no further reports on the occurrence of the

species outside of Brazilian waters. However, col-

lections in the U. S. National Museum contain

numerous specimens from Barbados (depths, 46 to

1 83 m ), Antigua (220 m ), the Virgin Islands and

Puerto Rico (73 to 256 m ), and Cat Cay, Baliamas

(550 m ). All of these specimens were found dead

with only the shells remaining on bottoms of coarse

sand or coral rubble. Living specimens have been

taken by the R/V Oregon off St. Lucia (66 to 75

m), south of Saba Banks (91 to 110 m), and off

Vol. 87 (2)

THE NAUTILUS

47

Tortola. Virgin Islands (49 m). Argopecten

noronlicnsis is not known to occur in the western

Caribbean or in the Gulf of Mexico. In Bermuda

fragments of A. noronhemis were recovered from

sediment samples taken on our deepest dive, at a

depth of 51 m (Station 39). The species is il-

lustrated in Figures 5-8.

Althougli not well-known because of its small

size (generally less than 1 5 mm), Chlamys benedicti

VerrUl and Bush, 1897, is the most widespread and

numerous Chlamys in the Western Atlantic region.

According to data with specimens examined thus

far, the species ranges from the coast of South

America off Surinam northward to a point south-

east of George's Bank and is also widespread in the

northern and eastern Gulf of Mexico. Shells of C.

benedicti have been found from beach level down

to 2,480 in, but living specimens are known only

from about 2 to 806 m. Out of 86 records of

living specimens, 75% are within the 46 to 90 m

range. At many localities the species is associated

with coral debris and sponges, and preserved spec-

imens indicate that, like all Chlamys, C. benedicti is

byssally attached. At Bermuda, fragments and one

small valve of Chlamys benedicti were recovered

from sediment samples taken at Stations 24 (34 m)

and 39 (51 m).

Despite an intensive search for living Chlamys

imbricata, only one live individual only 17 mm in

height was found (Station 18). It was byssally

attached with its riglit valve upward against the

undersurface of a hemispherical head of dead coral

at a depth of 7 m. The scallop voluntarily released

its byssal attachment when the coral was overturned

and, by clapping its valves, slid along the coral

surface.

In a bucket on the boat, the specimen quickly

attached itself with a new byssus to a clump of

coral. However, after a few hours in a laboratory

aquarium, the specimen detached itself from the

same clump of coral, moved up and out of a small

glass bowl in the bottom of the aquarium, and

reattached to the lower umbo of an Argopecten

gibbus on the bottom of the same aquarium. On the

basis of only a few trials with this one immature

specimen, C. imbricata seems to have only a weak

overturning reaction, generally not beginning to

right itself for several minutes after being over-

turned and then rigliting with a single clap and

rotation about the hinge line. Tlie specimen was

observed to swim with its commissure inclined and

left valve uppermost, taking off from the bottom at

a steep angle with very rapid valve adductions.

Bermuda is the northernmost occurrence of this

species, which on the North American coast occurs

no further north than Miaini. In the south, C

imbricata is known only from the Caribbean,where

it has been found as far south as Payardi Island,

Panama.

Chlamys multisquamata (Dunker, 1864) is the

rarest of the Western Atlantic Chlamys. Originally

described from Havana Bay, Cuba, the species was

subsequently redescribed as Pecten effluens by DaU

(1886) on the basis of two immature valves dredged

from 230 m. off Havana. Since this time, additional

specimens have been found at Barbados (USNM, 46

to 110 ni); Dominica (Nat. Marine Fish. Service,

Lab. at Pascagoula, Miss., 62 to 110 m); Puerto

Rico (USNM. 73 to 91 m); Cuba (USNM, 550 m);

Bahamas (USNM, 90 to 550 m): off Palm Beach,

Florida (F. M. Bayer Collection, 24 to 30 m); and

Bonaine, Dutch West Indies (S. D. Abbott ,17 m.

in crevice, on coral heads). At Bermuda, un-

mistakable fragments of the species were recovered

from sediment samples taken on the deepest dive

(Station 29, 51 m). According to data accom-

panying the Palm Beach specimen, the species was

living among rocks and between the shells of

Spondylus. Morphologically, Chlamys multisquamata

resembles several Indo-Pacific species that live in

coral crevices (see Waller, 1972). The Western At-

lantic species is illustrated in Figures 9-11.

Living Pecten ziczac was found only in the

Shallow Sandy Zone of Harrington Sound, althougli

shells of the species are common in other bays such

as Ely's Harbor and inner Port Royal Bay. The

living habits of F. ziczac are as described by Stanley

(1970, p. 140). The scallops live with their riglit

valve buried in the sediment to the level of the

commissure; a thin layer of sediment covers the

upper flat valve; and the only portion visible from

above is the ring of extended tentacles.

Pecten ziczac is more adept at swimming and

does so more readily than Argopecten gibbus. Al-

thougli the latter would not swim when sensing the

approach of a diver, Pecten would occasionally do

so. Other specimens would begin swimming when

lifted from the sediment. The animal takes off at a

steep angle and then levels out in flights of about 5

seconds duration or longer. Adduction frequency is

48 THE NAUTILUS

April 1973

Vol. 87 (2)

fairly rapid, generally about two claps per second,

and flight distances average slightly more than 1 in.

Like Argopecten, P. ziczac has a distinct rigliting

reaction but because of the flatness of its left valve,

righting is accomplished with great difficulty.

Studies of three specimens in an aquarium indicated

that rigliting begins one or two minutes after over-

turning. Rotation is about an axis parallel to the

hinge line, generally requiring three or four claps of

the valves during rotation.

Tlie ability of Pecten ziczac and many other

scallops to excavate a shaOow depression in the

sediment has been described by Waller (1969, p. 17)

and Stanley (1970, P- 140).

Anomiidae

Collections of Anomia simplex and Pododesmus

nulls from Harrington Sound suggest differences in

habitat between the two species. Anomia is most

abundant in the Shallow Sandy Zone where it

attaches to shells and other debris on the sandy

substrate. Pododesmus, on the otlier hand, is most

abundant in the Oculina Zone, where it attaches to

Ocullna and avoids contact with the bottom. In

areas on the North American coast, Stanley (1970,

p. 144) has observed that A. simplex generally

occurs attached to pebbles and debris on a firm

substrate where there is a moderate current flow.

Limidae

Lima lima attaches with a strong byssus beneath

rocks in open marine areas and avoids bays and

sounds.

Gryphaeidae and Ostreidae

The listing of Neopycnodonte cochlear is based

on a single, eroded riglit valve from the deepest

station (Station 39, 51 n^; Figs. 12, 13). The

specimen is roughly circular in form, measures 13

mm in diameter, and clearly displays the vesicular

shell structure and outer prismatic layer (Fig. 12)

that Stenzel (1971, p. 1105) considers to be char-

acteristic of the genus. According to Stenzel there is

only one living species in the genus, which has a

worldwide distribution in tropical and subtropical

oceanic waters ranging in depth from 27 to 1,500

m.

The familiar Ostrea frons of the West Indian

region has been relegated to the genus Lopha and to

the synonymy of Ostrea folium Linnaeus by Stenzel

(1971, pp. 1024, 1158). Tlie elongate, clasping form

associated with Ostrea frons is one of two

ecomorphic end members within the species. Al-

thougli commonly attributed to growth while at-

tached to mangrove roots (Warmke and Abbott,

1954, p. 173), this ecomorph is generally the result

of attachment to gorgonacean corals in subtidal

habitats. In Harrington Sound (Station 13), the

species also cements itself to shells lying on the

bottom, in which case it secretes clasps only where

the oyster shell overlaps the edge of the shell to

which it is attached. As with all oysters, the shape

of the attachment surface profoundly affects the

shape of the oyster, and those attached to shells are

less crenulated and less elongate than those found

on gorgonaceans.

Condylocardiidae

Viviparity is conmion in the superfamily

Carditacea (Dall, 1902, p. 696), and Cardltopsls

smlthl is no exception. One of the dead but tiglitly

closed specimens from Walsingham Pond yielded

three articulated prodissoconchs, one of which is

illustrated in Figure 19. Large, sculptured,

saucer-shaped prodissoconchs, of this type are gen-

erally associated with viviparity in several taxonomic

groups, as in the genera Condylocardla (Bernard,

1 897, p. 1 70) and Philobrya (Bernard, 1 896, p. 9).

Chamidae

Certain species of Bermudian chamids are sepa-

rated in open marine and partially enclosed en-

vironments. Pseudochama radians lives in abundance

in the Shallow Sandy Zone of Harrington Sound

and also occurs in the Ocullna bed at Station 27 in

association with well-developed Chama

maccrophylla. Another common chamid in Har-

rington Sound is Chama congregata, which is most

abundant in the Ocullna Zone. Chama sarda was

found only well off-shore, where it was abundant at

the three stations deeper than 30 m.

The shells of Chama maccrophylla found on the

bottom of Walsingliam Pond have extremely heavy

and deep left valves and resemble the Chama ber-

mudensls described by Heilprin (1889, p. 141) from

Harrington Sound. Because these have crenulated

inner margins and lack an exterior sulcus, they

cannot be considered a subspecies of Chama slnuosa

Broderip as in Bayer (1943, p. 122).

Vol. 87 (2)

THE NAUTILUS

49

In Table 1 the Chainidae have been placed near

the Carditidae rather than near the Lucinidae fol-

lowing the convincing evidence of relationships pre-

sented by Kennedy, Morris, and Taylor (1970).

Crassatellidae

FoUovk'iiig the study of Harry (1966), both

Crassinella parva (C. B. Adams, 1845) and C. guad-

ahipensis (d'Orbigny, 1846) are considered to be

junior synonyms of C. Iwndata (Conrad, 1 834).

Veneridae

The Western Atlantic species of Gouldia are

higlily variable and inadequately described. It is

curious that G. ceriiia (C. B. Adams, 1845), G.

bermudemis (E. A. Smith, 1885), G. imularis (Dall

and Simpson, 1901), and G. foresti Fischer-Piette

and Testud, 1967, were all described without ade-

quate published comparison to other known species.

Tliere is no adequate basis for separating G. cerina,

G. benmidensis, and G. foresti without an extensive

quantitative study of geographical variation; Gouldia

insularis appears somewhat more distinct, character-

ized by its rounded form and high degree of

inflation.

None of the numerous Gouldia from the Sub-

thermocline Zone of Harrington Sound were alive

when collected. All show some deterioration of

their aragonitic shells and lack preserved ligaments,

although some have traces of pigment remaining

(see Notes on Habitats).

Gastropoda

Rissoidae

Alvania platycephala was first described and il-

lustrated by Dautzenberg and Fischer (1896, p. 63,

plate 19, figs. 12, 13) on the basis of specimens

taken from a depth of 1,385 meters in the Azores.

The first Bermudian record is that of Verrill and

Bush (1900, p. 539), who identified the species

from "shell sand" taken from waters less than 15

m deep. Tlieir identification was apparently based

on Dautzenberg and Fischer's illustration, and the

specimens themselves were evidently not compared.

The species was subsequently reported by Piele

(1926, p. 77) from an unknown site and by Gould

(1968, p. 4) from Walsingliam Pond. These spec-

imens, shown here in Figure 21, bear an even closer

resemblance to the illustration of Dautzenberg and

Fischer than the rather poor line drawing of Verrill

and Bush would indicate. Tlie species was found by.

us only in Walsingliam Pond (Station 1 1 ), where it

is abundant.

A second species of Alvania, also reported by

Piele (1926, p. 77), is A. pagodula (Bucquoy,

Dautzenberg, and Dollfus, 1884), originally des-

cribed from the Mediterranean and Adriatic. Piele's

specimens were more likely Alvania didyma (Wat-

son, 1886), which in the present study was taken at

depths exceeding 30 m. (Stations 21 and 24, Fig.

20).

Turritellidae

Gould (1968, p. 4) recognized Vermicularia

spirata Philippi, 1836, as the most common moUusk

in Walsingham Pond and later (Gould, 1969) studied

the ecology and functional significance of uncoiling

in the species. In Bermudian waters V. spirata

cements itself to a variety of hard substrates but is

particularly common in association with the branch-

ing coral, Oculina. In Walsingliam Pond there is no

development of Oculina, and, as noted by Gould,

fewer than 1 percent of the largest Vermicularia

present show uncoiling. Instead, individuals live in

burrows with their apertures projecting sliglitly

above the mud.

Specimens of Vermicularia from stations on the

open Bermuda Platform have all been identified

here with V. knorri (Deshayes, 1843) only because

their early whorls are white, rather than tan or

brown as in V. spirata. This is a character that some

workers believe is genetically significant. However,

there are no differences in whorl profile or sculp-

ture other than those imposed by differing rates of

uncoiling, a process shown by Gould to be en-

vironmentally controlled, and it is possible that

further study will place V. knorri in the synonymy

of K spirata.

Caecidae

Caecum plicatum Carpenter, 1858, is a senior

synonym of Caecum termes Heilprin, 1 889, re-

ported by Piele (1926, p. 78) to be the most

common Caecum in Bermuda and by Gould (1968,

p. 4) to be the most common in Walsingham Pond.

Buccinidae

All living Colubraria swifti observed in Bermuda

(Table 1) were beneath rocks in open marine areas,

commonly associated with Barbatia domingensis.

50 THE NAUTILUS

ApiU 1973

Vol. 87 (2)

TABLE 2. Collecting stations grouped according to general environment or ecological zone. Ecological zones in

Harrington Sound follow Neunwnn ( 1 965).

Description

Harrington Sound. Shallow Sandy Zone

Harrington Sound and North Lagoon,

Oculina Zone

Harrington Sound, Subthermocline Zone

Walsingham Pond, muddy bottom

Bays and sounds other than Harrington

Sound

Open platform, sand pockets and

patch reefs, 5 to 18 m

Open platform, sand pockets, 34 to 51 m

Stations

3. 5, 9, 10, 13

2, 7, 27

1, 6

11

14, 17, 20, 29, 30, 31

33, 34b, 35, 36, 37, 38

12, 15, 16, 18, 23,25,

26, 28

21, 24, 39

Polyplacophora

Isclmochitonidae

Radsiella nigulata (Sowerby, 1832) is a senior

synonym of Ischnochiton boogii Haddon, 1886

(Keen, 1971, p. 869), which in turn is a senior

synonym of Ischnochiton bermudensis Dall and

Bartsch, 1911.

ACKNOWLEDGMENTS

Tlie field portion of this study was carried out at

the Bermuda Biological Station for Research, St.

George's West. I am grateful to the staff of the

Station, particularly Dr. Wolfgang Starrer, Director,

and Mr. Brunell Spurling, skipper of the Micmac.

Dr. Porter M. Kier, Chairman of the Department of

Paleobiology, Smithsonian Institution, kindly per-

mitted his assistant, Mr. Tliomas F. Phelan, to

accompany me as a companion diver. Both Mr.

Phelan and Miss Ellen P. Broudy, my research

assistant, ably assisted in the field study. Additional

help in Bermuda was received from Dr. J. D.

Herkes, Dr. David Pawson, and Mr. Arthur Guest.

Assistance in identification was received from Mr.

Russell Jensen and Dr. R. T. Abbott, Delaware

Natural History Museum; Dr. D. R. Moore, Uni-

versity of Miami; Dr. K. J. Boss, Harvard Uni-

versity; Dr. Ruth Todd, U. S. Geological Survey,

and Drs. J. P. E. Morrison, H. A. Rehder, J.

Rosewater, and G. A. Cooper, Smithsonian In-

stitution, and Mr. W. G. Lyons, Florida Department

of Natural Resources, St. Petersburg. I am grateful

to Dr. F. M. Bayer, University of Miami, for the

loan of a specimen of Chlamys nndtisquamata, and

to Mr. H. R. Bullis, Jr., National Marine Fisheries

Service, for permission to study his collections in

Pascagoula, Mississippi. Dr. D. R. Moore, Dr. R. T.

Abbott, and Mr. R. Jensen read the manuscript and

gave many helpful suggestions. Mr. Larry Isham,

artist for the Department of Paleobiology, drafted

the maps.

Funding for travel and field study was through

the Department of Paleobiology, Smithsonian Insti-

tution, and funding for all other aspects of the

work was througli grants to the author from the

Smithsonian Research Foundation.

LITERATURE CITED

Abbott, R. Tucker. 1954. American seashells:

Princeton, Van Nostrand Co., 541 p., 40 pis.,

100 figs.

Adams, C. B. 1845. Synopsis Conchyhorum

Jamaicensium, etc. Proc. Boston Soc. Nat. Hist.,

Jan. 1, 1845, 17 p.

Bayer, F. M. 1943. Tlie Florida species of the

family Chamidae. Tlie NautUus, 56 (4): 116-124,

pis. 12-15.

Bernard, Felix. 1896. Etudes comparatives sur la

coquille des lamellibranches. Condylocardia, Jour.

ConchyHologie. 44: 169-207, 5 figs., pi. 6.

Bernard, F€lix. 1897. Etudes comparatives sur la

coquille des lamellibranches. II. Les genres

Philobrya et Hochstetteria: Ibid. 45: 5-47, 7

figs., pi. 1.

Boss, Kenneth J. 1966. The subfamily Tellininae in

the Western Atlantic. The genus Tellina (Part I).

Johnsonia 4 (45): 217-272, pis. 127-142, 3

tables.

Boss, Kenneth J. 1968. The subfamily Tellininae in

the Western Atlantic. The genera Tellina (Part II)

and Tellidora. Ibid. 4 (46): 273-344, pis.

143-163.

Vol. 87 (2)

THE NAUTILUS

51

Boss, Kenneth J. 1969. The subfamily Tellininae in

the Western Atlantic. The genus Strigilla. Ibid. 4

(47): 345-368, pis. 164-171.

Boss, Kennetli J. and Donald R. Moore. 1967.

Notes on Malleus (Parimalleusj candeanus

(d'Orbigny) (Mollusca; Bivalvia). BuU. Mar. Sci.

17 (1): 85-94, 2 figs.

Bretsky, Sara S. 1967. Environmental factors in-

fluencing the distribution of Barbatia do-

mingensis (Mollusca: Bivalvia) on the Bermuda

Platform. Postilla, Peabody Museum of Nat.

Hist., Yale Univ., no. 108, 14 p., 2 figs., 2 tables.

Britton, Joseph Cecil. 1970. The Lucinidae (Mol-

lusca: Bivalvia) of the western Atlantic Ocean.

Washington, D. C, George Washington Univ.,

unpublished Ph.D. dissertation, 566 p., 23 pis., 7

tables.

Qench, William J. 1942. The genus Conns in the

Western Atlantic. Johnsonia 1 (6): 40 p., 15 pis.

Clench, William J. 1964. The genera Pedipes and

Laemondonta in the Western Atlantic. Ibid., vol.

4, no. 42, p. 117-127, pis. 76-79.

Clench, William J. and Ruth D. Turner. 1950. The

Western Atlantic marine mollusks described by C.

B. Adams. Occasional Papers on Mollusks, Mus.

of Comparative Zool., Harvard Univ. 1 (15):

233-403, pis. 28-49.

Dall, William Healey. 1886. Reports on the results

of dredging ... by the U. S. Coast Survey

Steamer "Blake". Report on the Mollusca, Part I

- Bracliiopoda and Pelecypoda. BuU. Mus. Com-

parative Zool. 12 (6): 173-318, 9 pis.

Dall, William Healey. 1 889. A preliminary catalogue

of the shell-bearing marine ■ mollusks and

brachiopods of the southeastern coast of the

United States, with illustrations of many of the

species. Bull. U. S. Natl. Mus., no. 37, 232 p., 95

pis.

Dall, William Healey. 1902. Synopsis of the

Carditacea and of the American species: Proc.

Acad. Natural Sci. Pliila. 54: 696-716.

Dall, William Healey. 1925. Notes on the nomen-

clature of some of our east American species of

Pecten with descriptions of new species. Nautilus,

vol. 38, no. 4, p. 112-120.

Dall, William Healey and Paul Bartsch. 1911. New

species of shells from Bermuda. Proc. U. S. Natl.

Mus. 40 (1820): 277-288, pi. 35.

Dall, William Healey and Charles Torrey Simpson.

1901. The Mollusca of Puerto Rico. U. S. Fish

Commission Bull. 1: 351-524, pis. 53-58.

Dautzenberg, P., and H. Fischer, 1896. Compagnes

scientifiques de S. A. le Prince Albert 1 de

Monaco. Dragages effectives par VHirondelle et

par \a Princess- Alice , 1885-1895. Mem. de la Soc.

Zool. de France 9: 1-104, pis. 15-22.

Davis, J. D., 1967. Ervilia concentrica and Meso-

desma concentrica. Clarification of synonymy.

Malacologia 6 (1-2): 231-241, 1 fig., 2 pis.

De Laubenfels, M. W. 1950. An ecological dis-

cussion of the sponges of Bermuda. Trans. Zool.

Soc. London 27: 155-201, 4 figs.

Fischer-Piette, E. and A.-M. Testud. 1967.

MoUusques Lamellibranches: Veneridae. Resultats

scientifiques de campagnes de la "Calypso", Fas-

cicule 8, no. 13, p. 205-220, 4 pis.

Gould, Stephen Jay. 1966. Notes on shell mor-

phology and classification of the Siliquariidae

(Gastropoda). The protoconch and slit of

Siliquaria squamata Blainville: Amer. Mus.

Novitates, no. 2263, 13 p., 7 figs., 2 tables.

Gould, Stephen Jay. 1968. The moUuscan fauna of

an unusual Bermudianpond: A natural experiment

in form and composition: Breviora, no. 308, 13

p., 2 figs., 2 tables.

Gould, Stephen Jay. 1969. Ecology and functional

significance of uncoiling in Vermicularia spirata:

An essay on gastropod form. Bull. Marine Sci. 19

(2): 432-445, 3 figs., 1 table.

Harry, Harold W. 1966. Studies on bivalve molluscs

of the genus Crassinella in the Northwestern Gulf

of Mexico: Anatomy, Ecology and Systematics.

Publ. Inst. Marine Sci., Texas 11: 65-89, 17 figs.

Heilprin, Angelo. 1889. On some new species of

Mollusca from the Bermuda Islands. Proc. Acad.

Nat. Sci. Philadelphia for 1889, p. 141, 142, pi.

8.

Jackson, Jeremy B. C. 1970. The ecology and

paleoecology of near shore tropical Lucinacea

(Bivalvia). Geol. Soc. America Abstracts, 2 (7):

586-587.

Keen, Angeline Myra. 1971. Sea shells of tropical

west America, 2nd Edition. Stanford, California,

Stanford Univ. Press., 1,064 p., 22 pis.

Kennedy, William James, N. J. Morris and John D,

Taylor. 1970. The shell structure, mineralogy and

relationships of the Chamacea (Bivalvia).

Palaeontology 13, part 3, p. 379-413, 6 figs.,

pis. 70 - 77.

McGinty, Thomas L. 1940. New land and marine

Tertiary shells from southern Florida. The Nauti-

lus 53 (3): 81-84, pi. 10.

52 THE NAUTILUS

April 1973

Vol. 87 (2)

Moore, Raymond C. (Editor). 1969. Treatise on

invertebrate paleontology, Part N, vols. 1 and 2,

moUusca 6, bivalvia: Univ. of Kansas and Geol.

Soc. Amer., 951 p., illustr.

Neumann, A. Conrad. 1965. Processes of recent

carbonate sedimentation in Harrington Sound,

Bermuda. Bull. Marine Sci. 15: 987-1035, 8 figs.,

1 table.

Piele, A. J. 1926. The MoUusca of Bermuda. Proc.

Malac. Soc. London 17 (2 and 3): 71-98.

Smith, Edgar A. 1885. Report on the

LameDibranchiata. Report on the scientific re-

sults of tlie voyage of the H. M. S. Challenger

during the years 1873-76, Zoology 13 (35); 341

p., 25 pis.

Stanley, Steven M. 1970. Relation of shell form to

life habits in the Bivalvia (MoUusca). Geol. Soc.

Amer. Mem. 125. xiii -i- 296 p., 48 figs., 40 pis.

Stenzel, H. B. 1971. Oysters: in Raymond C. Moore

(Editor), Treatise on invertebrate paleontology,

Part N, vol. 3, MoUusca 6, Bivalvia, p. 953-1224,

1 53 figs.

Taylor, Dwiglit W. and Norman F. Sohl. 1962. An

outline of gastropod classification. Malacologia 1

(1): 7-32, 2 figs.

Verrill, A. E. and Katherine J. Bush. 1900. Addi-

tions to the marine MoUusca of the Bermudas.

Trans. Connecticut Acad. Sci. 10; 513-544, pis.

63-65.

Waller, Tliomas Richard. 1969. Tlie evolution of the

Argopecten gibbus stock (MoUusca: Bivalvia),

with emphasis on the Tertiary and Quaternary

species of eastern North America. Paleont. Soc.

Mem. 3 (Jour. Paleont. 43 (5), suppl.), 125 p.,

13 figs., 7 pis., 43 tables.

WaUer, Thomas Richard. 1972. The Pectinidae (Mol-

lusca: Bivalvia) of Eniwetok Atoll, Marshall Is-

lands. Veliger 14(3); 221-264, 22 figs., 8 pis., 13

tables.

Walton, Anthony W. 1969. Walsingham Pond: A brief

study of its sediments and water chemistry. Re-

ports of Research, 1968 Seminar on organism-sedi-

ment interrelationships, Bermuda Biol. Sta. for Re-

search, Special Publ. no. 2: 41-50, 3 figs.

Warmke, Germaine L. and R. Tucker Abbott. 1961.

Caribbean seashells. Narberth, Pennsylvania, Liv-

ingston Publ. Co., 346 p., 44 pis., 34 text-figs.

Warmke, Germaine L. and Luis R. Almodovar.

1963. Some associations of marine moUusks and

algae in Puerto Rico: Malacologia 1 (2): 163-177,

1 fig., 10 tables.

Yonge, C. M. 1968. Form and habit in species of

Malleus (including the "hammer oysters") with

comparative observations on Isognomon isogno-

mon Biol. BuH. 135 (2): 378-405, 13 figs.. 1

table.

BOOK REVIEW

SHELLS AND SHELL COLLECTING. By S. Peter

Dance. 128 pp., 110 photos, 30 in color. Index.

HamlynPubl, London, N. Y., Toronto. $4.95.

This is one of the more attractive of the recent

rash of popular books on conchology. It is

well-written, accurate and Ulustrated by excellent

photographs. It is an interesting introduction to the

major groups of moHusks. shell shapes and colors,

the history of shell collecting, rare shells, and man's

use of moUusks as food, money and objects of art.

Peter Dance has included several accounts new to

the popular literature, and has illustrated a number

of land and marine shells that previously had not

been published in books for the layman. Missing

from tills otherwise excellent book on sheU col-

lecting are instructions on how to coUect shells and

how to clean and preserve them. The art director

has reversed the negative on the first double-page

spread, thus creating sinistral Nucella dogwinkles.

For its low price, this book is a worthwhile pur-

chase.

R. Tucker Abbott

Delaware Museum of Natural History

I

Vol. 87 (2)

THE NAUTILUS

53

EVIDENCE FOR SPAWNING BY GONATUS SP. (CEPHALOPODA: TEUTHOIDEA)

IN THE HIGH ARCTIC OCEAN

Richard Edward Young

Department of Oceanography

University of Hawaii, Honolulu, Hawaii 96822

ABSTRACT

A specimen of the squid, Gonatus sp., was captured through an ice hole at 79° 58'

N. lat., 170° 23' E. long, (about 500 miles north of Wrangel Island, Siberia) whose

condition suggests it had spawned just before its capture on March 30, 1962. The

emaciated condition is described in specimens of other Califomian female Gonatus that

presumably had spawned just before capture, and a brief review is given of the records

of the degeneration and fatal effects from spawning by females in other genera of

squid and octopods.

The cephalopod fauna of the high Arctic Ocean

is poorly known; pelagic cephalopods have been

reported only on a few occasions. Berry (1925)

described from seal stomachs a small collection of

cephalopod beaks, one of which taken at

70°13'N, 140°50'W, probably belongs to a squid.

MacGinity (1955) recorded a juvenile and adult

Cirroteuthis sp. (finned octopods) that were

dip-netted near shore and 3 specimens of Gonatus

"fabricii" (72, 63, 59 mm mantle length) that had

washed ashore at Pt. Barrow, Alaska. Voss (1967),

in a footnote, mentioned the capture of Cir-

rothauma (finned octopod) in the high Arctic. The

locality of this capture is 86°N, 173°E (Roper and

Brundage, 1972). Nesis (1971a) found 2 specimens

of Gonatus "fabricii" (175 and 87 mm Mantle

Length) in an ice hole at §7°24.2'N, 132°01.5'E

and one specimen of G. "fabricii" (130 mm ML) in

an ice hole at 80°13.3'N and 143°01'E. He further

reports that Gonatus is abundant at the border of

the Arctic Basin and the Greenland Sea, particularly

young specimens ranging from 30-71 mm ML. The

only pelagic cephalopods known from the high

Arctic Ocean, therefore, are members of the squid

family Gonatidae and two species of finned oc-

topods.

The squid reported here was captured from ice

island Arlis II (Arctic Research Laboratory Ice

Station No. 2) in the high Arctic Ocean on March

30, 1962. The squid, alive at the time of capture,

was found floating head downward in a hole cut

through the ice for hydrographic and plankton

studies.

DESCRIPTION

Since the identity of this specimen is of con-

siderable importance but impossible to determine to

species at present due to the condition of the

animal, a brief description is presented (Fig. 1 & 2).

The pen is 210 mm in length. The manfle and fins

are gelatinous and flaccid. A well-developed conical

"tail" is present posterior to the conus of the pen.

In preservation in 70% ethanol the fins measure 102

mm in total length and 145 mm in total width. The

fins extend posteriorly along the "tail."

The funnel is large and reaches to the level of

the midpoint of the eyes. The funnel locking-carti-

lages bear simple straight grooves (Fig. 2, F). The

dorsal pad of the funnel organ has an inverted

V-shape, the anterior half of each limb has low

ridges along the lateral margin (Fig. 2, G). A small

anterior papilla is present. The ventral pads are

smaU and nearly teardrop-shaped with the blunt end

anterior. A large funnel valve is present.

The head is short and bears large eyes with a

distinct sinus on the anterior margin of each eyelid.

A nuchal crest with three indistinct pairs of nuchal

folds occurs on the head. "Olfactory" lobes lie on

the second pair from the funnel on each side. The

nuchal cartilage, which is long and sliglitly rounded

at either end, bears a median ridge containing a

central groove.

The arms are gelatinous and relatively short (Arm

I = 73 mm. III - 97 mm, IV = 98 mm). A weakly

developed aboral keel is present on each arm III and

large lateral keels are present on each arm IV. All

arms bear very broad, thick, gelatinous trabeculae

that are joined by thick, poorly defined protective

membranes which converge and conceal the hooks

in the distal portion of the arms. The arm tips are

not attenuate. The armature of arms I-III consists of

two alternating rows of small hooks (Arm I = 42

54 THE NAUTILUS

April 1973

Vol. 87 (2)

hooks, II = 41 hooks, III = 44 hooks). The marginal

and terminal rows of suckers are absent and only

small grooves indicate their former presence. Suckers

are absent from arms IV (Fig. 2, B). The tenta -

cles have been lost except for rounded stubs.

The specimen is a female. Oviducal and

nidamental glands are slightly swollen (lengths: 28

mm and 14 mm respectively). The ovary is rather

small but still contains many elongate immature ova

and a few scattered, larger sperical ova of various

sizes, the largest of which measures about 1.5 mm

in diameter.

The radula (Fig. 2, C) contains only five teeth in

a transverse series. The rachidian tooth has a short

central cusp and two small lateral cusps. The first

lateral teeth are absent. The second laterals have

broad bases each with a rather blunt medial cusp.

Marginal plates are absent. The beaks (Fig. 2, D) are

heavily pigmented. The lower mandible shows a

faint ridge on the lamella. Many sperm reservoirs are

attached to the inner wall of the buccal membrane.

Although the generic boundaries in the family

Gonatidae appear to be somewhat indistinct

(Okiyama, 1969; Fields & Gauley, 1971; Nesis,

1971b), this specimen clearly belongs within the

genus Gonatus sensu stricto as indicated by the

presence of a tail extending beyond the gladius,

tentacle stubs and a radula with only five teeth.

DISCUSSION

The following evidence suggests that the spec-

imen has recently spawned:

1. The specimen had mated as indicated by the

presence of sperm reservoirs (discharged

spermatophores) embedded in the buccal

membrane.

2. The nidamental and oviducal glands are of

intermediate size (i. e. these glands are larger

than would be expected in immature specimen

but much smaller than would be expected in a

gravid specimen.

3. The ovary is almost totally depleted of mature

ova.

4. The specimen has undergone degeneration as

indicated by the gelatinous nature of the

muscular tissues and the loss of suckers from

all arms, as well as the loss of the tentacles.

The features which typify spent females are not

well-known. In order to confirm the above list as

characteristic of spent females of the family

Gonatidae, I have examined the extensive col-

lections of cephalopods at the University of South-

em California for females that appear to have

recently spawned. Four species of Gonatus are

found in the waters off southern California; females

of three species were found which appear to have

spawned.

Gonatus pyros Young, 1972. Two specimens,

135 mm pen length (P. L.) and 130 mm pen length,

easUy identified by the presence of a large oval

photophore on the ventral surface of each eye,

showed extreme signs of degeneration. In con-

sistency they are flaccid and gelatinous which con-

trast strongly wdth the muscular condition of im-

mature specimens. Tentacles are absent except for

small stubs. All arm suckers are absent; small poc-

kets or shght puckerings of the skin on the gela-

tinous trabeculae mark the spots where the suckers

had once been. The nidamental glands in both

specimens are considerably larger ( 30 and 25

mm) than one would expect to find in a large

immature specimen. The oviducal glands are fairly

large (22 and 16 mm). In both cases the ovary is

small and filled with small, deteriorated eggs. In one

specimen a mature egg which was presumably dis-

lodged from the oviducal gland during dissection

was found in the mantle cavity. No other eggs were

found in the oviducal glands or oviducts. The

mature egg, oval in shape, measures 3 mm long by

1.7 mm at the widest point. The deteriorating eggs

are sliglitly more than .5 mm in length. A number

of sperm reservoirs were found attached to the

buccal membrane. Clusters of small oval vesicles also

were found in the buccal membrane; broad

funnel-shaped ducts connected the clusters to the

oral surface of the buccal membrane (one funnel

per cluster). Presumably these are organs for storing

sperm. All of the vesicles examined appear to be

empty.

Gonatus berryi Naef. 1923. A single female (185

mm P. L), tentatively identified as this species on

the basis of the massive size of the arms, showed

evidence of having spawned. In most respects the

features of this specimen are the same as in G.

pyros. Tlie consistency is gelatinous; suckers are

absent; the ventral arms are completely bare and the

tentacles are absent. Tlie buccal membrane lacks

sperm reservoirs but most of the membrane is

Vol. 87 (2)

THE NAUTILUS

55

missing. Nidamental and oviducal glands are some-

what enlarged (35 mm and 25 mm respectively).

(For comparison, nidamental and oviducal glands of

an immature specimen (103 mm P. L.) measured 5

and 4 mm respectively). The ovary is small but has

a large number of small, elongate immature ova.

Interspersed among these cells are a number of

spherical, pale orange ova in various stages of ma-

turity, the largest of which is about 3.5 mm in

diameter.

Gcmatus onyx Young, 1972. One presumably

spent female has tentatively been identified to this

species, primarily on the basis of body proportions.

This specimen is somewhat different than the

others; the tissues are much firmer and more mus-

cular but still not as muscular as in an immature

specimen of this species. Both tentacles are missing

and suckers are absent from the dorsal three pairs

of arms. The ventral arms are bare except for a few

suckers at the arm bases. The nidamental and

/

..-'ft;®;'/'.'/'- ■'■■ ' ■

•V,.';

FIG. 1. Gonatus sp. from the Arctic Ocean. A, Ventral view; B, Dorsal view.

56 THE NAUTILUS

April 1973

Vol. 87 (2)

oviducal glands are somewhat enlarged (34 mm and

22 mm respectively). A small portion of the buccal

membrane is missing; the remaining portion is

strewn with sperm reservoirs and some of these are

attached to the bases of the arms. The ovary is

somewhat larger than in the other species and is

packed with long (about 1 mm in length), slender

immature eggs with a few larger orange, sperical,

partically mature eggs (1 mm diam.) interspersed.

Apparently a larger percentage of the eggs of this

specimen failed to mature completely before spawn-

ing.

These three species of Gonatus correspond rather

closely in their appearance with the specimen from

the Arctic Ocean. All show similar signs of degen-

eration, and comparable conditions of the ovary,

oviducal and nidamental glands; and several have

sperm reservoirs stOl attached to the buccal mem-

brane. The evidence strongly suggests that all of

these specimens are spent females.

Relatively little information is in the hterature

regarding the degeneration effects related to spawn-

ing in squids. Fields (1965) showed that the mantle

of spent females of Loligo opalescens decreased

greatly in thickness and girth and he concluded that

the mantle is the chief site for storing food reserves

consumed during the spawning period. He also

noted that oviducal and nidamental glands decreased

greatly in size after spawning but were still rela-

tively larger than these glands in immature females.

In an immature female (ML 87 mm) he found that

the nidamental gland was 0.6% of the total body

weiglit. In a mature female (ML 151 mm) the

nidamental gjand was 22.8% of the body weiglit

while in a spent female (ML 151 mm) the figure

was 5%.

McGowan (1954) observed a mass mortality of

the squid Loligo opalescens following copulation

and spawning. He noted that in the dead and dying

squid that the mantles were thin and limp. Sasaki,

1913 (according to Hamabe, 1963) found that in

the squid, Watasenia scintillam, males die after

mating in off-shore waters while females die after

spawning in coastal waters. Hamabe (1963) noted

that the cuttlefish Sepia esculenta apparently dies

after spawning. For Todarodes pacificus, Hamabe

(1963) found that in females with a reduced num-

ber of eggs in the ovary the mantle was thin and

flabby, the liver was reduced in size and firmness

and the stomach walls were thin and flabby. These

animals had large numbers of eggs in the oviducts

and Hamabe assumed that they had not yet

spawned but were about to do so.

It has been thought for many years that Octopus

dies after brooding its eggs (Nixon, 1969). Recently

W. Van Heukelem (in press) has noted rather rapid

degenerative changes occurring in the muscles of

female octopods in aquaria upon completion of the

brooding period and just prior to death.

The evidence suggests that degeneration and

death foUowing spawning by female squid is a

common if not universal phenomenon, and that the

same situation occurs in octopods although delayed

to the end of the brooding period (See also dis-

cussion in Arnold, in press).

Presumably the Arctic specimen had spawned

and was about to die when captured. Spawning

therefore probably occurred near March 30 and

about 79°58' N lat. and 170°23'E long., the time

and place of capture. This point is over water 2655

m deep in the Hyperborean Basin of the Arctic

Ocean. No information is available on the amount

of time which passes from spawning to hatching in

any gonatid. It is also not known whether gonatids

lay pelagic or benthic egg masses. Nesis (1971a)

estimates that the young specimens of Gonatus

from the northern Norwegian Sea hatched in April

or May.

ACKNOWLEDGMENTS

I thank the following people for reading and

commenting on the manuscript: J. Arnold, Pacific

Biomedical Research Center; M. Nixon, University

College London; M. Okiyama, Japan Sea Reg. Fish-

eries Res. Lab. Niigata; W. VanHeukelem and J.

McMahon, University of Hawaii. Tlie squid was

collected througli a program supported by NONR

228(19), NR 307-270 Office of Naval Research

contract with the University of Southern California

- work being pursured now under Dr. Hector Fer-

nandez: contract NOOO14-67-A-0269-0013 NR

307-270. I also thank Mrs. Constance McSweeny for

preparing the illustrations.

LITERATURE CITED

Arnold, J. M. 1972. Cephalopoda, Decapoda. In:

Reproduction of Marine Invertebrates. Giese, A.

and J. Pierce, edit.. Academic Press, In press.

Berry, S. S, 1925. The Cephalopoda collected by

the Canadian Arctic Expedition, 1913-18. Rept.

Can. Artie Exped. 1913-18, 8(B): 3-8.

Vol. 87 (2)

THE NAUTILUS

57

Fields, G. 1965. The structure, development, food

relations, reproduction, and life history of the

squid Loligo opaleseens Berry. Fish Bull. 131,

1-108.

Fields, W. G. and V. A. Gauley. 1971. Preliminary

description of an unusual gonatid squid

(Cephalopoda: Oegopsida) from the North Pa-

cific. Jour. Fish. Res. Bd. Canada 28(11):

1796-1801.

FIG. 2. Gonatus sp. from the Arctic Ocean. A, Left third arm; B, Left ventral arm; C, Radula; D, Beaks; E,

Large arm hook; F, Funnel-locking cartilage; G, Funnel organ.

58 THE NAUTILUS

April 1973

Vol. 87 (2)

Hamabe, M. 1963. Exhaustion process of the genital

organs of common squid, Ommastrephes sloani

pacificus. Bull. Japan Sea Reg. Fish. Res. Lab.

11: 1-11 [in Japanese] .

MacGinitie, G. E. 1955. Distribution and ecology of

the marine invertebrates of Point Barrow, Alaska.

Smithsonian Misc. Coll. 128(9): 1-201.

McGowan, J. A. 1954. Observations on the sexual

behavior and spawning of the squid Loligo

opalescens at La Jolla, California. Calif. Fish

Game 40: 47-54.

Naef, A. 1923. Die Cephalopoden. Systematik.

Fauna u. Flora Neapel, Monogr. 35, 1(1):

149-863.

Nesis, K. N. 1971a. The squid Gonatiis fabricii at

the center of the Arctic Basin. Hydrobio. Jour.

7(1); 93-96 [in Russian].

Nesis, K. N. 1971b. The family Gonatidae - abund-

ant squids of the North Pacific (their dis-

tribution, ecology, system and phylogeny).

Fourth Meeting of Investigation of Molluscs,

Academy of Sciences, U. S. S. R., Zoological

Institute, pp. 63-65 [in Russian].

Nixon, M. 1969. The hfespan of Octopus vulgaris

Lamarck. Proc. Malac. Soc. London 38: 529-540.

Okiyama, M. 1969. A new species of Gonatopsis

from the Japan Sea, with the record of a spec-

imen referable to Gonatopsis sp. Okutani, 1967

(Cephalopoda: Oegopsida, Gonatidae). Pub. Seto

Mar. BioL Lab. 17(1): 19-32.

Roper, C. F. E. and W. L. Brundage, Jr. 1972.

Cirrate octopods with associated deep-sea or-

ganisms: new biological data based on deep

benthic photographs. Smithson. Contr. Zool. No.

121: 1-46.

Sasaki, N. 1913. Ecology of hotaru-ika (Watasenia

scintillans (Berry)). Zool. Mag. 30(302): 581-590.

Van Heukelem, W. 1972. Life span of Octopus

cyanea Gray. Jour. Zool. (London), in press.

Voss, G. L. 1967. The biology and bathymetric

distribution of deep-sea cephalopods. Studies in

Tropical Oceanography, Miami 5: 511-535.

Young, R. E. 1972. The systematics and areal

distribution of the cephalopods from the seas off

southern California. Smithson. Contr. Zool., No.

97: 1-159.

BOOK REVIEW

SEA SHELLS OF TROPICAL WEST AMERICA.

Marine Mollusks from Baja California to Peru.

Second Edition. By A. Myra Keen; with assis-

tance by James H. McLean, xvi + 1064 pp., 4000

illus., 22 pages in color. Index. Stanford Uni-

versity Press, Stanford, Calif. 1971. $29.50.

It seems almost superfluous to recommend this

classic compendium to both professional and

amateur students of the tropical West American

marine mollusk fauna. The second and greatly en-

larged edition is the labor of love of A. Myra Keen

and a number of her able associates, such as James

H. McLean who undertook the herculean task of

organizing the section on the Turridae and most of

the Archaeogastropoda. Twila Bratcher and Robert

Burch contributed the revision of the Terebridae;

Eugene Coan and Barry Roth, the Marginellidae;

and Spencer Thorpe, the chitons.

Over 3,325 species are included, most of which

are well-illustrated and accompanied by succinct

descriptions, ranges, and synonyms. The updated

bibliography is one of the most complete ever

assembled for the mollusks of a major marine

province. The index, a measure of the magnitude of

the book, contains over 7,000 scientific entries. A

bonus comes to the users of the book in the form

of a good glossary, a geographical guide to and a

series of maps of the area, and tables for fath-

om-meter-feet conversion.

A very welcome addition is a fairly full treat-

ment of the known nudibranchs, thanks to the

efforts of Jame Lance. This rapidly growing subfield

of malacology has been greatly stimulated by the

inclusion of these shell-less orphans of conchology.

Twelve new plates of stunning color photographs

of 72 living mollusks add to the beauty and use-

fulness of this remarkable faunal guide.

In the face of such a huge, meticulous and

detailed documentation, one cannot resist repeating

Dr. Keen's quote from another woman writer,

George Eliot; "Why, you might take up some light

study - conchology now; I always thought that must

be a liglit study."

R. Tucker Abbott

Delaware Museum of Natural History

Vol. 87 (2)

THE NAUTILUS

59

SHORT PAPERS

RANGE EXTENSIONS OF CORBICULA

MANILENSIS (PHILIPPI) IN THE ATLANTIC

DRAINAGE OF THE UNITED STATES

Samuel L. H. Fuller

and

Charles E. Powell, Jr.

Academy of Natural Sciences of Philadelphia

Philadelphia, Pennsylvania 19103, U. S. A.

ABSTRACT

Corbicula manilensis (Philippi) is newly recorded

from the Savannah, Pee Dee, and Delaware river

systems in the Atlantic drainage of the United

States.

In a recent issue of this journal, Sickel (1973,

The Nautilus, 87 (1); 11-12) discussed the appear-

ance of Corbicula manilensis (Philippi) in the

Altamaha river system of Georgia, where this Asian

species was first detected in 1971. According to the

age of Sickel's oldest material, establishment of

Corbicula in this system had probably occurred as

early as 1968. "It will be only a matter of time

[Sickel concluded] before Corbicula moves north-

ward to inhabit the remainder of the Southern

Atlantic Slope region." To the probable detriment

of the native Atlantic drainage benthos, that time is

come.

Several living Corbicula were taken 20 September

1972 by Dr. Selwyn S. Roback and one of us

(SLHF) from the Savannah River, about 9 miles

northeast of Milhaven, Screven County, Georgia.

The specimens lived in shifting sand on a bar at the

mouth of an enormous slough; none was found at

depths greater than about 1 foot of water. Visits to

this locality in 1971 had failed to produce

Corbicula, but this material falls into at least two

size classes, and, at 20.0 mm in length, the largest

individual may have been in its second year of life.

During the previous week (12 September), Mr. C.

Kirkland Dunlap, Jr., and one of us (SLHF) had

found one living Corbicula and several "gapers" in

the Pee Dee River, about 3 miles southeast of

Society Hill, Darlington County, South Carohna. At

this point the Pee Dee floor is chiefly of hard clay,

with pockets of sand, which drifts in from springs

along the river margin. The living specimen was

found on bare clay in a few inches of water. The

largest individual (a gaper) measures 28.5 mm in

length, shows extensive erosion at the beaks, and

appears to have reached at least two years of age.

During September and October, 1972, one of us

(CEP) found Corbicula at numerous localities in the

Delaware River between Philadelphia, Pennsylvania,

and Trenton, New Jersey. The species was common

in no more than 1 8 inches of water on a variety of

substrates, primarily muds and fine gravels. The

largest individual collected is 16.5 mm in length and

appears to have been in its second year.

Thus Corbicula has now invaded the Savannah,

Pee Dee, and Delaware river systems, in each case

probably as eariy as 1971. Sickle argued per-

suasively that the species reached the Altamaha

system from the Apalachicola river system of the

eastern Gulf of Mexico drainage by means of man's

inadvertent carriage overland. We, on the other

hand, have no evidence which would incriminate

any given vehicle in the rapid and far-reaching

dispersal of Corbicula recorded here.

•

STREAM POLLUTION

REPORTED 104 YEARS AGO

"In the Erie Canal, Anodonta Lewisii Lea, for-

merly very abundant, is now quite scarce, prob-

ably on account of chemical contamination from

the gas works and armory at Uion, a mile and a

half west from Mohawk" [N.Y.] . - James Lewis,

M.D., Amer. Jour. Conch., vol. 4, p. 245, 1869.

•

JUNE A. M. U. MEETING

PLANS COMPLETE

Delaware will be the site of the 39th Annual

Meeting of the American Malacologjcal Union from

Monday, June 25, through Thursday, June 28,

1973. The main meetings will be held at the new

air-conditioned conference center on the campus of

the University of Delaware, Newark, Del. On Tues-

day there will be a guided tour of the Delaware

Museum of Natural History, with special films and

additional A. M. U. papers being given there. Pro-

spective new A. M. U. members are also welcome,

and are asked to send their annual dues ($4.00 per

person and $1.00 for an additional family member)

to obtain registration and dormitory reservation

forms from Mrs. Marian S. Hubbard 3957 Marlow

Court, Seaford, New York 11783.

60 THE NAUTILUS

April 1973

Vol. 87 (2)

EMARGINULA DENTIGERA HEILPRIN, 1889,

A LITTORAL MOLLUSK

Margaret C. Teskey

P. 0. Box 273

Big Pine Key, Florida 33043

From more than sixty shallow water stations in

the Florida Keys, both reef and shore, no specimen of

Emarginula was collected until a colony was dis-

covered in 1972 at the north (GulQ end of Big Pine

Key. Here the live mollusks cUng to slab limestone

rubble that was pushed into the water to form a nar-

row, sloping shore when a bordering road was con-

structed two years ago. They occur occasionally at

tide line but are most abundant at the two-foot

depth. Density of population may be judged by the

fact that on about fifty feet of shore, one hundred

specimens were taken by one collector in less than an

hour. Only Aaiiaea pustulata (Helbling) is more

numerous at the site.

Identification was made by Dr. R. Tucker Abbott

who remarked ( in litt.) that althougli Emarginula

dentigera has been considered an ecological form of

E. pumila A. Adams (Farfante, 1947), he is inclined

to recognize it as a valid species. E. pumila is reported

as extremely variable in size and shape; The Big Pine

Key shells are remarkably uniform, high-spired and

somewhat narrow. Adults vary little from a length of

6 to 7 mm,, width 4 mm., height 3 mm. The apical

whorls form a definite hook at about 2/3 of the

posterior length of the shell; color chalky-white or

FIGS. Emarginula dentigera /ye(7pn», 1889, from Big

Pine Key, Monroe Co., Florida. X5.

stained green; interior glossy; sculpture of 18 to 26

radial ribs of varying size and of numerous, smaller

concentric threads, giving a slightly cancellate appear-

ance.

Emarginula dentigera Heilprin, 1 889, was original-

ly described from Bermuda, as was its synonym, E.

pileum Heilprin, 1 889.

LITERATURE CITED

Farfante, Isabel Perez, 1947. Johnsonia, 2, (24):

107-109.

Heilprin, A. 1889. On Some New Species of Mollusca

from the Bermuda Islands, Proc. Acad. Nat. Sci.

Phila. for 1889: 141-142, pL 8, figs. 6 and 7.

•

New Council of Systematic Malacologists

On November 17, 1972, a group of 11 American

malacologjcal systematists, representing some of the

leading moUusk collections in the country, gathered

at the Academy of Natural Sciences of Philadelphia

to form a "CouncU of Systematic Malacologists." The

purposes of this organization are 1 ) to prepare recom-

mendations concerning standards and techniques of

collection management for malacological resources;

and 2) to define the goals and priorities of malaco-

logy. Working subgroups were appointed to prepare

recommendations concerning:

1) Compatible EDP (electronic data processing)

and curatorial programs for use by malacologists in

collection management and research;

2) Articulation of national goals for malacological

research;

3) Coordination of specimen acquisition policies

for malacological collections;

4) Preparation of position papers outlining priority

programs for malacology.

Membership to the Council is open to all profes-

sional persons in the Americas who use or manage

systematic malacological resources. Further informa-

tion may be obtained from the Chairman of the

Steering Committee, Dr. Alan Solem, Dept. of

Zoology, Field Museum of Natural History, Chicago,

111. 60605. It is hoped that other institutions and

their managerial malacologists will be added to the

initial participants: Acad. Nat. Sci. Phila.; Delaware

Mus. Nat. Hist.; Field Mus. Nat. Hist. (Chicago); Los

Angeles Co. Mus. Nat. Hist.; Mus. Comp. Zool. at

Harvard; National Mus. Canada (Ottawa); Nat. Mus.

Nat. Hist. (Wash., D.C.); Ohio State Univ. Mus.; Nat.

Hist. Mus. San Diego; Mus. Zool., Univ. Mich.

INFORMATION FOR SUBSCRIBERS

The annual subscription rate for The Nautilus is

$7.00 for individuals and $1 2.00 for institutions (do-

mestic or foreign). Subscriptions may be commenced

in January when a new volume begins. Send check or

money order to "The Nautilus" to Mrs. Horace B.

Baker, Business Manager, 1 1 Chelten Road, Haver-

town, Pa. 19083.

Back issues from volume 72 to date are obtainable

from the Business Manager. Volumes 1 through 71 (if

available) may be obtained in reprint or original form

from Kraus Periodicals, Inc., 16 East 46th Street,

New York, N.Y. 10017.

Advertising rates may be obtained from the

Business Manager or Editor.

CONTRIBUTORS

Manuscripts: Authors are requested to follow the

recommendations of the Style Manual for Biological

Journals, which may be purchased from the American

Institute of Biological Sciences, 2000 "P" Street,

N.W. Washington, D.C. 20036. Manuscripts should be

typewritten and doublespaced; original and one copy

are required, to facilitate reviews. Tables, numbered

in arable, should be on separate pages, with the title

at the top. Legends to photographs should be typed

on separate sheets. Explanatory terms and symbols

within a drawing should be neatly printed, or they

may be pencilled in on a translucent overlay, so that

the printer may set them in 8 pt. type. There is a

charge of 50 cents per word for this extra service. All

authors or their institutions will be charged 50 cents

per line of tabular material and taxonomic keys. The

publishers reserve the right, seldom exercised, to

charge $32 per printed page.

An abstract should accompany each paper.

Reprints and covers are available at cost to

authors. When proof is returned to authors,

information about ordering reprints will be given.

They are obtained from the Economy Printing Co.,

Inc., R. D. 3, Box 169, Easton, Maryland 21601.

JULY, 1973

THE

NAUTILUS

;ie=B!-J

Vol. 87

No. 3

A quarterly

devoted to

malacology and

the interests of

conchologists

Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker.

Editors: R. Tucker Abbott and Charles B. Wurtz

O

EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of Mollusks

National Museum of Canada

Ottawa, Ontario, Canada K1A-0M8

Dr. WUliam J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. €2 138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 1 0024

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

Dr. James H. McLean

Los Angeles County Museum of Natural History

900 Exposition Boulevard

Los Angeles, California 90007

Dr. Arthur S. Merrill

Biological Laboratory

National Marine Fisheries Service

Oxford, Maryland 21654

Dr. Donald R. Moore

Division of Marine Geology

School of Marine and Atmospheric Science

1 0 Rickenbacker Causeway

Miami, Florida 33149

Dr. Joseph Rosewater

Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. Gilbert L. Voss

Division of Biology

School of Marine and Atmospheric Science

10 Rickenbacker Causeway

Miami, Florida 33149

EDITORS

Dr. R. Tucker Abbott

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807

Dr. Charles B. Wurtz

3220 Penn Street

Philadelphia, Pennsylvania 19129

Mrs. Horace B. Baker

Business and Subscription Manager

1 1 Chelten Road

Havertown, Pennsylvania 19083

OFFICE OF PUBLICATION

Delaware Museum of Natural History

Kennett Pike, Route 52

Box 3937, GreenvUle, Delaware 19807

Second Class Postage paid at Wilmington, Delaware

Subscription Price: $7.00 (see Inside back cover)

THE

NAUTILUS

Volume 87, number 3 - July 1973

CONTENTS

Ronald P. Thomas and Lee Opresko

Observations on Ocropus joiibim: Four Laboratory Reared Generations 61

Dorothea S. Franzen

Oxyloma dcprimida, a New Species of Succineidae (Pulmonata) 66

David Nicol and Graig D. Shaak

Late Eocene Distribution of the Pelecypod Exputens in Southeastern United States 72

John N. Kraeuter

Notes gn Mollusics Ostrea and Siphonaria from Georgia (U.S.A.) 75

Maria Luiza S. Mello and Marlene T. Ueta

Observations by Polarized Liglit on the Radula of Aplexa marmorata 79

Alan K. Craig

A New Record for Liginis: The Boynton Beach Colony 83

Stuart A. Harris

PisiJium heiisluwanwu (Sheppard) in Western Canada 86

James T. Carlton

Corbicula in San Francisco, California 87

Robert Robertson

Cyclostremella: A Planispiral Pyramidellid 88

Donald R. Shasky

Obituary of G. Bruce Campbell, M.D 89

Frank Perron and Thomas Probert

Vivipanis malleatus in New Hampshire 90

Book Reviews

(of) Tom Rice, 77; J. C. Bequaert and W. B. Miller, 78; Recent Publications, 74.

J.ne iJneU Cabinet

Successor to :

Richard E. Petit

and

John Q. Burch

OLIVE SHELLS

OF

THE WORLD

SPECIMEN SHELLS

NATURAL HISTORY

BOOKS

ROWLAND F. ZEIGLEK

HUMBERT a PORRECA

International Directory of Conchologists - $3.00

A current worldwide list of 2,000 people interested

in mollusks and in exchanging shells. Gives addresses

and interests. Available only from The Shell Cabinet.

Olive Shells of the World bv Zeigler and Porreca -

$12.95

13 full-page color plates illustrating over 150 species

and named varieties. The only current treatment of

the popular family. Clothbound. Available from your

book store or directly from us.

} Other Recently Published Shell Books Available

I Post Office Box 29. Falls Church, Virginia 22046

A Rare Callectar^s I town .

now available at a startlingly low price!

Now only $n.7'> (formerly $30.00)

The few remaining copies of this unusual facsimile have

been purchased by the Delaware Museum of Natural History

and are now made available practically at cost to all libraries,

scientists and amateur conchologists. 48 beautiful, colored

plates, representing a faithful reproduction of Swainson's

1834 and 1841 classic, together with the original text and a

modern analysis by R. Tucker Abbott and Nora NfcMillan.

Handsomely bound, gilt-edged on 3 sides and in full 9x12

inch page size. A useful and rare work belonging in the

library of every lover of mollusks. Both available from your

favorite book dealer or directly from the

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807, U.S.A.

INDO-PACIFIC

MOLLUSCA

MONOGRAPHS OF THE MARINE MOLLUSKS OF

THE WORLD WITH EMPHASIS ON THOSE OF

THE TROPICAL WESTERN PACIFIC

AND INDIAN OCEANS

The most technical and most beautifully illustrated

journal now being published on Recent and Tertiary

marine mollusks. Over 20 professional malacologists are

currently contributing. Edited by R. Tucker Abbott.

Among the groups treated are Strombidae, Cassidae,

Tridacnidae, Turridae, Littorinidae. Phasianelhdae, and,

soon to come. Patellidae, Harpidae and Mitridae.

Issued to date in looseleaf form with three sturdy,

permanent binders — 1100 pages, 810 plates (31 in full

color). Limited number of complete .sets left, $91.90 U.S.

(foreign: $94.00), postage paid. Any number of extra

binders available at $6.00.

Published by

The Delaware Museum of Natural History, Box 3937, Greenville, Delaware 19807 U.S.A.

VI

Vol. 87 (3)

THE NAUTILUS

61

OBSERVATIONS ON OCTOPUS JOUBINI: FOUR LABORATORY

REARED GENERATIONS'

Ronald F. Thomas and Lee Opresko

University of Miami, Rosenstiel School of Marine and

Atmospheric Science, Miami, Florida 33149

ABSTRACT

Four consecutive generations of Octopus joubini Robson were reared in the

laboratory. Spawning occurred at 4-5 months after hatching. At 27-29° C,

development of the eggs was completed in 34-42 days. Fiddler crabs /'Uca spp.j were

used almost exclusively for food, although other animals were also presented to the

octopods. Results indicate that there are probably two generations of 0. joubini per

year in the field.

INTRODUCTION

Few laboratory studies have been carried out

on the life history of octopods. Discussion of

hatching and rearing of the fry of sepioid squids

were presented by Choe and Oshima (1963), Choe

(1966). Schroder (1969a, b) and Boletzky et al.

(1971). The loliginid squids Sepioteuthis sepioidea

and Doryteuthis plei were discussed by LaRoe

(1971). Wolterding (1971) reared Octopus briareus

from egg to maturity and Arnold et al. (1972)

have reared Euprymna scolopes.

Owing to their accessibility and relative hardi-

ness in aquaria, some shallow water octopods, and

particularly species of Octopus, have been studied.

Octopus vulgaris was used extensively by J.Z.

Young in his classic series of experiments on

learning and behavior. Nixon (1969) studied the

rate of growth of O. vulgaris kept in the

laboratory. Boletzky (1969) studied the early

development of O. vulgaris, O. joubini and O.

briareus. In the same year Boletzky and Boletzky

reported on the first success in raising O. joubini

to maturity in the laboratory. From one brood of

about 50 eggs collected in the field, one female

survived to maturity and spawned about S^A.

months after hatching.

'Contribution No. 1592 from the University of

Miami, Rosenstiel School of Marine and Atmos-

pheric Science. This work was supported in part

by National Science Foundation grant

GB-24994X.

Octopus joubini is a species of small octopus

which is found in shallow waters along the west

coast of Florida and the northwestern Caribbean.

Its small size (about 10 cm. in total length at

maturity) makes it well-suited for laboratory

culture. The present paper reports on four genera-

tions of 0. joubini reared in laboratory. Observa-

tions on culture, feeding, brooding and life span

are given.

METHODS AND MATERIALS

This study was begun in February 1971 and is

still in progress. The octopods were maintained in

aquaria of 5-, 10-, and 20-gallon capacities and

FIG. 1. Adult male Octopus joubini //; aquarium.

Natural size. Photo by Dennis Opresko.

62 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

FIG. 2. Eggs being brooded by female 0. joubini

on inside of empty flower pot. x I'A. Photo by

Dennis Opresko.

supplied with running sea water piped from

Biscayne Bay. The bay water was first passed

through settling tanks and was then filtered

througlr a series of gravity-fed compound filters

containing layers of CaCO pebbles and sand and

glass wool. With this system, adequate water

quality and aeration for the survival of the

octopods was maintained. Additional aeration was

provided by means of a mechanical pump as a

safeguard against water stagnation due to possible

blockage of the filters and subsequent loss of

water flow.

The initial brood and female were obtained

from inside a bottle found in shallow water of

about 1-2 meters at Key Biscayne, Miami, Florida.

The newly-hatched animals were supplied refuge

by astro turf floating on the water-surface. After 4

weeks, flower pots, rocks and shells were provided

for the animals. Mature females would readily

deposit eggs in the flower pots. Octopus joubini

attaches each egg individually to the substrate.

After hatching, the attachment point of the egg

on the substrate can be seen as a dark spot; this

permitted the estimation of the size of the brood.

For the first several days after hatching, the

octopods were still receiving nourishment from the

internal yolk sac and would not feed. During the

following 2 to 3 weeks, the juveniles would not

accept live ilea as food and were individually fed

daily. A segment of a Uca appendage placed in

contact with the octopus was usually seized and

eaten. After several weeks, the young animals

accepted live Uca of appropriate size.

The water level was kept several inches below

the tops of the aquaria to discourage animals from

crawling out of the tanks. This method was found

to be successful for this species.

OBSERVATIONS

At an early age O. joubini selected a home site,

either in a flower pot or in a rock crevice. Seven

adult octopods, each provided with a suitable

refuge and placed together in a 20-gallon aquari-

um, did not display any territorial aggressive

behavior. Under these conditions no deaths in any

aquaria were considered to be caused by other

octopuses.

The animals consumed a considerable amount

of food (Uca). The young octopods successfully

attacked crabs equal to or larger than themselves.

They were strongly nocturnal, remaining in crev-

ices during the day and searching for food at

night. If a liglit were turned on at niglit while an

octopus was feeding, it would drop the partially

consumed crab and return to its den. It would

later attack another Uca rather than continue to

feed on the dead, half-eaten crab.

In table 1. temperatures in the aquaria are

given. Small fluctuations of aquarium temperatures

(from 1-2 C) tended to reflect those of the room

rather than those of the field, but in general,

aquarium temperatures were comparable to those

in the field.

Fj GENERATION

The F generation hatched on 13 February

1971. The newly hatched animals grew slowly,

probably because of an inadequate supply of food.

The most critical stage in rearing O. joubini occurs

between 2 and 21 days of age, since the animals

would not attack live food. Consequently this is

the time during which the higliest mortality

occurred. Later mortalities usually occurred when

the sea water system became blocked and addi-

tional aeration was not used. As the water in the

tank began to foul, the animals responded by

leaving their dens and moving to the water

surface.

Three F females laid eggs 4'/i months after

hatching, all within several days of each other.

Vol. 87 (3)

THE NAUTILUS

63

Thus the maturation ot~ laboratory-raised O. joii-

bini was sutTiciently rapid to allow two genera-

tions to be completed within a year.

One female laid about 40 eggs. The length of

these eggs (7.6 mm.) did not differ significantly

from measurements reported in the literature

(8.0-8.5 mm.) by Boletzky (1969).

Once the eggs were laid, the female remained

with them until they hatched. Her arms continu-

ously swept over the eggs to remove detritus that

settled on them. She would not leave the eggs to

feed. If a crab entered the octopus' den while she

was brooding eggs, she would generally attack and

consume it. Due to the higli density of crabs in

each tank, females were able to feed while

brooding. Although it is generally believed that the

female octopus dies soon after the eggs hatch, this

did not always occur in the laboratory, probably

due to the availabihty of food during brooding.

At 27-30°C, hatching occurred within 34-42

days, usually at niglit. Near the time of hatching,

mechanical stimulation caused by examining the

eggs resulted in premature hatching of some eggs.

Although these octopods hatched prematurely,

they were still able to survive.

The females did not lay a second brood of

eggs. After the eggs hatched both males and

females showed a reduction in feeding intensity

(the animals now fed sporadically rather than

regularly). At about 6 months of age the F males

and females began to die. The mortality rate

steadily increased so that by 8 months of age

about 50% adult mortality was attained. Old,

dying O. joubini typically stopped feeding and

abandoned their dens. They often sat on the floor

or sides of the aquarium, slowly coiling and

uncoiling their arms in an uncoordinated manner.

F^ GENERATION

Eggs were laid by F females on 23 June 1971.

The F generation hatched between 34 and 42

days later. Growth of these animals was similar to

that stated for the F generation. Egg laying by

F^ females began at the end of November,

indicating that this generation reached maturity in

about 4 months. The water temperature in the

aquaria during the development of the F genera-

tion was 27-29°C, about 1-3°C higher than the

temperature during the development of the first

generation.

TABLE 1. Temperature values (°Cj measured in the

aquaria.

Date

21 July 1971

28 July

3 August

10 August

16 August

24 August

31 August

7 September

1 1 September

20 September

24 September

30 September

Temp (°C)

28.9

28.2

29.0

27.8

27.1

29.0

27.8

26.7

27.0

FIG. 3. Young O. joubini in typical posture, x 2.

Photo by M. Wolterding.

64 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

TABLE 2. Dates of hatching o/ Octopus joubini eggs

laid 23 June 1971. Water temperatures: 27-29° C.

Fj GENERATION

Approximately 95 eggs were laid by one of the

F females. Hatching of these eggs began on 26

December 1971. At the end of six weeks. 52

young were still alive. This generation was raised

very successfully, and about 42 animals reached

maturity, with 1 5 females laying broods of eggs.

The size of the F animals was quite small

(average mantle length of 5 males: 26.7 mm.; of 5

females: 21.9 mm.). We feel that stunting occurred

as a result of crowded conditions in the aquaria.

Inadequate food may have been a contributing

factor during the early growth period. Egg-laying

began at AVi months (15 May 1972) even in the

small females, althougli the number of eggs per

brood was low. It is interesting to note that O.

joubini in Biscayne Bay, Florida, were also

observed to have eggs at the same time as the F^

females (James Espy, personal communication).

F GENERATION

4

These octopuses began to hatch on 20 June

1972 and hatching continued to the middle of

July. A culture of brine shrimp (Artemia salina)

was established and maintained in an attempt to

replace the individual feeding of each juvenile

octopus. For this experiment, octopuses between 1

and 7 days old were supplied with brine shrimp of

various sizes, excluding newly hatched brine

shrimp. The brine shrimp were placed in the tank

with the octopus at 4:00 PM daily and the

running water was turned off. The Artemia were

observed to swim slowly around the tank, during

which time they often came in contact with an

octopus. Many times brine shrimp were seen to

crawl under the octopus. Instead of seizing the

brine shrimp, the octopus blanched, inked and

swam away. Brine shrimp held with forceps and

presented to the octopus were rejected. One

octopus, presented with brine shrimp in this

manner, accepted the food after many trials and

was later observed to capture live Artemia. This

procedure unfortunately met with little success

and it became necessary to resume the hand

feeding.

The number of F juveniles that hatched could

not be accurately estimated because many were

able to hide in the rocks that were provided for

the adults. About 70 were isolated and placed in

separate aquaria of 3- to 5-gallon capacity. One

month after the initial hatching occurred 15 of

the largest F O. joubini were placed in a single

20-gallon aquarium and were fed exclusively with

live Uca spp. The small individuals which remained

in the 3- to 5-gallon aquaria were fed pieces of

Uca appendages and were also provided with

suitably-sized live Uca.

DISCUSSION

Boletzky & Boletzky (1969) were able to raise

one female Octopus joubini to maturity. This

animal laid eggs 5'/i months after hatching. The

present study confirms the rapid development of

this species of octopus. Mangold- Wirz (1963)

found that females of Eledone cirrosa, E. mos-

chata. Octopus vulgaris and Pteroctopus tetracir-

rhus reached sexual maturity at about 18-24

months, while Bathypolypus sponsalis matured at

8-17 months.

According to Boletzky (1969), at 24-25°C the

eggs of O. vulgaris hatch in 25 days, O. joubini in

30 days, and O. briareus in 50 days. In this study,

hatching of O. joubini began at about 34 days,

thus agreeing with Boletzky's findings.

As is typical with most octopods, the female of

O. joubini diligently cares for the eggs until they

hatch. Brooding behavior weakens the female and

she generally dies soon after the eggs hatch, but

death appears to be delayed if she is able to feed

during brooding.

The rapid maturation of O. joubini in the

laboratory suggests that there are two generations

Vol. 87 (3)

THE NAUTILUS

65

annually. Thus in certain parts of the range it is

likely that there is both a summer and a winter

population of tiiis species. This species of small

octopus with its short life span provides an

opportunity to study development in the labora-

tory and to establish techniques for cephalopod

maintainance. An important problem, water qual-

ity, can be minimized by a running sea-water

system with filtering equipment. With running

sea-water, additional aeration is unnecessary but is

advisable in the event that the water flow is

temporarily blocked.

Feeding presents the other major problem in

the culture of benthonic octopods. At the present

time, suitable live food for the young juveniles has

not been found. Gammarid and caprellid amphi-

pods and various other amphipods as well as

numerous isopods were offered to the juveniles

but were rejected by them. This was found to be

the case for Artemia salina, except with one O.

joubini The largest and probably oldest (7 days

old) F juvenile which was offered brine shrimp

did finaUy attack Artemia. Even though the

octopus accepted them as food, the fact that the

smaller O. joubini would not eat brine shrimp

prevents this method from being used on newly

hatched O. joubini Artemia seems to be more

suitable for octopods with planktonic larvae and

has been used in rearing O. vulgaris (Boletzky,

personal communication).

The method of feeding with Uca described here

is time consuming and inefficient, contributing to

most of the mortalities at this stage of growth. In

areas where Uca is abundant, it provides a

satisfactory and easily obtainable source of live

food for the larger animals.

ACKNOWLEDGMENTS

We wish to thank Mr. Dennis Opresko and Mrs.

Irene Thomas for their help in maintaining the

animals and for their assistance in the preparation

of the manuscript. We also wish to thank Mr. D.

Opresko for photographing the animals. Dr. Won

Tack Yang provided helpful suggestions in the

culture of marine organisms and also provided the

stock of live Artemia. We especially wish to thank

Dr. Gilbert L. Voss for his support and encour-

agement during this study and for his critical

reading of the manuscript.

LITERATURE CITED

Arnold, J.M., C. T. Singley and L. D. Wil-

liams-Arnold 1972. Embryonic development and

post-hatclring survival of the Sepiolod squid

Euprymna scolopes under laboratory conditions.

The VeHger 14(4); 361-364.

Boletzky, S. von 1969. Zum Vergleich der Onto-

genesen von Octopus vulgaris, O. joubini. und

O. briareus. Rev. Suisse Zool. 76: 716-726.

Boletzky, S. von • & M. V. von Boletzky 1969.

First results in rearing Octopus joubini Robson,

1929. Verh. naturf. Ges. Basel 80: 56-61.

Boletzky, S. von, M. V. von Boletzky, D. Frosch,

& V. Gatzi 1971. Laboratory rearing of Sepio-

linae (Mollusca; Cephalopoda). Mar. Biol. 8:

82-87.

Choe, S. 1966. On the eggs, rearing, habits of the

fry, and growth of some Cephalopoda. Bull.

Mar. Sci. 16: 330-348.

Choe, S. & Y. Oshima 1963. Rearing of cuttle-

fishes and squids. Nature 197: 307.

LaRoe, E. T. 1971. The culture and maintainance

of the loliginid squids Sepioteuthis sepioidea

and Doryteuthis plei Mar. Biol. 9: 9-25.

Mangold- Wirz, K. 1963. Biologie des Cephalopodes

benthiques de la Mer Catalane. Vie et Milieu

(Suppl.) 13: 1-285.

Nixon, M. 1969. The lifespan of Octopus vulgaris

Lamarck. Proc. malac. Soc. London 38:

529-540.

Schroder, W. 1969a. Observations made during the

breeding of cuttlefish {Sepia officinalis L.).

Drum and Croaker 69: 9-15.

Schroder, W. 1969b. Notes concernant Cepha-

lopoda et la reproduction de la seich commune.

Zool. Anvers. 34th year (3).

Wolterding, M. 1971. The rearing and maintainance

of Octopus briareus in the laboratory with

aspects of their behavior and biology. M.S.

thesis, Univ. of Miami, 120 pp.

66 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

OXYLOMA DEPRIMIDA, A NEW SPECIES OF SUCCINEIDAE (PULMONATA)

Dorothea S. Franzen

Ulinois Wesleyan University

Bloomington, Illinois 61701

ABSTRACT

Oxyloma deprimida, a new species of Succineidae (Piilmonataj is described from

Utica, LaSalle County, lilinoiSj (type locality) and near Canton Lake, NE of Canton,

Fulton County, Illinois. The nuclear whorl of the shell is depressed, hence, the species

name deprimida. The habitat of O. deprimida is on cattails generally 3 to 5 ft. above

the wet ground or v^ater level The snails mature in the fall and not in the summer as

does O. retusa, a sympatric species. Anatomical features include a broadly based penial

appendix and an albumengland which is smaller than the prostate gland.

The search for succineas in the state of Illinois

has resulted in the recognition of a hitherto

undescribed species, Oxyloma deprimida, new spe-

cies.

Description of Holotype: The amber-colored,

ovate, imperforate shell, composed of three inflated

whorls, is thin, fragile, shiny, and marked only by

tme striae. The nuclear whorl is depressed (Fig. 1,

upper riglit), rough and pitted (Fig. 1, lower left).

The whorls increase rapidly in size from the nuclear

to the body whorl. The suture is sharply incised

resulting in a sliglit shouldering along the upper

border of the whorl. The peristome of the ovate

aperture is sharp and continuous over the surface of

the body whori in the form of a thin, white callus

(Fig. 2, left). The narrow, white columella follows

the inner border of the peristome (Fig. 2, left). The

dimensions of the shell are: heiglit, 12.45 mm.;

width, J. 6 mm.; height of aperture, 9.85 mm.;

width of aperture, 5.7 mm. Tlie ratios of the several

dimensions are shown in Table I.

The surface of the head and the body wall are

cream-colored and coarsely and irregularly tuber-

culate. The pigmentation of the anterior dorsal

portion of the head consists of black blotches

arranged in five, indistinctly defined, longitudinal

bands that form a shield-shaped pattern (Fig. 2,

riglit). Tliey converge medially between the pos-

terior tentacles and then spread apart continuing as

two bands towards the mantle. The shield pattern is

Hanked on either side by a pigmented band, narrow

anterioriy, but broadening at the base of the pos-

terior tentacles as it continues to the mantle. On

FIG. 1 . Upper left. Chromosomes of an ovotestis

smear of Oxyloma deprimida from the type

locality. Enlarged. Upper right, Scanning-elec-

tron-photomicrograph showing the nuclear whorl of

Oxyloma deprimida of a shell from the f}'pe locality.

Enlarged. Lower left, Scanning-electron photo-

micrograph of the nuclear whorl of Oxyloma

deprimida of a shell from the type locality, showing

surface detail. Enlarged. Lower right. Scanning-elec-

tron-photomicrograph of the nuclear whorl of a shell

of Succinea luteola, showing surface detaiL Enlarged.

Vol. 87 (3)

THE NAUTILUS

67

FIG. 2. Drawings of the holotype of Oxyloma

deprimida. Left, shell drawn to scale as indicated.

Right, head and anterior portion of mantle.

Enlarged.

the lateral body wall flecks of light pigmentation

form a broad band that tapers and diminishes

posteriorly. Tlie manfle collar is lightly pigmented

on the upper and outer surface and more heavily

pigmented on the anterior and inferior surface. The

pigmentation of the anterior portion of the mantle

occurs as widely-spaced blotches (Fig. 2, riglit). The

anterior border of the nephridium is outlined by a

narrow, broken, black band whereas the posterior

border is outlined by a broader, darker band.

Sparse, scattered patches of pigment occur on the

remainder of the body. The genital aperture, cres-

cent-shaped, 2 mm. in length, surrounded by a white,

tumid lip, is situated on the anterior right-hand side

of the body. A pedal groove, continuous from the

labial palp to the posterior tip of the body, sep-

arates the foot from the lateral body wall; the pedal

groove is paralleled dorsally by a less-pronounced,

suprapedal groove. Shallow, vertical grooves incise

the pedal and suprapedal grooves and the broad,

pigmented band. These vertical grooves produce a

series of shallow scallops along the margin of the

pedal groove especially when the animal is in a

somewhat contracted state.

Holotype: catalogue no. FMNH 176002; para-

types nos. FMNH 176003, FMNH 176004, and

FMNH 176005, Molluscan CoUection, Field Museum

of Natural History, Chicago, Illinois. Additional

paratypes are in the private collection of the author.

The author has examined shells of succineas of

the collection in the Natural History Museum, Uni-

versity of Illinois. A shell identified by Frank

Collins Baker (1939) as Succinea retusa peoriensis

Wolf and which Pilsbry (1948) identified as Oxy-

loma decampi peoriensis (Walker), as indicated be-

low, is herein identified as Oxyloma deprimida, new

species.

Synonymy —

Succinea retusa peoriensis Wolf. Frank Collins

Baker, 1939. Fieldbook of Illinois Land Snails,

Manual 2, Natural History Survey Division, p. 126,

Fig. E, left-hand figure only.

Oxyloma decampi peoriensis (Walker), Henry A.

Pilsbry, 1948. Land Mollusca of North America,

Vol. II, Ft. 2, p. 785, Fig. 420, b, left-hand figure

only.

Known geographic distribution and habitat: I.

Type locality: Field No. D.S.F. 350; NEVa SWA S 9,

Twp. 33 N, R 2 E, Utica (N. Utica), LaSaUe

County, Illinois, about Vz mile east of the crossing

of Illinois Highway 178 over the Rock Island and

Pacific railroad tracks, between the Rock Island and

Pacific railroad tracks and Clark Run Creek. The

habitat is the shore of a water-filled pit on the

northern edge of Clark Run Creek that supports

cattails (Typha sp.), goldenrod (Solidago sp.), and

willow (Salix sp.). Oxyloma deprimida lives on the

cattails several feet (2 to 5) above the wet ground

and/or above the water level.

An earher unpublished record of the species

taken from "Utica, Illinois" is in the Molluscan

Collection of the Field Museum of Natural History,

Chicago, Illinois. The shells, catalogue no. 58198,

were identified as Succinea (Oxyloma) decampi

Bryant Walker. They were formerly in the James H.

Ferriss collection and identified as Succinea retusa

peoriensis Wolf

Locality: 2. Field No. D.S.F. 194; NEy4 SEV*

NW54 S 19, Twp. 7 N, R 5 E; 3 miles NE of the

town square in Canton, Fulton County, Illinois. The

habitat is a lowland adjacent to a northeastern arm

of Canton Lake where moist ground supports a

stand of cattails and willows. Oxyloma deprimida

lives on this vegetation 3 to 5 feet above the

ground, and is not found on the ground where lives

O. retusa (Lea), a sympatric species. This association

does not, however, occur at the Utica locality.

Anatomy of paratypes: The large, coiled, heavily

pigmented hermaphroditic duct, coming from the

ovotestis, enters the fertilization sac which is con-

nected with the seminal vesicles. The relatively

large, twinned, unequal seminal vesicles are coarsely

peppered with black pigmentation whereas the in-

68

THE NAUTILUS

July 30, 1973

Vol. 87 (3)

flated fertilization sac is lightly pigmented. The

prostate gland is situated immediately anterior to

the albumen gland. The acinar prostate gland is

covered by a thin, lightly pigmented sheath, and is

larger than the albumen gland. The albumen gland,

about two-thirds the length of the prostate, is

composed of acini which are smaller than those of

the prostate gland. Tlie single duct leading from the

fertilization sac divides to form two ducts, one

leading into the prostate gland and the other form-

ing the oviduct (Fig. 3, F).

The vagina is straight, slightly pigmented, equal

to about seven-eighths of the length of the penis,

and opens along with the penis into the genital

antruiTL The oviduct is higlily convoluted, slightly

pigmented, and enters the distal end of the vagina

as does the spermathecal duct. The spermathecal

duct loops over the oviduct just prior to its entry

into the vagina.

The retractor muscle of the right posterior tent-

acle passes between the penis and the vagina. The

penis is enclosed in a thin, translucent sheath pep-

pered with black pigment in the middle or towards

its distal end. The vas deferens may or may not be

lightly stippled. As the vas deferens approaches the

distal end of the penial sheath it loops ventrally

under the penial retractor muscle and enters the

penial sheath terminaOy; then known as the

epiphallus, it coils, increases in caliber as it ap-

proaches the penis and becomes enlarged just before

it merges with the penis subterminally (Fig. 3, A).

The penial appendix, because of its broad base,

appears to be a terminal extension of the penis

rather than an appendix separated from the body of

the penis by a constriction. The size and shape of

the penial appendix is variable (Fig. 3, A - E).

The ovotestis ot many individuals of Oxyloma

deprimida, from both localities, were squashed and

l-R-L

2-R-M 4-R-M 21-R-M

FIG. 4. Representative teeth of a radula of a snail

taken from near Canton Lake; 3 mi. NE Canton,

Illinois. C, central tooth: l-R-L, 1st right lateral;

2-R-M, 2nd right marginal; 4-R-M. 4th right

marginal; 21-R-M, 21st right marginal.

stained with orcein. Examinations of the stained

chromosomes in metaphase revealed the haploid

number of nineteen to be characteristic of this

species (Fig. 1, upper left). This is the same number

found in O. retusa (Lea), O. haydeni (W. G. Binney)

and O. salleana (Pfeiffer), (Franzen, 1966, p. 67).

The radula (Fig. 4) is composed of many rows of

teeth and numerous teeth to a row. The number of

rows in 6 radulae examined, ranged from 90 to 116.

The numbers of marginals and laterals of re-

presentative rows of 5 radulae are to be noted in

Table II. The ratios of marginals to laterals ap-

proaches 1:3 and 1:4. Tliis compares with what has

been reported for Oxyloma ren<xa, (Franzen, 1963,

Table II, p. 89) and for O. haydeni, (Franzen, 1964,

Table II, p. 78).

The structural details of the individual teeth

resemble those of the genus in general (Fig. 4). The

central tooth has a broad basal plate which has a

lateral basal pointed projection on either side. The

pointed mesocone, somewhat variable in length,

does not usually extend to the lower margm of the

basal plate. A shorter, pointed ectocone flanks the

mesocone on either side. The laterals have a large,

pointed mesocone which, like tliat of the central

tooth, is usually not as long as the basal plate: the

single ectocone, sometimes divided into

two — especially of the more lateral teeth — is

pointed; a short, pointed endocone may be present.

The marginals, smaller than the laterals, have a long,

slender basal plate which is characteristic of the

genus Oxyloma (Quick, 1933:296, Fig. 1, Oxyloma

(Succineaj pfeifferi Rossm.). Of the cusps of the

marginals, the mesocone is the largest, the endocone

is small and pointed. The ectocone of about the six

medial-most marginals is divided into two unequal

cusps of which the lateral-most is the largest, and

curved. The ectocone of the more lateral marginals

is divided into three cusps of which the outermost

is the largest. The ectocone of the marginals nearest

the outer side of the radula may be divided into

four cusps of which the outermost is short and

pointed. The features of the amber-colored jaw are

typical of the genus. A prominent median fold

projects anteriorly and is flanked on either side by a

broadly rounded fold.

The surfaces of the bodies of the paratypes are,

as in the holotype, creanvcolored and coarsely and

irregularly tuberculate. The pattern of pigmentation

of the head and body is like that of the holotype

Vol. 87 (3)

THE NAUTILUS

69

althougli tlie intensity of pigmentation is variable.

The sole of the foot may be lightly or not at all

speckled. The most striking variation of pig-

mentation is that of the mantle. The pattern of the

anterior portion may be in the form of black,

widely-spaced blotches, as in the holotype, or

heavier blotches arranged in indistinctly defined

rows connected by less intense stripes of pigment,

or the blotches may merge, resulting in an irregular

pattern. The amount of pigmentation anterior to

the nephridium varies from light to heavy, the latter

in the form of a dark area interspersed by

fine-lined, light streaks. The intensity of the pig-

mentation outlining the nephridium is light in some

individuals and more intense in others. The pigmen-

tation of the area posterior to the nephridium varies

from a few scattered blotches to many patches

merging to form a more overall pattern extending to

the posterior tip of the animal.

Shell of paratypes: The shell of the mature snail,

attaining a height of almost 13.5 mm., is comprised

of 2^/2. to 3 inflated whorls. The surface of the

nuclear whorl is rougli and pitted (Fig. 1, lower

left) but not distinctly malleated as in Succinea

lutenla Gould (Fig. 1, lower riglU), S.

vaginacontorta Lee, and S. campestris Say.

Dimensions of the three largest shells, number of

shells measured, and the median of each series

measured, are recorded in Table L The range of the

greatest height of the series of shells included in this

study is from 9.8 inm. to 13.46 mm.; and the range

of the greatest width is from 6.7 mm. to 8.4 mm.

The greatest heiglit of shells obtained from the

Canton Lake locality is 11.8 mm.; the greatest

diameter is 8.4 mm. (October, 1960): the com-

parable dimensions of shells of the Utica locality

are, greatest height, 13.46 mm., greatest diameter,

8.04 mm. (September, 1967).

Distinctive characteristics of Oxyloma deprimida:

The name of the species, deprimida, denotes the

depressed position of the nuclear whorl (Fig. 1,

upper riglit) as contrasted with the prominent,

knoblike nucleus which is characteristic of other

known species of the genus. The tip of the nuclear

whorl appears to be somewhat "tucked in"" under

the penultimate whorl.

The epiphallus enters the penis subterminally

which produces a terminal extension of the penis,

the penial appendix. In O. deprimida the base of

the penial appendix is broad which makes it appear

to be a terminal extension of the penis rather than

an appendix distinctly separated from the body of

the penis by a constriction as in, for example, O.

haydeni, (Franzen, 1964, Fig. 1). The epiphallus

enlarges as it approaches the penis and merges

imperceptibly with it.

The albumen gland is smaller than the prostate

gland and is located posterioriy to it. The acini of

the albumen gland are distinctly smaller than are

those of the prostate.

Oxyloma deprimida lives on cattails and willows

generally 3 to 5 feet above the ground and/or the

water level, not on the ground nor on objects

floating on the water. The last time the type

locality was visited (September 28, 1971) the water

had receded to the level of the small stream as a

PENIAL RETRACTOR

MUSCLE

PENIAL SHEATH

FIG. 3. Drawings of genital organs of Oxyloma

deprimida: A - E, penis, penial appendix and

epiphallus shown inside of penis sheath cut open.

The figures are of snails from the following

localities: A, near Canton Lake, 3 mi. NE Canton,

Illinois: B, Utica. Illinois: C, Utica, Illinois: D,

Utica. Illinois: E, near Canton Lake. S mi. NE

Canton. Illinois. F. prostate gland and albumen

gland of a snail from the Utica, Illinois, locality:

PG, prostate gland: AG, albumen gland: FS,

fertilization sac: SV, seminal vesicles: HD, herma-

phroditic duct.

70 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

TABLE 1. Dimensions of the shells o/Oxyloma depressa. The measurements are of the 3 largest shells of each

of the 9 series as indicated. In the 4th column of measurements are listed the ratios of the width of the shell

over its height. In the last 3 columns are listed the ratios of the height of the aperture over the height of the

shell, width of aperture oner width of shell, width of aperture over height of aperture.

result of a period of about a year of low pre-

cipitation. The ground of the usually submerged

area was wet though not muddy. Even under such

conditions O. deprimida was not found on the

ground. Many of the snails were aestivating on the

cattails 2 feet above the ground.

Oxyloma deprimida differs from other species of

the genus, at least in the Mississippi Valley, as to

the time of year in which it reaches maturity. The

mature, fully grown, sexually active adults are to be

found from the latter part of September to the

early part of October. This is several months later

Vol. 87 (3)

THE NAUTILUS

71

TABLE 2. Formulae of representative rows of teeth o/Oxyloma deprimida new species.

No. of Rows

TABLE 3. Comparisons of the medians of ratios of several dimensions o/Oxyloma deprimida with

several other species of Oxyloma as previously reported (Franzen, 1969, Table III).

Species

O. deprimida

O. retusa

O. haydeni

O. salleana

0. sanibelensis

W. of Shell/

H. of Shell

59.7%.- 71.0%

52.0%

49.0%

56.3%

47.8%

H. Aperture/

H. Shell

73.7% - 78.0%

74.4%

73.0%

78.6%

60.7%

W. Aperture/

W. Shell

67.9% - 76.8%

80.5%

85.3%

79.2%

68.8%

W. Aperture/

H. Aperture

59.4% - 64.5%

57.1%

57.6%

55.2%

than the maturation time of O. retusa, O. haydeni,

and O. salleana whose mature adults disappear in

the latter part of June or in early July.

ACKNOWLEDGMENTS

National Science Foundation Grants-in-Aid No's.

NSF G18000 and NSF GB2715 provided laboratory

equipment and supported, in part, the field studies.

The author is grateful to Dr. B. V. Hall, Director,

Center for Electron Microscopy, Univ. of Illinois,

Urbana, and to Dr. James Anderson for the scan-

ning-electron-photomicrographs. Dr. Alan Solem,

Field Museum of Natural History, Chicago, Illinois,

and Dr. Donald F. Hoffmeister, Natural History

Musuem, Univ. of Illinois, Urbana, graciously lent

shells of the respective museum collections. Dr. A.

Byron Leonard read the manuscript and offered

helpful suggestions.

LITERATURE CITED

Baker, Frank Collins. 1939. Fieldbook of Illinois

Land Snails, Natural History Survey Division,

Manual 2, State of Illinois: 3-166.

Franzen, Dorothea S. 1963. Variations in the Anat-

omy of the Succineid Gastropod Oxyloma retusa.

The Nautilus 76 (3); 82-95, Tables I-II, Figs. 1-4.

Franzen, Dorothea S. 1964. Anatomy of the

Succineid Gastropod Oxyloma haydeni. The

NautUus 77(3): 73-81, Tables I-II, Fig. 1.

Franzen, Dorothea S. 1966. Anatomy of the

Succineid Gastropod Oxyloma salleana (Pfeiffer).

The Nautilus 80(2): 59-69, Tables I-II, Figs. 1-3.

Franzen, Dorothea S. 1969. Structural Character-

istics of Succineid Gastropod Oxyloma

sanibelensis. The Nautilus 82(3): 77-83, Tables

I-III, Figs. 1-2.

Pilsbry, Henry A. 1948. Land MoUusca of North

America (North of Mexico). Acad. Nat. Sci.

Philadelphia Mon. No. 3, Vol. 11, Pt. 2: xlvii -^

521-1113, 585 figs.

Quick, H. E. 1933. The Anatomy of British

Succineae. Proc. Mala. Soc. London 20(VI,

Nov.): 295-318 PI. 23-25, Figs. 1-18, Tables

1-V.

THE NAUTILUS

July 30. 1973

Vol. 87 (3)

LATE EOCENE DISTRIBUTION OF THE PELECYPOD KXPlTEys

IN SOUTHEASTERN UNITED STATES

David Nicol

Box 14376, University Station.

Gainesville, Florida 32601

and

Graig D. Shaak

The Florida State Museum

University of Florida

Gainesville, Florida 32601

ABSTRACT

The pelecypod Exputens (Malleidae) was confined To the Tethyan marine province

during the Middle and Late Eocene. In Peninsular Horida Exputens ocalensis

(MacNeil) is found in the two lowest zones of the Crystal River Limestone, which is

Late Eocene in age. Exputens has also been found in the Ocala Limestone of

southwestern Georgia and the Castle Hayne Marl in North Carolina, which shows

that at least one Tethyan mollusc lived farther north during the Late Eocene than

had previously been supposed.

Tlie presence of the pelecypod Exputens in

strata of Late Eocene age in Peninsular Florida has

been reported by Palmer, 1957. p. 70; 1967, pp.

189-190; and Palmer and Brann, 1965. p. 143.

However, Exputens was not mentioned as occur-

ring in Florida by Hertlein and Cox, 1969, in the

Treatise on Invertebrate Paleontology. Moreover,

the type species, Exputens llajasensis Clark, 1934,

was stated as fixed by original designation. In his

original description of Exputens, Clark described

two new species, E. llajasensis and E. alexi, but he

selected neither species as the type of Exputens.

The type species was subsequently designated by

Yokes, 1939, p. 51, as E. llajasensis Clark from

Middle Eocene beds from Simi Valley, California.

Palmer, 1957. 1967. has particularly emphasized

the fact that Exputens was confined to the

Tethyan marine province, and it has been reported

from Jamaica and southern California as well as

Florida. The stratigraphic range of the genus

appears to be Middle and Upper Eocene.

MacNeil described Vulsella ocalensis in 1934

from specimens collected at the Sumter Rock

Quarry about two miles northeast of Sumterville.

Sumter County. Florida. Also in the type lot was

a specimen collected from a limestone quarry

about a mile south of Newberry. Alachua County.

Florida. V. ocalensis was later allocated to the genus.

Exputens, and the outline of the valves is similar

to the type species, E. llajasensis. In his original

description, MacNeil mentioned the fact that the

shell of V. ocalensis could be composed of

lamellar calcite. In every specimen of V. ocalensis

we have seen, much of the original shell is

preserved, and tliis leads us to the conclusion that

TABLE 1. The stratigraphic range of Exputens

ocalensis in the Crystal River Formation of

Florida.

CLASSIFICATION OF THE CRYSTAL RIVER

FORMATION IN FLORIDA

{adopled from Puri,I957)

Vol. S7 (3)

THE NAUTILUS

73

the shell was composed nuiiiily of calcite. Except

for the Ostieidae and Pectinidae. almost all of the

other molluscs are preserved only as internal and

external molds and casts in the Crystal River

Limestone. The correct orientation of the valves is

questionable, but we are assuming that the beaks

and the small wing-like projection on the dorsal

border are located anteriorly. The location of the

beaks varies from nearly a central position to near

the anterior end of the dorsal border.

There are more than 40 specimens of Exputens

ocalensis in the collections at The Florida State

Museum, indicating that this species is not rare.

On one small piece of limestone, there are seven

right valves and some embedded double valves. At

some localities the riglit valve is much more

common; at others the reverse is true. This fact

indicates selective sorting by currents and wave

action.

Exputens ocalensis is most abundant in the

lowest zone in which it appears, Puri's (1957)

FIGS. 1, 2, 3. Exputens ocalensis (MacNeil). The

porous material showing around the shells is the

limestone matrix. Locality: Limestone quarry west

of old U. S. 441 at south edge of Kendrick,

Marion County, Florida, N. W. !4 Sec. 25, T. 14

S., R. 21 E.

FIG. 1. Left valve exterior, hypotype Florida State

Museum 3481, length 28.1 mm, height 16.6 mm.

FIG. 2. Left valve interior, hypotype Florida State

Museum 3482, length 27.4 mm, height 14.1 mm,

FIG. 3. Right valve exterior, hypotype Florida

State Museum 3483, length 25.0 mm, height 14.7

mm.

Spiroloculina newberryensis faunizone of the

Crystal River Limestone. It apparently is found

less frequently in the zone above, the Lepido-

cyclina-Pseudophragmina faunizone. It is doubtful

that E. ocalensis is found in any of the higher

zones of the Crystal River (see Table 1). The

Tethyan gastropod Gisortia harrisi Palmer seems to

have about the same stratigraphic distribution as

E. ocalensis because G. harrisi has been found at

nearly aU localities where E. ocalensis has been

reported. Thus, some Tethyan molluscs appeared

in Florida during the time that the lower part of

the Crystal River Limestone was being deposited

and after the Avon Park, Inglis, and Williston

Formations were laid down.

Exputens ocalensis has been found in the

following counties in surface outcrops in Penin-

sular Florida: Columbia, Suwannee, Dixie, Gil-

christ, Alachua, Levy, Marion, Citrus, and Sumter.

Exputens has been collected by U. S. Geological

Survey geologists in Mitchell and Dougherty Coun-

ties in southwestern Georgia in the Ocala Lime-

stone. More surprisingly, they have also collected

Exputens from the Castle Hayne Mad in Pender

and Wayne Counties, which are in southeastern

and east-central North Carolina. A specimen from

Wayne County is considerably larger than any

specimens of Exputens ocalensis from Florida and

may be a different species. At least one Tethyan

mollusc, Exputens, lived farther north during the

Late Eocene than had previously been supposed.

ACKNOWLEDGMENTS

We are particularly indebted to Mr. Druid

Wilson of the U. S. Geological Survey, who

showed the senior author specimens of Exputens

from Georgia and North Carolina. Dr. Katherine

V.W. Palmer of the Paleontologjcal Research

Institution sent the senior author a list of

localities in Florida where Exputens has been

collected.

LITERATURE CITED

Clark, B. L. 1934. A new genus and two new

species of Lamellibranchiata from the Middle

Eocene of California. Jour. Paleo. 8 (3):

270-272.

Hertlein, L. G. and L. R. Cox. 1969. Family

Malleidae. in Treatise on Invertebrate Paleon-

74 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

tology (R. C. Moore, editor). Part N, Vol. I,

Mollusca 6, Bivalvia. N326-N332.

MacNeil, F. S. 1934. The pelecypod genus Vulsella

in the Ocala Limestone of Florida and its

bearing on correlation. Jour. Wasliington Acad.

Sci. 24 (10): 428-431.

Palmer, K. V.W. 1957. A new Gisorda from the

Crystal River Formation, Ocala Group, of

Florida, with explanatory notes on the Tethyan

influence in the Floridian Middle and Upper

Eocene. Jour. Palaeo. Soc. India, Lucknow. D.

N. Wadia Jubilee Number. 2: 69-72.

Palmer, K. V.W. 1967. A comparison of certain

Eocene molluscs of the Americas with those of

the western Tethys. Systematics Assoc. Pub.

No. 7. Aspects of Tethyan biogeography.

183-193.

Palmer, K. V.W. and D. C. Brann. 1965. Catalogue

of the Paleocene and Eocene Mollusca of the

southern and eastern United States. Part I.

Pelecypoda, Amphineura, Pteropoda, Scapho-

poda, and Cephalopoda. Bull. Am. Paleo. 48

(218): 1-466.

Puri, H. S. 1957. Stratigraphy and zonation of the

Ocala Group. Florida Geol. Survey. Geol. Bull.

38: 1-248.

Vokes, H. E. 1939. Molluscan faunas of the

Domengine and Arroyo Hondo Formations of

the California Eocene. Ann. New York Acad.

Sci. 38: 1-246.

RECENT PUBLICATIONS

Cheatuin, E. P. and R. W. FuUington. 1971. The

Aquatic and Land MoUusca of Texas. Pt. 1.

The Recent and Pleistocene Members of the

Gastropod Family Polygyridae in Texas ($2.50).

Pt. 2 — Pupillidae and Urocoptidae (Gastropoda)

in Texas ($4.50). Bull. 1 Dallas Museum of

Natural History, P. O. Box 26193, DaUas,

Texas, 75226. Illustrated and Useful.

Ladd, Harry S. 1972. Cenozoic Fossil MoUusks

from Western Pacific Islands; Gastropods (Turri-

teUidae Through Strombidae). U. S. Geol.

Survey Professional Paper 532, 79 pp., 20 pis.

Descriptions or citations of 174 species of 18

gastropod families from seven island groups.

Excellent.

Britton, J. C. Jr. 1972. Two New Species and a

New Subgenus of Lucinidae (Mollusca: Bivalvia),

with Notes on Certain Aspects of Lucinid

Phylogeny. Smithsonian Contributions to Zoo-

logy, no. 129, 19 pp. Liicina hendersoni n. sp.

from Antigua; Pannluciua rehderi n. sp. from

Brazil, and Radiolucina n. subgenus (type:

amiantus Dull, 1901).

Sandved,K. B. and R. Tucker Abbott. 1973. Shells

in Color. 101 pis. in color. 48 pp. accompany-

ing text; bibUography; index. Viking Press, Inc.,

625 Madison Ave., N. Y., N. Y., 10022.

$12.95. A pretty book, but over-priced.

Wriglit, C. A. 1971. Flukes and Snails. Science of

Biology Series no. 4. 168 pp. George Allen and

Unwin, Ltd., London. Hardback, about $7.50.

Biology of schistosomes and planorbid and

bulinid molluscan hosts, mainly in Africa.

Saul, Louella R. 1973. Evidence for the Origin ot

the Mactridae (Bivalvia) in the Cretaceous. Univ.

Calif. Publications in Geol. Sci., vol. 97, 51 pp.,

3 pis. $4.00.

Parenzan, Pietro. 1970. Carta d'identita delle

conchiglie del Mediterraneo. Vol. 1, Gastropodi,

283 pp., 53 pis. Bio Taras Editrice, paperback

$11.00. Useful guide, but many of the 1000

ink drawings are inadequate for identification.

Kensley, Brian. 1973. Sea-Shells of Southern

Africa - Gastropods. 236 pp., 910 species illus.,

some in color. Maskew Miller, Ltd., Cape Town.

About $17.00. Contains misidentifications,

many out-of-date genera, some wrong author

names, non-binomial names, incorrect numbers

under the illustrations and a pasted-in errata

sheet containing two obsolete names. Recom-

mend waiting for second printing.

Ocean Management, Vol. 1, no. 1. A new

international, multi-disciplinary journal dealing

with the economics and exploration of oceans.

About 500 pp. per year; quarterly. Annual U.

S. subscription, $31.00. Elsevier Scientific Jour-

nals, P. O. Box 211, Amsterdam, Netherlands.

Vol. 87 (3)

THE NAUTILUS

75

NOTES ON MOLLUSKS OSTREA AND SIPHONARIA FROM GEORGIA (U. S. A.)

John N. Kraeuter

University of Georgia Marine Institute'

Sapelo Island, Georgia 31327

ABSTRACT

The sponge-dwelling oyster Ostrea permoUis Sowerby, is now known to occur off

Georgia at 20 meters depth; and the pulmonate limpet, Siphonaria pectinata (Linne)

has been found laying eggs on tree stumps on Cabretta Island, Georgia.

Recent systematic studies have illustrated how

much is still to be learned about the marine fauna

of Georgia. Darby (1965) described 13 new species

from the 24 species of podocopid ostracods he

collected. Menzies and Frankenberg (1966) des-

cribed 1 1 new species and one new genus of

isopods among the 30 species represented in the

University of Georgia Marine Institute collections.

Malacologjsts have also reported high numbers of

new species in this area. Marcus and Marcus

(1967) described three new species and a new

subspecies from the 12 opistobranch species pre-

sent in the above collections. Merrill (1963) and

Frankenberg (1966) have reported on smaD col-

lections of moDusks off Georgia. Merrill and Petit

(1965, 1969) compiled 26 and 19 range records

for moUusks from South Carolina. They reported a

ratio of one range extension for each six species

previously reported from the area. Similar ratios

are beUeved to be found in Georgia.

Range modifications for two recently studied

species are presented: one, Ostrea permollis,

Sowerby creates a continuum where a disjunct

distribution has been suggested; the other. Siphon-

aria pectinata (Linne), extends the range of this

common intertidal pulmonate beyond its northern

Umit. This paper is preliminary to a more

thorough study of the molluscan fauna of Georgia

and is intended to alert malacologjsts to the

opportunities available through the study of the

Georgia fauna.

After examination of published records and

museum specimens, Forbes (1964) concluded that

the distribution of the oyster Ostrea permollis

Sowerby was disjunct between Molasses Reef,

Florida (25° r N, 80°23' W), in 154 m, and Cape

Fear, North Carolina (33°50' N, 78°02'40" W), in

15 ra He Beheves that the absence of its host

sponge Stelletta grubii (?) Schmidt from the

continental shelf in the South CaroUna-Florida area

has isolated O. permollis populations since at least

the Sangamon Interglacial.

The discovery of O. permollis and its host

sponge Stelletta grubii (?) in Georgia emphasizes a

continuum between the Keys (Molasses ReeO and

Cape Fear. The station locations provided by

Forbes (1964) are plotted and the two collections

from Georgia are included (Fig. 1). The Georgia

specimens of O. permollis were from 31°22'55" N,

80°52.2' W, Sta. 44 & 45, M. Gray coU., 20 m,

3/7/61, 8 live individuals, and 31°24'15" N,

80°52'35" W, J. Kraeuter, coU., 20 m, 5/23/71, 2

live individuals. These lots are on deposit in the

^^;"t-^

'Contribution No. 267 from the University of

Georgia Marine Institute.

FIG. 1. Distribution of Ostrea permollis after

Forbes (1964) with Georgia data added.

76 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

collections of the University of Georgia Marine

Institute, Sapelo Island. In both cases spiculation

on the sponge fragments from the shells typify

spicules figured by de Laubenfels (1953). The

spicules measured (terminology from de Lauben-

fels, 1953):

1. Oxeas 350 - 437 ju x 10 - 21 m

2. Hagiotriane 392 - 432 m x 11.7 - 40 ^

(across head)

3. Anatriaene 392 - 422 m x 11.7 - 38 fi

(across head)

4. Eutylaster 6 - 8 /i across

5. Oxyeuaster 5 - 8 /.i across

The eutylasters were much more numerous than

the oxyeuasters and the former did not seem to

have as rough a termination as illustrated by de

Laubenfels (1953).

The presence of the host sponge and the

commensal oyster in Georgia indicates that the

lack of populations between Molasses Reef and

Cape Fear is not due to disjunct distributions, but

is due to either (1) lack of collections in offshore

waters or (2) lack of substrate for the sponge, or

(3) a combination of these factors.

The extreme narrowness of the continental shelf

near Miami obviously limits the number of

possible sites; however, submerged reefs or out-

crops have been found in North Carolina (Menzies

et al. 1966) and tliis reef is similar to those of

the Gulf of Mexico (Parker, 1956). A reef off

Georgia has been described by Hunt, Woolsey and

Henry (1971), and other reefs or "live bottom

habitats" have been listed by Struhsaker (1969)

for South Carolina, Georgia, northeast Florida and

central Rorida. It seems probable that as these

areas are sampled more extensively, fewer gaps will

exist in offshore distribution patterns.

Siphonaria pectinata, an intertidal pulmonale

limpet, has its reported northern range as eastern

Florida (Abbott, 1954) or Fernandina. Florida

(Voss, 1959). Dr. Erik Rasmussen has found it

with egg masses on tree stumps in the surf of

Cabretta Island, Georgia, adjacent to Sapelo Island.

He has been kind enougli to let me record its

northern limit and breeding during January 1972.

Tlus reproductive time is very close to that found

for the same species in south Florida by Nancy

Voss (1959). She correlated the northern limit of

S. pectinata with the 55°F isochryme on a

worldwide basis. Sapelo Island is not much beyond

the previous record of Fernandina. but the

presence here of S. pectinata suggests the northern

distributional limit may be controlled by lack of

suitable substrate on the exposed sandy shores of

the mid-Atlantic region. The large numbers and

breeding individuals found at Cabretia seem to

indicate that S. pectinata could survive even

farther north.

Historically. Dall (1927) has provided the most

conspicuous source of moUuscan daia for the

north Georgia to Florida area. In this one paper

he listed 332 species of which he considered 213

to be new. Thus, the area between 30 44" N.

79°26' W (948 m) and 30°58" N, 79°38' W (678

m) provided data which seems to be primarily

responsible for the major faunal separation at the

Georgia-north Florida boundary found by Fischer

(1961) in his analysis of diversity gradients of the

U. S. East coast. The magnitude of Dall's paper

can best be appreciated by comparing Fischer's

pelecypod and gastropod distributions. Fischer

(1961) was not able to demonstrate a clear, faunal

discontinuity of pelecypods in the north

Florida-Georgia area. It appears from his data,

however, that perhaps ten species have their

northern latitudinal limits at this point. Converse-

ly, the gastropods have approximately 160 species

reaching their northern latitudinal limits at the

Georgia-Florida boundary - a clear faunal break.

DalFs paper listed only 41 pelecypods, four new

species, but he catalogued 277 gastropod species

of which 200 were new. It appears that the

gradual loss of species with latitude as described

by Fischer (1961) for the lamellibranchs may

come closer to distributional reality than that of

the sharp zoogeographic boundary illustrated by

the Gastropoda.

The obvious influence of one major work such

as Dall's and the records of Ostrea permollis and

Siphonaria pectinata demonstrate the need for

more thorougli ecologjc and systematic work in

the Georgia-north Florida area. The importance of

hard substrates as limiting factors in marine

distributional patterns along the East coast of the

United States is also emphasized.

The critical reading and suggestions given by

Drs. Dirk Frankenberg and William Shaw are

gratefully acknowledged.

Vol. 87 (3)

THE NAUTILUS

77

LITERATURE CITED

Abbott, R. T. 1954. American Seashells. D. Van

Nostiand Co., Inc., New York, 541 pp.

Dall, W. H. 1927. Small shells from dredgings off

the southeast coast of the United States by the

United States fisheries steamer "Albatross" in

1885 and 1886. Proc. U. S. Nat. Mus. 74(18);

1-134.

de Laubenfels, M. W. 1953. Sponges from the

Gulf of Mexico. Bull. Mar. Sci. Gulf and

Caribbean 2(3): 511-557.

Darby, D. G. 1965. Ecology and taxonomy of

Ostracoda in the vicinity of Sapelo Island,

Georgia. National Science Foundation project

GB-26. Report No. 2, 59 pp. University of

Michigan.

Fischer. A. G. 1961. Latitudinal variations in

organic diversity. Amer. Sci. 49: 50-75.

Forbes, M. L. 1964. Distribution of the commen-

sal oyster, Ostrea pennollis, and its host sponge.

Bull. Mar. Sci. Gulf and Caribbean 14(3):

453-464.

Frankenberg, D. 1966. Southern limit oi Nassariits

trivittatus. The Nautilus 79(3): 89-90.'

Hunt. J. L, Jr., J. R. Woolsey. Jr. and V. J.

Henry, Jr. 1971. Geological investigation of a

submerged reef off Sapelo Island, Georgia.

Abstract Volume, Second National Coastal and

Shallow Water Research Conference, 268 pp.

Marcus, E. and E. Marcus. 1967. Some opistho-

branchs from Sapelo Island, Georgia, U. S. A.

Malacologia 6(1-2): 199-222.

Menzies, R. J. and D. Frankenberg. 1966. Hand-

book on the common marine isopod Crustacea

of Georgia. University of Georgia Press, 93 pp.

Menzies, R. T., O. H. Pilkey, B. W. Blackwelder,

D. Dexter, P. Huling and L. McCloskey. 1966.

A submerged reef off North Carolina. Int. Rev.

ges. Hydrobiol. 51(3): 393-431.

Merrill, A, S. 1963. Mollusks from a buoy off

-Georgia. The NautUus 77(2): 68-70.

Merrill, A. S. and R. E. Petit. 1965. Mollusks new

to South Carohna. The Nautilus 79(2): 58-66.

Merrill, A. S.' and R. E. Petit. 1969. Mollusks new

to South Carohna: II. The Nautilus 82(4):

117-122.

Parker, R. H. and 1. R. Curray. 1956. Fauna and

bathymetry of banks on the continental shelf,

northwest Gulf of Mexico. Bull. Amer. Assoc.

Petrol. Geol. 40: 2428-2439.

Struhsaker, P. 1969. Demersal fish resources:

Composition, distribution, and commercial

potential of the continental shelf stocks off

southeastern United States. Fishery Industrial

Res. 4(7): 261-300.

Voss, N. A. 1959. Studies on the pulmonate

gastropod Siphonaria pectinata (Linnaeus) from

the southeast coast of Florida. Bull. Mar. Sci.

Gulf and Caribbean 9(1): 84-99.

BOOK REVIEW

A SMELLER'S DIRECTORY of Clubs, Books,

Periodicals, and Dealers. By Ton} Rice. Third

Edition, 1 9 72. 64 pp. , paperback. Of Sea and

Shore Publications, P. O. Box 33, Port Gamble,

Wash. 98364. $L00.

There is no handier guide to shell dealers, shell

clubs and popular literature than this greatly im-

proved and neatly printed booklet. 88 shell clubs

are listed, including some new ones, such as the

Whitsunday SheU Club (of Queensland) and the

Wilmington Shell Club of Delaware. There is an

imposing list of shell and book dealers, and a useful

and complete list of periodicals dealing with mol-

lusks.

R. Tucker Abbott

Delaware Museum of Natural History

78 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

BOOK REVIEW

THE MOLLUSKS OF THE ARID SOUTHWEST

WITH AN ARIZONA CHECK LIST. By Joseph

C. Bequaert and Walter B. Miller. Tucson:

University of Arizona Press, pp. 271, maps.

$8.00.

A seasoned veteran wise in the ways of

traditional systematics and a younger well-versed in

newer techniques of study have combined to

summarize the mollusks of an important faunal

area where both have had extensive field experi-

ence. The result is a massive, but somewhat

disappointing, compilation of literature citations

and locality records. This book will be a fun-

damental information source for all who are

interested in the molluscan fauna of the arid

southwestern United States. Wlien combined with

the descriptions and illustrations in Pilsbry's

1939-1948 "Land Mollusca of North America",

the student of this region now has better

bibliograpliic and identification tools than for any

equivalent area of the United States.

The book presents first a general zoogeographic

analysis, then a species by species analysis of

distribution for the terrestrial species, and con-

cludes with an annotated check list of the Arizona

land and fresh water moUusks. The last two

sections contain very extensive duplication of data

and could have been condensed to good effect.

The locality lists give no clear indication in most

cases as to whether the cited record is based on

literature compilation or previously unreported

material.

Throughout the book there are many changes

proposed in synonymy and species limits. The

changes in Sonorella are taken from Miller's

unpublished thesis, while those proposed in other

genera generally are not documented. It is quite

useful to know the opinion of the authors

concerning the rank and relationships of the many

named forms in such genera as Ashmunella,

Holospira, Gastrocopta, Oiaenaxis and Vertigo, but

presentation of their evidence for changes would

have been preferable.

In the systematic portion of this work there

seems to be only a few errors and omissions.

Work in Europe indicafing the presence of sibling

species or species complexes in such groups as

Cionella, Pupilla, Columella and Vertigo, has not

been alluded to in their discussions. This is an

important omission, since "circumpolar" represen-

tatives of these genera occur in Arizona. The

section on foreign desert dwellers (pp. 98-102) is

cursory. The genus Sculptaria is misclassified in

the Endodontidae rather than the Corillidae. More

important, the Australian bulimulid Bothriembryon

is called a desert genus, while in reality it is

characteristic of the at least seasonally moist

forested areas of Southwest Australia.

The most disappointing portion of the book is

the discussion on zoogeography. It is concerned

with the classic and rather sterile problem of

delimiting faunal area boundaries (pp. 7-10) and

descriptive spatial accounts of each species (pp.

14-98). It accepts as axiomatic the now generally

discredited idea that the center of diversity for a

genus represents its center of origin (p. 20).

Dynamic aspects of biogeography are virtually

ignored, with a discussion of Sonorella (p. 21)

talking of "genetic drift" and "invasion of new

territory", whereas the area involved apparently

still is undergoing progressive drying. Consequent

restriction of ranges and increased colony isolation

is a more probably recent history. Despite the

advances in theoretical and experimental biogeog-

raphy in recent years, questions of species diver-

sity and species-area relationships are not dis-

cussed, although the mollusk-inhabited mountains

of Arizona form ideal "islands" in a zoogeograpltic

sense.

In summary, the authors are to be congrat-

ulated on producing an encyclopaedic work of

great usefulness. I shall make continual reference

to it. Unfortunately, it is not the significant work

in terms of new ideas or syntlieses that could have

been hoped for from the niixing of diverse talents.

That it is a mix and not a blend can be inferred

from divergent views of shell versus anatomy. On

page 1 5, the absence of generic or specific shell

characters is called exceptional, while on page 100

the shell is called untrustworthy for generic or

even family placement. Given the difficulties of

mollusk collecting in Arizona and the massive data

gathering needed to produce the present volume, it

is perhaps unfair of me to want something more

than usefulness.

Alan Solem, Curator of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Vol. 87 (3)

THE NAUTILUS

79

OBSERVATIONS BY POLARIZED LIGHT ON THE RADULA OF

APLEXA MARMORATA

Maria Luiza S. Mello and Mariene T. Ueta

Institute de Biologia, Universidade Estadual de Campinas,

13100 Campinas, Sao Paula, Brazil

INTRODUCTION

Many ultrastructural aspects of mollusc radulae

have been recently pointed out by scanning

electron microscope studies (Runliam, 1969;

Thomas, 1971; Kohn et al., 1972). However,

nothing concerned with the organizational state of

the macromolecules which make up the radula

teeth has been ytt estabhshed.

In the present paper observations with polarized

light were performed in radulae subjected to some

special histochemical procedures as an approach to

the discovery of the molecular arrangement of the

structure.

MATERIAL AND METHODS

Specimens of Aplexa marmorata (Guilding,

1828) (Pulmonata, Physidae) were collected in a

small artificial tank of a country house in

Campinas, Sab Paulo, BrazU. The animals were

killed in hot water (nearly 70°C), removed from

their shells, fixed in 70% ethanol and dissected.

After being extracted from their bulbs, some

radulae were treated with a 10% KOH aqueous

solution for 24 hours. They were washed, stained,

dehydrated in ethanol and mounted in Canada

balsam.

The following staining solutions were used: 1.

Congo Red saturated solution in 96% ethanol

(Barth, 1958); 2. Orange G saturated solution in

70% ethanol (Barth, 1958); 3. Xylidine-Ponceau at

pH 1.7 (Vidal, 1972); 4. Toluidine Blue at pH 4.0

(Uson, 1960).

At least five radulae were subjected to each

staining procedure.

The investigation of anisotropic properties was

carried out with a Zeiss Pol-photomicroscope

equipped with a MPM photometer and an EMI

6256 photomultipher. Histophotometric readings

were performed in different regions of the teeth

of the 1st and 2nd distal rows, oriented paraUel

(d||) and perpendicular (djj to the plane of

polarized light. A Plan 40/0.65 objective, 2.0

optovar, lOx projection lens and 2mm measuring

diaphragm were used; the area of the specimen

measured corresponds to a 2.5 /n diameter. Differ-

ent wavelengths were provided by a Zeiss mono-

chromator ruler.

The wavelength of the absorption peak was

determined for each set of measurements at the

visible spectrum. Maxima of linear dichroism (=Ad

= d|| - dj) and diochroic ratios (d||/dj) were also

calculated.

The dispersion of the birefringence (DB) was

investigated in the same material by determining

the optical path differences at several wavelengths

with a rotatory analyser and a X/4 compensator,

according to Senarmont's methods.

RESULTS

The teeth stained strongly with Congo Red and

Xylidine-Ponceau and slightly with Orange G and

Toluidine Blue.

Anisotropic phenomena are found after staining

on the basal portion of the teeth (prop and region

A) and on the denticular region (B) (Fig. 2, 3, 6,

7 and 8; Table I). Some aspects of the linear

dichroism (selective absorption of the polarized

hght) and birefringence are shown in Figures 1 to

10. The absorption values are larger when the

different regions analysed are placed parallel to the

plane of polarized light (positive dichroism), ex-

cept when the radula is stained with Toluidine

Blue (negative dichroism). When Orange G is used

as staining solution the dichroism (Ad) is very

weak and only found at the region B (Table I).

The anisotropical properties are usually less

evident at the prop region (Fig. 3 and 7).

Tlie dichroism is stronger in teeth stained with

Congo Red (Fig. 2, 3, 4 and 5; Table I) and in

this case the radicular membrane also stains red

and displays birefringence.

DB curves were determined but they did not

show inflection points within the X range used

under present experimental conditions. However,

when the radula is stained with Xylidine-Ponceau

the compensated image remains unchanged in

80 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

directions NE-SO and NO-SE at wavelengths lower

than 520nm (Fig. 1 1 - dotted line).

The dichroism and birefringence values are

slightly decreased in KOH-untreated radulae.

DISCUSSION

Optical properties such as those observed in the

present work (dichroism and birefringence) indicate

that there are oriented macromolecules in the

teeth of Aplexa mannorata suitable to bind the

dyes (Congo Red. Xylidine-Ponceau and Toluidine

Blue) in a preferential way and that this level of

molecular orientation can be measured. However,

from the fact that a precise inflection point has

not been found in the DB curves, it can be

considered that there is a very complex organiza-

tion of the stained molecules in the teeth, yeilding

inner compensations and a background effect

which would also account for the low dichroism

values. Mollusc radulae have been reported by

FIG. I. A radula stained with Congo Red (5Wnmj. x 48

FIG. 2 and 3. Details from Fig. 1. The tooth region A is oriented parallel (Fig.

3) to the plane of polarized light, x 480

FIGS. 4 and 5. Idem Fig. 2 and 3 (5l0nmj. x 185

1j and perpendicular (Fig.

Vol. 87 (3)

THE NAUTILUS

81

many authors as being composed of cliitin;

however other components are probably found in

the teeth because they stain with Xyhdine-Ponceau

at pH 1.7 and Orange G which usually attach to

protein radicals and also with Toluidine Blue

wliich generally shows dichroism when bound to

oriented polyanion substrates. Congo Red also

binds to mucins (Conn, 1967). These data suggest

that the stained substrate is probably a muco-

protein.

The staining sliglitly increases after the KOH

treatment indicating that it functions as a definite

saponification, liberating the anionic groups of the

polyanion and allowing them to become reactive.

This treatment, in addition, does not remove these

groups from their structural complex.

Present data can be considered as a starting

point for further demonstration of differences as a

function of several physiological states in the

teeth. Preliminary observations point out differ-

ences of birefringence intensity and dichroism

between the apical and basal zones of the radula,

which would mean decreasing orientation of the

macromolecules as the teeth become older and

worn.

FIG. 6. KOH-untreated radula stained with Orange G and observed with crossed polarizer and analyser, x

480

FIG. 7. The analyser was turned for compensation of region B. x 480 P = prop

FIG. 8. Compensation of the region A and of the denticles (arrow), x 480

FIG. 9. Congo Red-stained radula as seen with crossed analyser and polarizer, x 500. d = denticles

FIG. 10. Idem Fig. 9. Arrow indicates birefringence at the prop region, x 250

THE NAUTILUS

July 30, 1973

Vol. 87 (3)

TABLE 1. Polarized light extinction values and dichroism data at the wavelengths of the peaks of the absorption

curves (number of measurements = 5).

ACKNOWLEDGM ENTS

Authors are indebted to Dr. J. L. Moreira Leme

(Museu de Zoologia, Univ. S. Paulo) for the

identification of the specimens.

LITERATURE CITED

Barth, R. 1958. Metodos usados em micro-

anatomiae histologia entomologjca. Mem. Inst.

Oswaldo Cruz 56: 453-471.

Conn, H. J. 1967. Stains and staining. In Mc-

Clung's Handbook of Microscopical Technique.

Hafner Publ. Co., N. Y., pp. 72-111.

Kohn, A. J., J. W. Nybakken and J.-J. Van Mol.

1972. Radula Tooth Structure of the Gastropod

Conus imperialis Elucidated by Scanning Elec-

tron Microscopy. Science 176: 49-51.

Lison, L. 1960. Histochimie et Cytochimie Ani-

males. Paris, Gauthier-Villars.

Runhani, N. W. 1969. The use of the scanning

electron microscope in the study of the gastro-

pod radula: the radulae of Agriolimax reticu-

latus and Nucella lapillus. Malacologia 9:

179-185.

Tliomas, R. F. 1971. A scanning electron micro-

scope study of the marginal teeth of Nerita

peloronta Linnaeus. Tlie Nautilus 84: 118-119.

Vidal, B. C. 1970. Dichroism in Collagen Bundles

Stained with Xylidine-Ponceau 2R. Ann. His-

toch. 15: 289-296.

FIG. 1 1 . Curves of the dispersion of the birefringence.

P = optical path differences (nm)

Vol. 87 (3)

THE NAUTILUS

83

A NEW RECORD FOR LIGUUS: THE BOYNTON BEACH COLONY

Alan K. Craig

Department of Geography

Florida Atlantic University

Boca Raton. Florida 33432

ABSTRACT

A new and disjunct northern range limit for Liguus fasciatus has been found at

Boynton Beach, Florida. Coloration indicates affinity with extinct adjacent

populations extending southward to Ft. Lauderdale. The relict beach ridge hammock

habitat is highly restricted and destined to be destroyed.

A surprising occurrence of Liguus fasciatus in a

remnant of beach ridge hammock was noted by

university personnel in 1970 in conjuction with

routine botanical studies at Boynton Beach, Flor-

ida. Subsequent investigation by the writer has

confirmed the presence of a small colony existing

at the northernmost limit of range under some

interesting conditions not previously reported by

earlier contributors such as Moore (1921), Pilsbry

(1912. 1946). Rhoads (1899). Simpson (1921.

1922), and Young (1960).

From historic accounts (Pierce: 1970) we know

tropical hammock vegetation formerly extended in

a narrow, almost unbroken band along the crest of

beach ridges formed between Ft. Lauderdale and

Lake Worth. Further northward hammock vegeta-

tion was displaced by a more xeric dune cover

consisting of Caribbean pine and palmetto. This

habitat effectively precluded any diffusion of

Liguus in that direction. Mangrove forests and the

sawgrass environment of the Everglades created an

impassable western boundary for these arboreal

snails.

Intense real estate development has destroyed

almost every vestige of beach ridge hammock in

southeast Florida. The only example remaining

survives in an inconspicuous strip (400 x 50 yds.)

crowded between U. S. Highway AlA and the

Atlantic Ocean (Fig. 1). Thousands of residents

and tourists daily pass by this relict habitat as

they travel the busy highway or visit the public

beach.

The Boynton Beach hammock has developed a

canopy 30-40 feet in height dominated by mature

gumbo limbo, ficus, and seagrape. Many trees

show marked wind shear in response to the

prevailing southeast tradevwnd. In addition, a

survey by Austin (1972) has shown the presehce

of the plants indicated in Table 1. Trees from this

group are commonly encountered in a very mature

state, with many signs of storm damage and

general senility. The hollow trunks, cavities and

other damaged areas provide abundant estivation

sites for the Liguus colony. Shading by the upper

storey effectively precludes the growth of most

under storey plants althougli a rather conspicuous

layer of leaf mould and litter has accumulated

over much of the hammock floor.

A careful analysis of the vegetation indicates

that it has been moderately to severely disturbed

in the southern half of the hammock so that

conditions approaching a natural equilibrium state

are restricted largely to the northern half -

amounting to little more than 10.000 sq. yds.

total area. The Liguus population is further

concentrated in a protected microhabitat situated

to the leeward of the seagrape stand that occupies

the beach ridge crest.

On penetrating the hammock the abundance of

empty shells of the carnivorous Euglandina rosea is

FIG. 1. Interior of the Boynton Beach Hammock.

84 THE NAUTILUS July 30, 1973

TABLE 1. Important botanical components of the Boynton Beach Hammock.

Vol. 87 (3)

Cenchnis tribuloides

Sabal palmetto

Serenoa repens

Smilax bona-nox

Coccoloba uvifera

Yucca aloifolia

Agave decipiens

Hymenocalis latifolia

Salix caroliniana

Ficus aiirea

Nectandra coriacea

Capparis cynophallophora

Capparis flexuosa

*Forestiem segregata

Chiococca alba

Chrysobalaniis icaco

Caesalpinia bonduc

* - trees frequented by Ligiius

Source: Austin and Weise (1972)

Dalbergia ecastophyllum

Pithecellobium keyense

*Zanthoxylum fagara

Simarouba glauca

*Bursera simaruba

Cnidoscolus stimulosus

Croton punctatus

*Metopium toxiferum

Schinus terebinthifolius

Toxicodendron radicans

Knigiodendron ferreum

Eugenia axillaris

Eugenia myrtoides

Ardisia escallonioides

*Mastichodendron foetidissium

Psvchotria nen'osa

immediately noticeable. This strictly terrestrial

snail is the control predator for the arboreal

Ligiais. One has the impression that any Liguus

unfortunate enough to be blown from a tree

branch probably has little chance of returning to

safety if it should fall more than a few feet from

a tree trunk. A thorough search of the Boynton

hammock has not revealed other terrestrial snails

of substantial size that would provide suitable prey

for E. rosea so I conclude that it subsists almost

exclusively upon Liguus. Both animals are basically

nocturnal and predation has not been witnessed by

the writer although it has been reported elsewhere.

It is possible that land crabs constitute another

significant predator. Their burrows are present in

low areas of the hammock and some Z./,gwi/x shells

have been noted with damage suggestive of

pincers.

Specimens of Liguus from the Boynton Beach

hammock have certain physical characteristics that

may be especially significant. Adults are often

quite elongate (40 mm x 15 mm), exceptionally

thin-shcUed, and higlily porcellaneous on both

interior and exterior surfaces. Adult coloration

consists of two or more broad golden bands with

narrow orange borders on a white background.

These colors usually disappear beyond the penul-

timate whorl. Many specimens disclose an affinity

with the variety L. fasciatus septentrionalis (Pils.),

having three or four fine olivaceous lines super-

imposed on the gold bands. Althougli coloration as

described is not duplicated by Liguus from other

locations, we can reasonably assume that a

gradation toward olive on white formerly existed

in what was once a continuous population extend-

ing southward to Ft. Lauderdale. However, the

majority of dead shells collected from the ground

are weathered to a dull white and often much

broken.

Juveniles of a different type appear frequently.

These specimens are relatively robust and

thick-shelled. Coloration is largely brownish fiames

similar to L. marmoratus but no adults with this

coloration have been seen. If L. marmoratus does

prove to exist in the hammock together with L.

fasciatus a problem in distribution exists since the

former subspecies was never found in hummocks

to the south at Yamato, Pompano, and Ft.

Lauderdale according to Pilsbry (1946:102). Con-

sequently its disjunct presence at the extreme

northern limit of the range under natural circum-

stances seems quite unlikely. However, artificial

Vol. 87 (3)

THE NAUTILUS

85

implantation of this relatively common variety by

some well-meaning conservationist in the past is a

likely explanation. The possibility of aboriginal

involvement must not be excluded as there is

some indirect evidence of Indian presence in the

hammock.

Having found Ligiius in the densely populated

locale of Boynton Beach, further efforts were

made to seek it in some of the previously known

localities listed above by Pilsbry and indicated on

his earlier (1912) map. In every case the former

hammock habitat was completely destroyed. The

search was extended to the inland oak-palmetto

hammocks but these likewise proved barren.

Survival of Liguus in the Boynton hammock is

not by mere accident (that is, beyond preservation

of the hammock itself). Nearness to the sea with

its ameliorating effect on winter temperatures is an

obvious advantage to these cold-sensitive animals.

Higli relative humidity fosters the growth of

epiphytic algae and fungi providing abundant

forage for tree snails (see Craig 1972 for details).

Beach ridge substrate material along this part of

the southeastern Florida coast is surprisingly

siliceous (Craig et al: 1969), whereas Liguus are

known to prefer habitats having a rocky, calca-

reous substrate. In fact, the survival of a Liguus

colony in an area not having a well-indurated

substrate is difficult to demonstrate (with the

possible exception of the Cape Sable area).

Initially this lack of a rocky floor appeared to be

an anomaly of the Boynton Beach site but in the

course of investigation an outcrop of calcareous

beach ridge rock was found along part of the

western side of the hammock. Case-hardening

(secondary cementation) of this strata may have a

subtle effect on plant food intake of the vegeta-

tion and thus be reflected in the shell structure of

the tree snails.

At the time this account was being written

(Dec, 1971) the Boynton Beach hammock was

still intact. It is, however, an extremely valuable

tract from the real estate viewpoint and privately

owned. Preservation of this unique site to date

apparently is due to temporary lack of develop-

ment funds as construction of a large multi-storied

residential building was announced several years

ago.

It is unfortunate indeed that no organized

effort has been made to save this last remnant of

the primeval landscape for the benefit of future

generations who can never know the splendid

beauty of this forest once it is gone forever.

LITERATURE CITED

Austin, D. F. and J. G. Weise 1972. Annotated

checklist of the Boynton Beach Hammock.

Jour. Florida Acad. Sci. 35.

Craig, A. K. 1972. Observations on the arboreal

snail Orthalicus floridemis. Jour. Florida Acad.

Sci. 35(1): 15-20.

Craig, A. K., S. Dobkin, R. B. Grimm and J. B.

Davidson 1969. The gastropod Siphonaria pecti-

nata: a factor in destruction of beach rock.

Amer. Zool. 9(3, ed. 2): 895-901.

Moore, C. B. 1921. Liguus at Marco, Florida. The

NautUus 34: 139-140.

Pierce, C. W. 1970. Pioneer life in Southeast

Florida. Univ. Miami Press, Coral Gables, Flor-

ida. 1-264.

Pilsbry, H. A. 1912. A study of the variation and

zoogeography of Liguus in Florida. Jour. Acad.

Nat. Sci. Philadelphia 15: 429-471.

Pilsbry, H. A. 1946. Land mollusca of North

America (north of Mexico) Acad. Nat. Sci.

Phil., PhOadelphia, Pennsylvania. 2(1): 1-520.

Rlioads, S. N. 1899. Annotated list of land and

fresh-water shells recently coUected in the

vicinity of Miami, Florida. The Nautilus 13:

43-47.

Simpson, C. T. 1921. Florida West Coast Liguus.

The Nautilus 35: 66-73.

Simpson, C. T. 1922. A Search for Liguus. The

Nautilus 35: 66-73.

Simpson, C. T. 1929. The Florida Tree snails of

the genus Liguus * Proc. U. S. Nat. Mus.

73(Art.30): 1-44.

Young, F. N. 1960. Color pattern variations

among snails of the genus Liguus on the

Florida Keys. Bull. Fla. St. Mus. 5: 259-266.

86

THE NAUTILUS

July 30, 1973

Vol. 87 (3)

PISIDIUM HENSLOWANUM (SHEPPARD) IN WESTERN CANADA

Stuart A. Harris

Department of Geography

The University of Calgary, 2920 24 Ave., N.W.

Calgary, Alberta, Canada T2N1N4

ABSTRACT

New localities of Pisidium henslowanum (Sheppardj are recorded for Alberta,

British Columbia, the North-West Territories and Ontario. This species is found in

snwll numbers in boreal North America where there are alkaline oligotrophic to

somewhat entrophic waters of low to moderate salinity. Since it also occurs in

post-glacial alluvial deposits over 7000 years old, it does not appear to be

introduced species.

Collections of freshwater mollusca by the writer

from Western Canada between 1970 and 1972

have frequently yielded valves identical to Pisidium

henslowanum (Sheppard) as described by Herring-

ton (1962). Although it has been found in the

Upper Pliocene and Early Pleistocene of Idaho (U.

S. Geological Survey in Herrington, 1962, p. 41),

Herrington (loc. cit.) has concluded that "this

species appears to be introduced." Accordingly, it

would seem appropriate to list the new occur-

rences, and re-examine this conclusion. The mate-

rial is in the collection of the author and a

sampling in the Delaware Museum of Natural

History, Greenville, Delaware.

NEW LOCALITIES

Alberta

Cardinal Lake, near Peace River; ponds between

Cochrane and Calgary; Crimson Lake, near Rocky

Mountain House; Elkwater Lake, 26 iniles

south-east of Medicine Hat; Gull Lake, 10 iniles

north of Red Deer; beaver pond on Lusk creek

along the Kananaskis road, 3 miles south of Seebe;

Ma-me-o beach. Pigeon Lake, 40 miles south of

Edmonton; Vermilion Lakes at Banff; pond,

one-quarter mile inside Waterton National Park;

Lake Winnegami, near Peace River (also form

supiuum).

It is also known from the post-glacial alluvial

deposits of the area around the Porcupine Hille

(Harris and Pip. 1973).

British Columbia

Altrude Lake, Vermilion Pass, Kootenay Nation-

al Park; Lake Columbia at Canal Flats; Dog Lake,

Kootenay National Park; Kootenay Lake.

Manitoba

Lynch 's Pt., Lake Manitoba.

North West Territories:

Great Slave Lake at Hay River; latitude 63°9'

N., and longitude 122° 14' W. in terrace deposits

above the MacKenzie River. The last two samples

were collected by A. N. Boydell.

Ontario

Lake Bancroft, at Bancroft; Lake Ontario at

Hamilton.

GEOGRAPHICAL RANGE

Wlien these localities are added to those listed

by Herrington (loc. cit.), and Sterki (1916), it is

clear the Pisidium henslowanum is a fairly wide-

spread boreal species ranging from the north-east

United States almost to the Arctic Circle. It is

never very abundant since it composed 3-13% of

the valves collected at any particular site. Unless a

large collection, say, 100 valves is made, it could

easily be missed.

HABITAT AND ECOLOGY

Contrary to the statements in Herrington, this

species occurs in ponds (e.g. at Waterton), in

creeks (e.g. Lusk Creek, Kananaskis) and in small

lakes (e.g., Altrude Lake, Lake Bancroft, Cardinal

Lake and Dog Lake) in North America. The fact

that it occurs in post-glacial, alluvium beneath

Vol. 87 (3)

THE NAUTILUS

87

Mazamash (7000 years B.P.) agrees willi this

interpretation and suggests that it was hving in tire

area long before Emopean man set foot on this

continent.

The range of water quality noted so far is pH

7.2-9.4; 91-142 alkalinity (CaCO^ mg/litre): total

hardness 101-179 (CaCO^ mg/htre) 138-797 ppm.

dissolved salts; dissolved oxygen 7.7-13.4 ppm;

bicarbonate 1.97-11.06 meq. /litres; chloride

0.02-3.00 meq./litre; nitrate 0.09-0.24 meq./litre;

sulphate 0.02-1.25 meq./litre; phosphate 0.004-0.07

meq./litre; potassium 0.01-0.51 meq./litre; mag-

nesium 0.58-14.17 meq./litre; calcium 0.50-2.87

meq./litre; and sodium 0.08-8.70 meq./litre. Thus

it favors alkaline waters of low-moderate salinity

in zones of calcareous soil and rock. The water

may be oligotrophic to somewhat entrophic and

is normally characterized by dominance of the

bicarbonate anion. Since it occurs in Lake Mani-

toba, it may also survive where sulphate ions are

fairly abundant.

Presumably, this explains why it has not been

reported from the extremely low salinity waters of

central British Columbia and parts of the shield of

Northern Ontario. It has yet to be found in the

waters with higlier salinities in the south-central

Prairie Provinces.

LITERATURE CITED

Harris, S. A., and Pip, E., 1973. MoUusca as a

means of determining late and post-glacial

climatic history in Alberta. Can. Jour. Zool. in

press.

Herrington, H. B., 1962. A revision of the

Sphaeriidae of North America (Mollusca: Pelecy-

poda). Museum of Zool; Univ. of Michigan

Misc. Paper No. 118, 74 pp.

Sterki, V., 1916. A preliminary catalogue of the

North American Sphaeriidae. Ann, Carnegie

Mus., 10 {%). pp. 429-74.

CORBICULA IN SAN FRANCISCO, CALIFORNIA

James T. Carlton

California Academy of Sciences

San Francisco, California 94118

Living specimens of the introduced Asiatic

Clam, Corbicula manilensis (Philippi, 1 844) (the

same as C. fluminea of authors), were collected on

February 24, 1973, from Stow Lake, a small,

shallow, artificial body of water in Golden Gate

Park, San Francisco, California. While Corbicula is

well-known from the Sacramento-San Joaquin Del-

ta of central California (and especially in the

Delta-Mendota Canal), as well as from the South

Bay Aqueduct, this is the first record of this

exotic clam on the San Francisco Peninsula. Only

two living specimens were found (measuring ap-

proximately 17.2 mm and 17.6 mm in length.

indicating an age of perhaps one and a half years),

but several dead and smaller valves (the smallest

8.5 mm in length) collected at the same time

suggest the presence of a reproducing population.

The largest dead valve measures approximately

21.7 mm in length. Tlie clams were found in

about 12 inches of water on a muddy substrate,

along the southeastern shore of the lake. The

presence in large numbers of the native mussel,

Anodonta wahlamctensis Lea, 1838, in Stow Lake

may provide an opportunity for students to study

the effects of the introduction of Corbicula on the

mussel population.

THE NAUTILUS

July 30, 1973

Vol. 87 (3)

CYCLOSTREMELLA. A PLANISPIRAL

PYRAMIDELLID

Robert Robertson

Academy of Natural Sciences

Philadelphia, Pennsylvania 19103

The genus Cyclostremella Bush (1897) was

originally placed in the family Vitrinellidae. Tliiele

(1929) ranked it as a doubtful subgenus of

Daronia A. Adams in the subfamily Skeneinae

(family Trochidae). Defining it conchologically.

Donald R. Moore (1966) has proposed the family

Cyclostremellidae. One character setting it apart

from vitrinellids, trochaceans and most other proso-

branchs (except mathildids and architectonicids) is

its hyperstrophic protoconch.

The type-species of Cyclostremella (by original

designation) is C. humilis Bush (1897) which is

probably the most abundant minute gastropod on

the Texas coast, although rarely collected alive

(Ode, 1966-1968; Boone, 1969). Mrs. Constance

(Mollis Q.) Boone of Houston has kindly sent me

two living specimens which she found among

polychaete tubes washed ashore at Bryan Beach,

Freeport, Texas, on November 26, 1972.

Tlie external body morphology closely resem-

bles that of the five species of east American

odostomias (sensu late) with which I am familiar;

the resemblance to Odostomia dianthophila Wells

& Wells is particularly close. The pair of purplish

black eyes is posteromedian to the base of the

fairly short tentacles. (Bush also observed that

"two large black eyes [are] close together on the

top of the head"). The tentacles are ciliated and

concave ventrolaterally; the cilia near the tip are

longer than the others. A narrow mentum is

present and this can extend beyond the anterior

end of the foot, which is slightly cleft medially.

The operculum is as described and illustrated by

Bush and Moore. Behind the head there is some

grey speckling. A shallow anal sinus is present

both on the shell and the mantle edge. The

pigmented mantle organ ('? hypobranchial gland) is

greenish white anteriorly and briglit yellow and

brown to the posterior and right. All these

characters are in accord with the European

pyraniidellids studied by Fretter and Graham

(1949).

For final proofs of the pyramidellid nature of

Cyclostremella I squashed each animal under a

cover slip. No radula is present (Bush had been

unable to find one either). Instead there is an

elongate buccal stylet together with an oral sucker

at the tip of a long acrembolic proboscis (deeply

inverted in the specimens studied). C humilis is

hermaphroditic: both oocytes and spermatozoa

were observed in the gonad of one of the

specimens. The spermatozoa are long-tailed and

have corkscrew-shaped heads that are like those of

the pyramidellid described and figured by Franzen

(1955, fig. 84). Such spermatozoa are unknown

among prosobranchs (ibid., pp. 449-450).

No other known pyramidellid approaches Cyclo-

stremella in shell shape. Thus spire height per se

is not a good familial character. With its low spire,

Cyclostremella is convergent with the Architec-

tonicidae which have, however, much larger proto-

conchs (those of C humilis are only 0.28 mm in

diameter) and differ conchologically in other ways

too.

LITERATURE CITED

Boone, C. 1969. Bring 'em back alive. Texas

Conchologist 5(8): 85.

Bush, K. J. 1897. Revision of the marine

gastropods referred to Cyclostrema, Adeorbis,

Vitrinella. and related genera; with descriptions

of some new genera and species belonging to

the Atlantic fauna of America. Trans. Con-

necticut Acad. Sci. 10(3): 97-144. 10 figs., pis.

22-23.

Franzen, A. 1955. Comparative morphological

investigations into the spermiogenesis among

Mollusca. Zoologjska Bidrag fran Uppsala 30:

399-456. 146 figs., 2 pis., 2 tables.

Fretter, V. and A. Graham. 1949. The structure

and mode of life of the Pyramidellidae, para-

sitic opisthobranchs. Jour. Mar. Biol. Assoc.

United Kingdom [ser. 2] 28: 493-532. 12 figs.

Moore, D. R. 1966. The Cyclostremellidae, a new

family of prosobranch moUusks. Bull. Mar. Sci.

16(3): 480-484. 6 figs.

Ode, H. 1966. Relative abundance of shells at

Galveston. Texas Conchologist 2 (6): [8].

Ode, H. 1967. Some Galveston records of new and

unusual shells. Texas Conchologist 3(9): [6-8]

Ode, H, 1968. Cyclostremella humilis Bush 1897.

Texas Conchologist 5(4): 38-39, 1 fig.

Thiele, J. 1929. Handbuch der systematischen-

Wcichtierkunde. Jena: Fischer 1(1): 1-376, 470

figs.

Vol. 87 (3)

THE NAUTILUS

89

OBITUARY

G. Bruce Campbell, M.D.

(1934-1973)

G. Bruce Campbell was born October 14, 1934,

at Wichita, Kansas, and died suddenly, February

28, 1973, at Victorville, California. He graduated

from the school of medicine, Loma Linda Univer-

sity, in 1959, at the top of his class, and interned

at the Glendale Adventist Hospital. After one year

of residency in neurosurgery at the White Memor-

ial Hospital in Los Angeles, he transfered to a

dermatology residency at the same hospital. Since

1970, he had practiced dermatology at Victorville,

California. He married Anita Martin in 1955.

Besides his widow, he leaves three children.

As a boy. Dr. Campbell was an ardent butterfly

collector, and assembled a fine collection from the

United States and Cuba. He was also interested in

orchid culture and in raising cats for show. While

a medical student, he developed a deep interest in

marine shells of West Mexico. He made many

shore collecting, diving, and dredging trips to Baja

California, Guaymas, and Mazatlan. He was the

president of the Conchologjcal Club of Southern

California in 1962, and the vice-chairman of the

American Malacological Union-Pacific Division in

1968. For the past few years, due to health

problems, he was inactive with his collection.

However, he had recently renewed his interest, and

was planning a collecting trip to Panama at the

time of his death. He was especially interested in

the fossO and recent Typhinae and was compiling

data for a monograph of the family. He will be

deeply missed by liis many friends and especially

the writer, since we shared many collecting trips

and also shared our professional office. Trigono-

stoma campbelli Shasky, 1964, was named in his

honor.

Bibliography of G. Bruce Campbell

1961a. Range extension for Terebra ornata Gray,

1834. The Veliger, 3(4): 112.

1961b. Colubrariidae (Gastropoda) of tropical West

America, with a new species. The Nautilus,

74(4): 136-142, pi. 10.

1961c. Four new Panamic gastropods. The Veliger,

4(1): 25-28, pL 5.

1961d. Range extension of Anatina cyprinus

(Wood 1828). ibid, 4(2): 115, 3 text figure.

1962. A new deep-water Anadara from the Gulf

of California, ibid, 4(3): 152-154, pi. 37, 1 text

figures.

1963a. Rediscovery of Terebra formosa Deshayes,

1857. ibid. 5(3): 122.

1963b. A new marine research station at Guaymas,

Mexico, ibid, 5(3): 122.

1963c. (With Burch, J. Q.) Four new Olivella from

the Gulf of California. The Nautilus, 76(4):

120-126, pis. 6, 7.

1963d. (With Burch, J. Q.) A new genus for

deep-water California naticid. Proc. Malac. Soc.

London, 35: 221-225, pi. 5.

1963e. (With Burch, J. Q.) Shaskyus, new genus

of Pacific Coast Muricidae (Gastropoda). Jour.

de Conchyl. 103(4): 201-206, pL 6.

1964a. New terebrid species from the Eastern

Pacific (Mollusca: Gastropoda). The Veliger,

6(3): 132-138, pi. 17.

1964b. (With Keen, A. M.) Ten new species of

Typhinae, (Gastropoda; Muricidae). ibid, 7(1):

46-57, pis. 8-11, 3 text figures.

1964c. (With Shasky, D. R.) New and otherwise

interesting species of mollusks from Guaymas,

90 THE NAUTILUS

July 30, 1973

Vol. 87 (3)

Sonora, Mexico, ibid, 7(2); 114-120, pis. 21,

22, 1 text figure, 1 map.

Names Proposed by G. Bruce Campbell

adairensis Campbell, Terebra (Strioterebnim). 1964

altatae Burch and Campbell, Olivella (Olivellaj,

1963

anitae Campbell, Nassarina (Zanassarina). 1961

berry'i Campbell. Terebra (Strioterebnim). 1961

Calinaticina Burch and Campbell, 1963

churea Campbell, Terebra (Strioterebnim). 1964

clarki Keen and Campbell, Typhis (Typhinopsis),

1964

cortezi Shasky and Campbell. Crassispira (Stri-

ospira). 1964

crebriforma Shasky and Campbell. Clathurella (Li-

oglyphostoma). 1 964

Distichotyphis Keen and Campbell, 1964

diishanae Campbell. Terebra (Strioterebnim). 1964

eri'tlirostigma Keen and Campbell. Siphonochelus

(Sipfionochelus), 1964

fayae Keen and Campbell. Pterotyphis (Tripter-

otyphis). 1964

hyphalopilema Campbell, Amdora(Scapharca), 1962

imperialis Keen and Campbell. Typhis (Typhina).

1964

ludbrookae Keen and Campbell. Laevityphis (Laevi-

typhis). 1964

mcleani Shasky and Campbell. Calliostoma, 1964

myrae Campbell, Trivia (Piisula). 1961

ninfae Campbell. Terebra (Strioterebnim). 1961

nipponensis Keen and Campbell. Sipiionochcliis

(Siphonochelus). 1 964

precursor Keen and Campbell, Tvphis (Talitvphis).

1964

schencki* Keen and Campbell, Laevityphis (Laev-

ityphis). 1964

Shaskyus Burch and Campbell. 1963

sphoni Burch and Campbell. Olivella (Olivella).

1963

steveni Burch and Campbell. Olivella (Olivella).

1963

steveni campbelli Burch and Campbell, Olivella

(Olivella). 1963

teramachii Keen and Campbell, Typhis (Tyhpina).

1964

vemae Keen and Campbell. Distichotyphis, 1964

xavieri Campbell, Columbraria. 1961

by Donald R. Shaskv, 229 Cajon Street

Redlands. California 92373

VI VIP A R US MA L LEA TUS

IN NEW HAMPSHIRE

Frank Perron and Thomas Probert

Box 2146, Babcock Hall

University of New Hampshire

Durham, N. H. 03824

A large population of Vivipanis malleatus

(Reeve) was found on May 25, 1972, in the

Cocheco River in the town of Rochester. New

Hampshire. William J. Clench kindly verified the

identification, and we believe that this is the first

record of V. malleatus in this state. Specimens

were deposited in The Ohio State Museum,

Columbus. Ohio.

The snaUs occupy an extensive area of slack

water backed up above the Rochester town dam.

The substrate is quite sandy, but is covered with a

thick layer of algae. It has been noted that the

snail population is densest near a large trailer park

where concrete septic tanks have been built very

close to the river. Finally, these snails share the

substrate with Elliptio complanatus. Alasmidonta

undulata. Amnicola limosa, and Helisoma anceps.

Heretofore, this species has been recorded by

Abbott (1950) from San Francisco (1891); San

Jose, California (1900); Muddy River, Boston

Massachusetts. W. J. Clench (1914); Lake Quin-

sigamond, Worcester. Massachusetts (1917); Fair-

mount Park, Philadelphia, Pennsylvania (1925); St.

Petersburg. Florida (1950); near Niagara Falls, New

York (1942). The Delaware Museum of Natural

History (R. T. Abbott, in lift.) has specimens from

Putnam Pond, Fitchburg. Massachusetts, K. and R.

Wriglit, January 1973; Roosevelt Common Pond.

Tenafly, New Jersey, Alice D. Barlow. October

1968; Pine Tree Brook, Milton, Massachusetts,

Clench and R. D. Turner, May 1957; Central Park,

New York City, 1970; Sawntill Lake, Higlt Point

Park. N. J., Dorothy Fackert, 1971.

LITERATURE CITED

Abbott, R. Tucker. 1950. Snail Invaders. Natural

History 59(2): 80-85.

INFORMATION FOR SUBSCRIBERS

The annual subscription rate for The Nautilus is

$7.00 for individuals and $12.00 for institutions

(domestic or foreign). Subscriptions may begin in

January. Send check or money order to "The

Nautilus" to Mrs. Horace B. Baker, Business

Manager, 11 Chelten Road, Havertown, Pa. 19083.

Back issues from volume 72 to date are

obtainable from the Business Manager. Volumes 1

through 71 (if available) may be obtained in

reprint or original form from Kraus Reprint Co.,

Route 100, Millwood, New York 10546.

Advertising rates may be obtained from the

Business Manager or Editor.

CONTRIBUTORS

Manuscripts: Authors are requested to follow the

recommendations of the Style Manual for Biological

Journals, which may be purchased from the American

Institute of Biological Sciences, 2000 "P" Street,

N.W. Washington, D.C. 20036. Manuscripts should be

typewritten and doublespaced; original and one copy

are required, to facilitate reviews. Tables, numbered

in arable, should be on separate pages, with the title

at the top. Legends to photographs should be typed

on separate sheets. Explanatory terms and symbols

within a drawing should be neatly printed, or they

may be pencilled in on a translucent overlay, so that

the printer may set them in 8 pt. type. There is a

charge of 50 cents per word for this extra service. All

authors or their institutions will be charged 50 cents

per line of tabular material and taxonomic keys. The

pubhshers reserve the right, seldom exercised, to

charge $32 per printed page.

An abstract should accompany each paper.

Reprints and covers are available at cost to

authors. When proof is returned to authors,

information about ordering reprints will be given.

They are obtained from the Economy Printing Co.,

Inc., R. D. 3, Box 169, Easton, Maryland 21601.

MOLLUSK VOUCHER SPECIMENS

It is becoming increasingly important for future

research purposes that an identified sampling of

species mentioned in pubUcations be deposited in

a permanent, accessible museum specializing in

mollusks. This is particularly true of mollusks used

in physiological, medical, parasitological, ecological,

and experimental projects.

The Delaware Museum of Natural History has

extensive, modern facilities and equipment for the

housing and curating of voucher specimens. Mater-

ial should be accompanied by the identification,

locality data and its bibliographic reference. There

is no charge for this permanent curating service,

and catalog numbers, if desired, will be sent to

authors prior to publication.

OCTOBER 1973

THE

NAUTILUS

Vol. 87

No. 4

A quarterly

devoted to

malacology and

the interests of

conchologists

Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker.

Editors: R. Tucker Abbott and Charles B. Wurtz

EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of Mollusks

National Museum of Canada

Ottawa, Ontario, Canada K1A-0M8

Dr. William J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. 02 138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 1 0024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

Dr. James H. McLean

Los Angeles County Museum of Natural History

900 Exposition Boulevard

Los Angeles, California 90007

Dr. Arthur S. Merrill

Biological Laboratory

National Marine Fisheries Service

Oxford, Maryland 21654

Dr. Donald R. Moore

Division of Marine Geology

School of Marine and Atmospheric Science

10 Rickenbacker Causeway

Miami, Florida 33149

Dr. Joseph Rosewater

Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. Gilbert L. Voss

Division of Biology

School of Marine and Atmospheric Science

1 0 Rickenbacker Causeway

Miami, Florida 33149

EDITORS

Dr. R. Tucker Abbott

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807

Dr. Charles B. Wurtz

3220 Penn Street

Philadelphia, Pennsylvania 19129

Mrs. Horace B. Baker

Business and Subscription Manager

1 1 Chelten Road

Havertown, Pennsylvania 1 9083

OFFICE OF PUBLICATION

Delaware Museum of Natural History

Kennett Pike, Route 52

Box 3937, Greenville, Delaware 19807

Second Class Postage paid at Wilmington, Delaware

Subscription Price: $7.00 (see inside back cover)

THE

NAUTILUS

Volume 87, number 4 - October 1973

CONTENTS

R. Tucker Abbott

Acteon eloiseae, a New Opisthobranch from Arabia 91

Michael A. Rex and Kenneth J. Boss

Systematics and Distribution of the Deep-sea Gastropod, Epitonium ( Eccliseogyraj nitidum 93

Francisco J. Palacio

On the Double Hectocotylization of Octopods 99

William A. Sodeman, Jr.

The Influence of Light on Biomphalaria glabrata 1 03

Morris K. Jacobson

On the Identity of Helicina granulum Pfeiffer (Prosobranchia) 1 07

Wayne D. Cable

The Valvular Membrane in Young Mactrid Clams, Spisula solidissima 110

Kenneth W. Turgeon and Richard A. Fralick

Size and Sex Ratio Differences in Urosalpinx cinerea (Say) (Muricidae)

from Great Bay, New Hampshire 112

Willard N. Harman

The Mollusca of Canadarago Lake and a New Record for Lasmigona compressa (Lea) 114

Richard W. FuUington

Obituary of Elmer P. Cheatum (1901-1973) 115

Book Reviews

(of) Percy A. Morris, 102; Edwin A. Joyce, Jr 114

INDEX TO THE NAUTILUS

The extensive 279-page Author Index to The

Nautilus, vols. 1-75, compiled and published by

Aurele La Rocque in 1963 is still available

(unbound) for the extremely low price of $2.00

postpaid. A continuation, covering vols. 76-85, was

recently published in no. 49 of Sterkiana, 1972.

Reprints of this very useful addition are also

available for 25 cents. When ordering both of

these indispensable indices, we suggest an addi-

tional 50 cents be added to cover postage. Write:

Dr. AurSle La Rocque, 125 South Oval Dr., Ohio

State Univ., Columbus, Ohio 43210.

vu

american malacologists

presenting

an

indispensable

desk

companion

for all

mollusk

research

workers and

historians . . .

a national register of living

professional and amateur conchologists

plus biographies of 500 great,

as well as little-known,

American malacologists of the past

Here are the facts about each biog-

raphee — date and place of birth, occu-

pation, educational record, protessional

career, present position, malacological

and civic memberstiips, writings (in-

cluding titles of most important papers

and books), current mollusk research

activities, extent of shell collection,

expeditions, sources of additional bio-

graphical information, marital status,

current address — all in a clear and

compact style.

Also contains Three Major Cross-

Indexings with over 1,000 biographees

arranged (1) geographically by states

and cities, (2) by current research ac-

tivities, and (3) by occupations.

ISBN No. 0-913792-02-0

American Malacolpgists

6314 Waterway Drive

Falls Church.

Va. 22044 normally lists for $12.50

I I for institutional libraries (20% discount) $10.00

I I special PRE-PUBLICATION cost

(for unlisted persons until Dec. 1, 1973) $10.00

;|-Ja^ cj/l^// Laolnet

Successor to :

Richard E. Petit

and

John Q, Burch

SPECIMEN SHELLS

NATURAL HISTORY

BOOKS

OLIVE SHELLS

OF

THE WORLD

ROWLAND F. ZEICLEf)

HUMBERT C PORRECA

International Directory of Conchologists ■ $3.50

A current worldwide list of 2,000 people interested

in mollusks and in exchanging shells. Gives addresses ;

and interests. Available only from The Shell Cabinet. ;

Olive Shells of the World by Zeigler and Porreca - i

$12.95 :

13 full-page color plates illustrating over 150 species ;

and named varieties. The only current treatment of ;

the popular family. Clothbound. Available from your ;

book store or directly from us. 3

I Other Recently Published Shell Books Available •

I Post Office Box 29, Falls Church, Virginia 22046 !

A. Hare Coltcctor*s Item - .

SWIIHS0H

£XOm €OI€IOLMI

now available at a startlmgly low price!

Now only |/i.75 (formerly S30,00)

The few remaining copies of tliis unusual facsimile have

been purchased by the Delaware Museum of Natural History

and are now made available practically at cost to all libraries,

scientists and amateur conchologists, 48 beautiful, colored

plates, representing a faithful reproduction of Swainson's

1834 and 1841 cl.issic, together with the original text and a

modern analysis by R. Tucker Abbott and Nora McMillan.

Handsomely bound, gilt-edged on 3 sides and in full 9x12

inch page size. A useful and rare work belonging in the

library of every lover of mollusks. Both available from your

favorite book dealer or directly from the

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807, U.S.A.

VIU

Vol. 87 (4)

THE NAUTILUS

91

ACTEON ELOISEAE, A NEW OPISTHOBRANCH FROM ARABIA

R. Tucker Abbott

Delaware Museum of Natural History

Greenville, Delaware 19807

In 1970, Dr. Donald Bosch of the American

Mission Hospital in the Muscat obtained eight

specimens of an undescribed species of Acteon.

Unfortunately, soft parts were not preserved,

although one shell contained a dried operculum.

When the radulae are known it may be possible to

place properiy this species which closely resembles

members of the genus or subgenus Maxacteon

Rudman, 1971. I take great pleasure in naming

this species after the discoverer's wife, Eloise

Bosch.

Acteon eloiseae new species, Abbott

Figs. 1-4

Diapiosis—kA\Ai shell 30 to 35 mm, stoutly

ovate, solid, spirally and closely grooved, with

about 7 whorls and a low spire. Color white with

3 spiral rows of large rose-colored, crescent-shaped,

irregular splotches, each of which is bordered by a

strong black Une. Columella white, with one spiral

cord. It is most similar to Acteon flammeus

(Gmelin) but the latter is smaller, less obese, with

more numerous axial flames of color which are

lacking the black borders.

Description—ShtW 30 to 35 mm in length (1

3/8 inches), stoutly ovate, soUd, with a deep but

minute umbilicus; whorls 7 to 8; nuclear whorls 2,

smooth, white. Postnuclear whorls in the spire

with 5 to 7 deep, narrow spiral grooves which are

crossed by regular, microscopic axial threads giving

a pitted appearance. Between the grooves there are

raised, slightly rounded cords crossed by numerous

weak growth threads. Last whorl with about 30

grooves. Suture slightly channeled. Aperture ovate,

narrow above white within. Spire slightly eleVated

and about 1/3 to 1/4 the length of the shell.

Outer lip thin and sharp. Parietal callus thick and

translucent white. Columella thick, glistening white

and bearing a single, smooth, large, spiral cord

which runs back into the aperture. Umbilicus very

narrow and deep, well-grooved. Periostracum thin,

translucent tan, but thicker in the umbilical

region. Color of shell white, with 3 spiral bands of

irregular, crescent-shaped, rose to crimson blotches

which are bordered by a narrow black line. The

latter is usually wider on the adapertural side.

There are 6 to 7 blotches on the top and bottom

rows and about 11 in the middle row. Operculum

thin, chitinous, translucent amber, elongate, 14

nun in length, 6 mm in width (from a shell 31

mm in length). Nucleus indistinct and at the

broader thick end. It resembles the operculum of

Acteon tomatilis (Linn^) in shape.

FIGS. 1-3. Acteon eloiseae Abbott, new species,

Figs. 1, 2. holotype, Delaware Museum Natural

History, no. 27213. 31 mm in length. Fig. 3.

Exterior of chitinous operculum 14 mm in length.

92 THE NAUTILUS

Vol. 87 (4)

no. whorls

7.0 holotype

8.0 paratype

7.0 paratype

Types— The type locality: Al Masirali Island,

Oman Protectorate, southeast Arabia. Collected by

fishermen in shallow water near shore, 1970.

Holotype in the Delaware Museum of Natural

History, no. 27213. Paratypes in Delaware Museum

of Natural History, no. 67746, and in the

collection of Dr. Donald Bosch.

Remarks- Acteon eloiseae is a very colorful and

attractive species of unusual large size, and is

characterized by the three rows of large,

black-bordered, rose-colored blotches. Its nearest

relative, found in the Indian and southwest Pacific

Oceans, A. flammeus (GmeUn), differs in having

twice as many, narrower, pinkish rose, axial flames

of color wliich lack the black borders. Other

curious marine moUusks inhabiting the waters of

this island include Cypraea teuleri Cazenavette,

1846, and Cymatium boschi Abbott and Lewis,

1970.

american malacologists

EDITOR-IN-CHIEF

R. Tucker Abbott

duPont Chair of

Malacology,

Delaware Museum

of Natural History

a national register of living

professional and amateur

conchologists

$12.50

plus biographies of 500 great,

as well as little-known,

American malacologists of the past

FIG. 4. Acteon eloiseae Abbott, new species.

Paratype, details of sculpture and color pattern

X6. FIG. 5. Details of sculpture and color pattern

of Acteon flammeus (Gmelin, 1791) from Praia de

Chocas, Mozambique. X6. Photos by the author

American Malacologists

6314 Watprway Drive

Falls Church,

Va. 22044

Vol. 87 (4)

THE NAUTILUS

93

SYSTEMATICS AND DISTRIBUTION OF THE DEEP SEA

GASTROPOD EPITONIUM (ECCLISEOGYRA) NITIDUM

Michael A. Rex

Department of Biology, University of Massachusetts

Boston, Massachusetts 02116

and

Kenneth J. Boss

Museum of Comparative Zoology, Harvard University

Cambridge, Massachusetts 02138

ABSTRACT

Recent deep sea dredging in the western North Atlantic and South Atlantic

Oceans has yielded new specimens of the rare, open-coiled gastropod Epitonium

nitidum (Verrill and Smith). It is the type-species of the subgenus Eccliseogyra Dall,

1892, which is now removed from the subfamily Liotiinae (family Turbinidaej and

placed in the genus Epitonium (family Epitoniidaej. The synonymy and Pan-Atlantic

distribution of this species are discussed.

INTRODUCTION

In 1 885, Verrill described a fragment of an

open-coiled gastropod shell dredged by the Alba-

tross as Delphinula nitida and provisionally re-

ferred the unusual species to the prosobranch

family Trochidae. Subsequently, Dall (1889a)

assigned nitida to Laxispira Gabb, a subgenus of

Liotia Gray in the Delphinulidae. But later, after

discovering that Gabb's taxon was based on a

fossil cast or mold of Siliquaria, Dall (1892)

proposed the subgenus Eccliseogyra for nitida. A

very similar, but intact, specimen was described

from the Challenger material by Watson (1886) as

vermetiformis and placed in the genus Scalaria

Lamarck, now a synonym of Epitonium Roding

(Clench and Turner, 1951). Locard (1897 a, b)

named two additional specimens collected during

the Travailleur and Talisman expeditions as

Scalaria dissoluta, distinguishing them from S.

vermetiformis Watson on the basis of minor

differences in shell size, shape and sculpture. Bayer

(1971) first suggested that, except for the relative

height of the spire, Verrill's and Watson's species

were "close ... if not identical."

Recent deep sea dredging operations on vessels

of the Woods Hole Oceanographic Institution

(Sanders, Hessler, and Hampson, 1965) have yield-

ed numerous additional specimens of nitida (Table

1 and Figure 1). This new material was compared

with the holotype of vermetiformis Watson, the

FIG. 1. Typical specimen of E. nitidum (Verrill

and Smith) from the western North Atlantic

(Chain station 103, see Table 1). Height 6.3 mm.

94 THE NAUTILUS

October 1973

Vol. 87 (4)

paratype of dissohita Locard, additional fragments

identified by Verrill as nitida (USNM 78566), but

evidently never reported on, and Bayer's excellent

photograph (1971: Figure 11, D). It is concluded

that all belong to the same species and that

previously noted small differences between speci-

mens are aspects of intraspecific character

variation.

Keen (1960) considered Eccliseogyra Dall a

genus and placed it in the family Turbinidae of

the superfamily Trochacea. On the basis of

morphological features of its type species, Ec-

cliseogyra is here considered a subgenus of Epi-

tonium and is transferred to the Epitoniidae.

SYSTEMATICS

The following synonymy summarizes the

nomenclatorial history of this species.

Genus Epitonium Roding 1798

Subgenus Eccliseogyra Dall 1892

Eccliseogyra Dall 1892, Trans. Wag. Free Inst. Sci.

3: 307 (type species: Delphinula nitida Verrill

and Smith 1885, by monotypy).

Solutiscala de Boury 1909, Bull. Mus. Hist. Nat.,

Paris 15: 482 (type species: Scalaria

[=Epitonium] dissoluta Locard, original

designation).

Epitonium (Eccliseogyra) nitidum

(Verrill and Smith 1885)

Figs. 1-4; Table 1

Delphinula nitida Verrill and Smith 1885 [in]

Verrill, Trans. Conn. Acad. Arts Sci. 6 (2):

424-425, pi. 44, fig. 11. (Albatross Sta. 2229,

off Chesapeake Bay, 37°38'40" N; 73°16'30"W,

1423 fathoms [= 2602 meters] ; holotype in the

USNM destroyed); paratype, USNM 78566,

Albatross Sta. 2714, SW of Nantucket,

38°22'N; 70°17'30"W, 1825 fathoms [= 3338

meters] .

Scalaria vermetifonnis Watson 1886, Challenger

Repts., Zool. 15: 142, pi. 9, fig. 6. (Challenger

Sta. 122, off Pemambuco (Recife], Brazil,

09°5'S; 34°50'W, 350 fathoms [= 640 meters] ;

holotype in BMNH).

Liotia (Laxispira) nitida (Verrill [sic] ). Dall 1 889,

Bull. Mus. Comp. Zool. 18: 386 [generic and

subgeneric affinities of nitida discussed] ; Dall

1889, Bull. U. S. Nat. Mus., 37: 166, pi. 46,

fig. 1 1 [Verrill and Smith's figure of nitida

reproduced] .

Liotia (Eccliseogyra) nitida (Verrill [sic]). Dall

1892, Trans. Wag. Free Inst. Sci. 3 (2): 307

[nitida assigned to a new subgenus] .

Scalaria dissoluta (P. Fischer MS) Locard 1897.

L'Echange, Revue Linneenne, 145, p. 3; Locard

1897. Exped. Sci. Travailleur et Talisman,

MoUusques Testaces 1: 407-408, pi. 19, figs.

1-3 (Talisman Sta. 113, off Cape Verde Islands,

495-618 meters [corrected to 80-110 meters by

Smith, 1 888] ; location of figured holotype

unknown; paratype, here figured, Travailleur

Sta. 3 [1881, Atlantic Series], W of Portugal,

39°47'50"N; 9°51'45"W, 3307 meters).

Solutiscala (Solutiscala) vermetiformis (Watson).

Clench and Turner 1952, Johnsonia 2: 347, pi.

170, figs. 1-2.

Epitonium (Solutiscala,) vermetiforme (Watson).

Bayer 1971, BuU. Mar. Sci. 21: 133, fig. 11 d

(Gerda Sta. G-966, Straits of Florida SW of

Marquesas Keys, 24°10'N; 82°22'W, 544-549

meters).

REMARKS

A specimen from the new material dredged in

the western North Atlantic (Table 1) is shown in

Figure 1 . For comparison, the paratype of Scalaria

dissoluta Locard and the holotype of Scalaria

vermetiformis Watson are illustrated in Figures 2

and 3 respectively. The protoconch was described

by Watson (1886, p. 142, pi. 9, fig. 6; see also

Figure 3 this paper). It is brown in color, has four

connected whorls and is sculptured with closely

placed axial ribs. The postlarval shell was

described by Verrill (1885). It is delicate and

whitish gray in color. Whorls are circular in

cross-section and are completely disconnected.

Sculpture consists of many closely placed spiral

ribs crossed by delicate axial lamellae which

completely circumscribe the whorls. As Verrill

(1885) pointed out, the lamellae are more elevated

on the upper, inner and lower surface of the

whorls (see, e.g.. Figure 1). This fact, along with

the broken appearance of the lamellae, suggests

that they are normally more extended, but have

been damaged during dredging.

Vol. 87 (4)

THE NAUTILUS

95

Except for the relative height of the spire,

there is Httle variation among the specimens that

we have examined. The width to height ratio at

the end of the fourth whorl ranges from 0.68

(specimen in Figure 1) to 0.48 (specimen in

Figure 2).

From its general appearance, nitidum is refer-

able to Epitonium and we concur with Clench and

Turner (1952) who placed it in the subgenus

Solutiscala which deBoury (1909) estabUshed for

the uncoiled species. However, deBoury's taxon, is

a junior subjective synonym of Eccliseogyra Dall

1892 and, thus, the proper nomen for the species

is Epitonium (Eccliseogyra) nitidum.

Unfortunately, both Verrill's surviving fragment

and specimens in the new material are missing the

distinctive protoconch illustrated by Watson (1886,

pi. 9, fig. 6). Several specimens in the new

material show remains of broken eroded apices

which are smooth and lack periostracum. However,

the similarities both in the sculpture and in the

FIG. 2. Paratype of Scalaria dissoluta Locard

f'Travailleur staHon 3, off Portugal, MNHP 642).

Specimen decollated, height 5.5 mm, coated with

magnesium oxide for photography.

FIG. 3. Holotype of Scalaria vermetiformis Watson

/■Challenger station 122, off Pemambuco [Recife],

Brazil, BMNH 87.2.9.500. Specimen complete with

protoconch, height 4.8 mm, coated with

magnesium oxide for photography.

form of postlarval whorls are sufficient to justify

the synonymy of vermetiformis and dissoluta with

nitidum

Epitonium revolutum (Hedley, 1899), reported

from shallow water in Funafuti Atoll, is reminis-

cent of E. nitidum because of its open-coiled shell,

and was included in Solutiscala by deBoury

(1909). However, it can be clearly distinguished by

its less numerous and more pronounced varices.

Ladd (1972) recently noted this species from the

Late Miocene of Palau.

An unusual feature of E. nitidum is its

open-coiled or alloiostrophic (Simroth 1896-1907)

FIG. 4. The distribution of Epitonium nitidum

(Verrill and Smith, 1885) in the Atlantic Ocean.

Solid stars indicate individual records, and the

open star in the western North Atlantic represents

a composite of the first nine new records

presented in Table 1 and two previous records

from Verrill discussed in the text.

96 THE NAUTILUS

October 1973

Vol. 87 (4)

condition. Coiling in snails was described by Raup

(1966) in terms of four geometric parameters:

whorl shape (S), rate of whori expansion (W),

distance of whorls from the coiling axis (D), and

translation rate along the coiling axis (T). Yochel-

son (1971) pointed out that "open coiling may

occur if any of these components is large enough

to prevent contact of the whoris." In isostrophic

(planispiral) shells it obtains when either D > - or

W >-(Raup, 1967; Yochelson, 1971). In ortho-

D

strophic (downward spiraling) forms such as E.

nitidum it is generally achieved by having an

exceptionally high translation rate. Alloiostrophy

occurred in numerous hneages of fossU gastropods

(cf Wenz, 1938) and is widespread in Recent

fresh-water and marine prosobranchs as well as in

terrestrial and fresh-water pulmonates (Boss and

Rex in prep.). In the Vermetidae, open coOing

provides increased surface area for cementation

(Gould, 1966), but the adaptive significance of

open coiling in other cases is not clear. In

sedentary deposit, or suspension feeding forms it

might function to maintain position in the

substratum.

DISTRIBUTION

New locality records for E. nitidum are present-

ed in Table 1, and all known localities are plotted

in Fig. 4.

The horizontal distribution pattern depicted in

Fig. 4 is largely an artifact of where extensive

dredging has taken place. It is evident, however,

that the geographic range of the species includes

both the eastern and western North Atlantic, and

a Pan-Atlantic distribution appears Ukely. Un-

fortunately, there is some confusion about the

location and depth of Talisman station 113.

Locard (1897 b) variously cited this station as

being located either in the Azores or the Cape

Verde Islands at depths ranging from 495 to 1983

meters. We have followed Smith (1888) in placing

Talisman station 113 in the Cape Verde Islands at

depths of 80-110 meters (Fig. 4).

Though a comparatively rare species', nitidum

exhibits nearly the same broad geographic range as

dominant deep sea gastropod species such as

Cithna tenella Jeffreys, Adeorbis umbilicatus

Jeffreys, Mangelia bandella Dall, and Lacuna

cossmanni Locard (Rex, 1972). Madsen (1961) and

Knudsen (1970) have shown that

macro-topographic relief of the ocean floor, such

as the Mid-Atlantic Ridge, does not exert an

important influence on the distribution of many

deep sea benthic invertebrate groups. This appears

to be true of some gastropods, including E.

nitidum

In the northwestern Atlantic, E. nitidum has

been collected only between the depths of 2022

and 4693 meters (Table 1) despite extensive

dredging both above and below these depths.

However, in more equatorial regions the species'

bathymetric range extends into much shallower

waters including the upper continental slope off

Brazil {Challenger Sta. 122, 640 meters), off

Florida {Gerda Sta. G-966, 544-549 meters), and

the continental shelf in the Cape Verde Islands

{Talisman Sta. 113, 80-110 meters according to

Smith 1888). Thus nitidum has a vertical distribu-

tion ranging from the continental shelf to the

abyssal plain. Deep sea benthic invertebrate species

with wide horizontal distribution are typically

eurybathic (Vinogradova 1959), but this tendency

has not previously been reported for gastropods

because of inadequate material. It should be noted

that this type of distribution is not universal

among deep sea gastropods and that its frequency

remains to be determined.

ACKNOWLEDGMENTS

We thank W. D. Hartman (PMNH), J.

Rosewater (USNM), J. Taylor (BMNH), and M.

Testud (MNHP), for the loan of specimens from;

these institutions. This research received support

from NSF Grant GB- 19922 to Harvard University

(R. C. Rollins, principal investigator). We especially

wish to thank H. L. Sanders who kindly provided

the new material presented in Table 1 which was

collected by WHOI under NSF Grants GB-563 and

GB-15638. The manuscript was read by R. D.

Turner, R. Bullock and R. Tucker Abbott.

' Eight live specimens collected between the depths

of 2022 to 3834 meters off North America

(Table 1) comprised less than 0.7% of the

number of live gastropods dredged between these

depths during the WHOI Gay Head-Bermuda deep

sea dredging project.

Vol. 87 (4)

THE NAUTILUS

97

ABBREVIATIONS

USNM United States National Museum, Washing-

ton, D. C.

PMNH Peabody Museum of Natural History, Yale

University

WHOI Woods Hole Oceanograpluc Institution

BMNH British Museum of Natural History

MNHP Museum of Natural History, Paris

LITERATURE CITED

Bayer, F. M. 1971. New and unusual mollusks

collected by R/V John ElUott Pillsbury and

R/V Gerda in the tropical western Atlantic.

BuU. Mar. Sci. 21: 111-236.

Boss, K. J. and M. A. Rex. In prep. Alloiostrophy

in Tertiary and Recent Gastropods.

deBoury, E. 1909. Observations sur les Scalidae

des Expeditions scientifiques du Tmvailleur et

du Talisman. Bull. Mus. Natl. Hist. Nat., Paris

15: 478-484.

Clench, W. J. and R. D. Turner. 1951. The genus

Epitonium in the western Atlantic. Johnsonia 2:

249-288.

Qench, W. J. and R. D. Turner. 1952. The genera

Epitonium (Part II), Depressiscala, Cylindriscala,

Nystiella and Solutiscala in the western

Atlantic. Johnsonia 2: 289-356.

Dall, W. H. 1889a. Report on the MoUusca, Part

II Gastropoda and Scaphopoda. Reports on the

results of dredging ... in the Gulf of Mexico

(1877-78) and in the Caribbean Sea (1879-80),

by the U. S. Coast Survey steamer "Blake" . . .

Bull. Mus. Comp. Zool. 18: 492 p.

Dall, W. H. 1 889b. A preUminary catalogue of the

shell-bearing marine mollusks . . . Bull. U. S.

Nat. Mus. 37: 1-221.

Dall, W. H. 1892. Contributions to the Tertiary

fauna of Florida, with special reference to the

Miocene Silexbeds of Tampa and the Pliocene

beds of the Caloosahatchee River. Part. II.

Streptodont and other gastropods, concluded.

Trans. Wagner Free Inst. Sci. (Philadelphia) 3:

201-473.

Gould, S. J. 1966. Notes on shell morphology and

classification of the Siliquariidae (Gastropoda).

The protoconch and slit of Siliquaria squamata

Blainville. Amer. Mus. Novitates 2263: 1-13.

Hedley, C. 1 899. The Mollusca of Funafuti. Part 1

Gastropoda. Australian Mus. Mem. 3: 395-488.

Keen, M. 1960. In Moore, R. C. [ed.] Treatise on

Invertebrate Paleontology. Part I, Mollusca 1,

Univ. Kansas, Lawrence, Kansas, 351 p.

Knudsen, J. 1970. The systemarics and biology of

abyssal and hadal Bivalvia. Galathea Rept. 11:

1-241.

Ladd, H. S. 1972. Cenozoic fossil mollusks from

western Pacific islands; Gastropods (Turritellidae

through Strombidae). Geol. Surv. Prof Pap.

532: 1-79.

Locard, A. 1897a.. Notices conchyliologiques.

XLII. Scalaridae nouveaux. L'Echange, Revue

Unneenne 145: 2-3.

Locard, A. 1897b. Expeditions scientifiques du

Travailleur et du Talisman. Mollusques Testaces

1: 1-516.

Madsen, F. J. 1961. On the zoogeography and

origin of the abyssal fauna, in view of the

knowledge of the Porcellanasteridae. Galathea

Rept. 4: 177-218.

Raup, D. M. 1966. Geometric analysis of shell

coiling: general problems. Jour. Paleontology

40: 1178-1190.

Raup, D. M. 1967. Geometric analysis of shell

coiling: coiling in ammonoids. Jour. Paleon-

tology 41: 43-65.

Rex, M. A. 1972. Species diversity and character

variation in some western North Atlantic deep

sea gastropods. Unpublished Ph.D. Thesis, Har-

vard University, 178 pp.

Sanders, H. L., R R. Hessler and G. R. Hampson.

1965. An introduction to the study of deep-sea

benthic faunal assemblages along the Gay

Head-Bermuda Transect. Deep-Sea Res. 12:

845-867.

Simroth, H. 1896-1907. Mollusca. In H. G. Bronn

[ed.]. Klassen undOrdnungen des Tier-Reichs 3:

1506 p. C. T. Winter'sche Verlagshandlung,

Leipzig.

Smith, S. 1888. Lists of the dredging stations of

the U. S. Fish Commission . . . Ann. Rept. U.

S. Comm. Fish and Fisheries, for 1886:

837-1017.

Verrill, A. E. 1885. Third catalogue of Mollusca

recently added to the fauna of the New

England Coast . . . Trans. Conn. Acad. Arts.

Sci. 6: 395-452.

Vinogradova, N. G. 1959. The zoogeographical

distribution of the deep-water bottom fauna in

the abyssal zone of the ocean. Deep-Sea Res. 5:

205-208.

98 THE NAUTILUS

October 1973

Vol. 87 (4)

TABLE 1. New Records o/ Epitonium (Eccliseogyra) nitidum (Verrill and Smith, 1885) in the Atlantic

Ocean.

Cruise

Depth

in

Station Meters

Locality

Date

Number of Specimens

Watson, R. B. 1886. Report on the Scaphopoda

and Gastropoda collected by the H.M.S. "Chal-

lenger" during the years 1873-1876. Challenger

Repts., Zool. 15: 1-756.

Wenz, W. 1938. Gastropoda, Teil 1: Allgemeiner

Teil und Prosobranchia. In O. H. Schindewolf

[ed.], Handbuch der Palaozoologie 6; 1-948,

Gebriider Borntraeger, Berlin.

Yochelson, E. L. 1971. A new Late Devonian

gastropod and its bearing on problems of open

coiling and septation. In J. T. Dutro, Jr. [ed.] .

Paleozoic perspectives: A paleontological tribute

to G. Arthur Cooper: Smithsonian Contr.

Paleobiology 3: 231-241.

BOOK REVIEW

A PARTIAL BIBLIOGRAPHY OF OYSTERS,

WITH ANNOTATIONS. By Edwin A. Joyce.,

Jr. State of Florida Department of Natural

Resources, Special Scientific Report No. 34,

June 1972. Paperback, 846 pp.

Among the many useful scientific works on

marine mollusks emanating from the Marine Re-

search Laboratory in St. Petersburg, Florida, this

enormous compilation leads this year's contribu-

tions in popularity and usefulness. Joyce's biblio-

graphy now replaces the famous, but hard to

obtain, oyster reference book that J. L. Baughman

issued in 1948.

Although the emphasis is naturally on papers

dealing with commercial oysters, there is excellent

coverage in other research areas, such as on the

venerid clams, the freshwater Corbicula and

unionid mussels, scallops and gastropod predators.

Of the 4,117 references listed 45% are annotated,

a feature invaluable to those without an extensive

hbrary. The key to the Uterature appears in a

Subject Index that lists all papers dealing with one

of 55 various categories, such as Age and

Longevity; Embryology; Mariculture; Pearls; Pollu-

tion and Taxonomy. Edwin Joyce and his Florida

associates are to be congratulated and thanked for

producing this much-needed compendium.

R. Tucker Abbott

Delaware Museum of Natural History

Vol. 87 (4)

THE NAUTILUS

99

ON THE DOUBLE HECTOCOTYLIZATION OF OCTOPODS

Francisco J. Palacio'

University of Miami

Rosenstiel School of Marine and Atmospheric Science

Miami, Rorida 33149

ABSTRACT

Two cases of double unilateral hectocotylization in Incirrate octopods are

reported for the first time. The supernumerary hectocotylized arms have smaller

ligulas, calami and seminal canals, and the last few rows of suckers are reduced in

size. The extra modified arms are believed to be sexually functional Double

hectocotylization is not accompanied by modifications of the genitalia. The extra

hectocotylization is possibly caused by a sex-linked mutatiQn, and it is manifested in

a uniform proportionality.

Most male octopods have the third right arm

modified for insemination; the tip of the arm is

transformed into a spoon-shaped or flat hgula

which may bear small transverse ridges; and a velar

membrane, converted into a seminal groove, fringes

the margin of the arm terminating distally into a

conical "cheeked" calamus. In shallow water

Octpodinae the ligula is small, usually not exceed-

ing 15% of the arm length. In deeper water forms

the modified portion may attain 60 or 70% of the

arm length.

Only three monotypic genera have the third left

arm hectocotyUzed: Scaeurgus unicurrhus

d'Orbigny of worldwide distribution, Euaxoctopus

panamensis, from the Gulf of Panama (Voss,

1971), and Pteroctopus tetracirrhus (delle Chiaje)

occurring in the Atlantic Ocean (a doubtful

species, Pteroctopus witjazi was described from off

the Kamchatka Peninsula (Akimsuhkin, 1963).

In the past only two cases of double bilateral

hectocotyUzation phenomena have been reported.

One case was in Eledone cirrosa from the

Mediterranean and North Sea (Appelof, 1892),

with the third left arm also modified. An idea of

the rarity of the case was given by Morales (1958)

who examined 1,538 specimens of this common

species without finding a single example. The

other case was in Octopus briareus, from Curacao

'Contribution No. 1640 from the Rosenstiel

School of Marine and Atmospheric Science,

University of Miami.

and the West Indies, which had the second left

arm also modified (Robson, 1929).

In the course of a study of the cephalopod

fauna of Brazil, I encountered a double unilateral

hectocotylization in Octopus vulgaris from off Rio

de Janeiro, in which the fourth right arm is also

hectocotyUzed. Another double, unilateral hecto-

cotylizafion was observed by Mr. Martin Wolter-

ding during morphometric measurements of Octo-

pus selene from the Gulf of Panama, in which the

second right arm is also modified.

From my observations and the previous records,

it appears that when this unique phenomenon

occurs, the double hectocotyUzafion is manifested

in a similar pattern. Fig. 1 illustrates the modifica-

tion of the tips of the arms of Octopus selene and

vulgaris the present observations indicate a regular

pattern of proporfional modificafion of the extra

hectocotylized arm to the normally hectocotylized

arm.

The normal third hectocotylized arm is usually

shorter and thinner than the other arms. The extra

hectocotylized arm is not diminished in length,

and maintains its normal relative proportionality in

being 1.06-1.21 times longer and 1.05-1.25 times

wider than the normal arm. The ligula of the

extra hectocotyUzed arm is reduced by 71-83% of

the normal hectocotyUzed arm; the extra calamus

is more variable, being between 48.0-91.6% of the

normal arm. The Ugula/arm proportion of the

extra to the normal hectocotyUzed arms is fairly

uniform (63.0-72.0%), and the calamus/Ugula pro-

portion of the extra to the normal arm ranges

100 THE NAUTILUS

October 1973

Vol. 87 (4)

from 57.5-128.0%. The extra hectocotylized arm

has about 1.16-1.27 times as many sucker rows

than the normal arm; however, the last 19.0 -

28.0% of the suckers of the extra hectocotylized

arm are conspicuously reduced in size.

As was noted in the double hectocotylized

Eledone cirrosa and Octopus briareus, there are no

structural differences in the genitalia of Octopus

selene and O. vulgaris. Nevertheless, the seminal

canal in the extra modified arm is smaller than

that of the normal hectocotylized arm, although

the length and width of the extra hectocotyhzed

arm are larger.

The comparative development of the hecto-

cotylus and the associated organogenesis has not

been studied, but the structural modification of

the arm takes place prior to the onset of

spermatogenesis. Because of the general morpho-

logy, I beheve the additional hectocotyhzed arm is

sexually functional.

The genetic control of hectocotyhzation has not

been studied, but I assume that it is governed by

sex-linked genes, and that mutations at these loci

are possibly responsible for the double hecto-

cotyUzations reported here.

ACKNOWLEDGMENTS

I am grateful to Dr. Gilbert L. Voss, Chairman,

FIG. 1. A-B, Octopus selene, C?. A, tip of second right extra hectocotylized arm; B, tip of third normal

hectocotylized arm. C-D, Octopus vulgaris, Cf ' C, tip of third right normal hectocotylized arm; D, tip of

fourth right extra hectocotylized arm. Notice the reduction in size of the distal suckers of the extra

modified arm.

Vol. 87 (4)

THE NAUTILUS

101

Division of Biology. Rosenstiel School of Marine

and Atmospheric Science. University of Miami, for

making the specimen of Octopus selene available

to me and for the use of his laboratory and

library. Special gratitude is due to Dr. P. E.

Vanzolini. Museu de Zoologia, Universidade de Sao

Paulo, Brazil, for making the Brazilian cephalopod

collections available for study. I am indebted to

Drs. G. L. Voss and Frederick M. Bayer for

reviewing the manuscript and to Miss Susan

Stevens for the illustrations.

TABLE 1. Measurements and proportions of the normal and supernumerary hectocotylized arms fin mm.).

TABLE 2. Sizes and percentages of extra to normal hectocotylized arms (in mm.).

102 THE NAUTILUS

October 1973

Vol. 87 (4)

LITERATURE CITED

Akimishki, I. I. 1963. Cephalopods of the U.S.S.R.

Acad. Sci. U.S.S.R, Institute of Oceanology.

U.S. Dept. Commerce (translation), 233 pp., 60

text-figs.

Appellof, A. 1892. Teuthologische Beitrage IV.

Uber einem Fall von doppelseitiger Hecto-

kotylisation bei Eledone cirrosa (Lam.) d'Orb.

Bergens Mus. Aarbog, :14-15.

Morales, E. 1958. Notas sobre la morfologja del

aparato genital en Eledone aldrovandi (Raf.) =

E. cirrosa (Lamarck). Rap. Proces-Verbaux

Reunios, 14: 389-394.

Robson, G. C. 1929. On a case of bilateral

hectocotylization in Octopus rugosus. Proc.

Zool. See. for 1929: 95-97, 1 text-fig.

Voss, G. L. 1971. Biological results of the

University of Miami Deep- Sea Expeditions. 76.

Cephalopods collected by the R/V JOHN

ELLIOTT PILLSBURY in the Gulf of Panama

in 1967. BuU. Mar. Sci. 21(1): 1-34, 6 figs., 12

tbls.

BOOK REVIEW

A FIELD GUIDE TO SHELLS OF THE ATLAN-

TIC AND GULF COASTS AND THE WEST

INDIES. By Percy A. Morris. Tlnrd Edition,

edited by William J. Clench 330 pp., 8 color

pis., 68 black-and-white pis. Houghton Mifflin

Co., 2 Park St., Boston, Mass. 02107. Hard-

back, $7.95. Paperback. $4.95.

For almost three decades, Percy Morris's field

guides to shells have been a constant companion

for legions of beginning conchologists. For clarity,

compactness and inexpensiveness, his books have

led the field and have been responsible for

encouraging and serving two generations of

American shelling enthusiasts. And now a

much-improved and enlarged edition, with eight

exquisite colored plates and 68 black-and-white

plates, has been issued under the technical direc-

tion of Dr. William J. Clench. The entire format

of the book is refreshingly different and affords

much easier consulting. The nomenclature, and

especially the use of technical and popular names,

has been put on a sounder basis.

In the expanded introduction, the new class,

Monoplacophora, is included, and many valuable

classificatory explanations for the beginner are

introduced. A list of shell clubs and an infor-

mative glossary are added. With the inclusion of

several dozens of West Indian sheUs, tliis field

guide now covers just over a thousand kinds of

Western Atlantic shells.

It is a happy note that a close friend of the

late Percy Morris and a competent malacologist,

Dr. WUliam J. Clench, should shepherd this book

through the rigors of publication. Dr. Clench spent

many tedious hours in bringing the finer points of

nomenclature up to date. Curiously, he did the

very same forty years ago for his deceased mentor,

Charles W. Johnson, in 1933, when he revised and

corrected the manuscript of the now historical

"List of Marine MoUusca of the Atlantic Coast

from Labrador to Texas." Were it not for Dr.

Clench, this last monument to Mr. Morris's many

Uterary contributions would not have attained the

perfection and credit that it riglitfully deserves.

For a handy S7.95 field guide to eastern

American shells, members of the shell public will

not go wrong in adding this book to their

conchological libraries.

R. Tucker Abbott

Delaware Museum of Natural History

Vol. 87 (4) THE NAUTILUS

THE INFLUENCE OF LIGHT ON BIOMPHALARIA GLABRATA'

William A. Sodeman, Jr.

University of Arkansas, School of Medicine

Department of Internal Medicine

4301 West Markham

Little Rock, Arkansas 72201

ABSTRACT

The effect of changes in incident light intensity on the activity of albino

Biomphalaria glabrata was studied utilizing a treadmill apparatus to measure crawling

rate of the snail. The study showed that B. glabrata have a threshold for light

perception which is at least as low as 8 foot candles. When this low intensity light

is turned off the snails respond with a transient pause in crawling activity. This

response was reproducible from trial to trial and was observed with all snails tested.

With sustained activity in the absence of food snails tire. The maximum crawling

rate observed for B. glabrata was 90 cm per hour.

103

Successful maintenance of the cycle of schisto-

somiasis in a natural environment depends upon

vector snails being sited to both receive infection

from and discharge it to suitable definitive hosts.

It is not clear whether the evolution of the

host-vector-parasite relationship depends in part

upon the presence of a critical density of snails,

thus permitting fortuitous snail parasite contact, or

whether other environmental factors cue the snail's

activity resulting in favorable positioning for both

receipt and discharge of the parasite. Isolation of

environmental factors that may cue behavior is

difficult under field conditions. There have been

substantial observations on the effect of environ-

mental variables on physiologic responses of snails,

such as growth, metabolic rates, reproductive

capacity and longevity, both in the field and in

the laboratory. Behavioral responses to environ-

mental variables, particularly in a controlled

laboratory setting, have received only minimal

attention. This paper reports changes in the

activity of Biomphalaria glabrata in relation to

fiuctuations in the intensity of incident light.

MATERIALS AND METHODS

The snails employed were Biomphalaria glabrata

of the albino mutant M-line. Stocks were obtained

'This work was supported by a grant from the

National Institutes of Allergy & Infectious

Disease. AI 10292-02.

in October, 1970 from the University of Michigan

Museum of Zoology, and have been maintained by

random breeding in 40-liter glass aquaria. Four, 10

mm-diameter snails were selected and individually

isolated in 6-liter glass tanks. Snails were fed 3

times weekly with boOed dried lettuce and

commercial tropical fish food (Bio-rell, Sternco,

Harrison, New Jersey). The four test snails were

Photocell

Treadmill

E^^disc

FIG. 1. Cross section of treadmill appartus (right)

and the treadmill in % view (left). The lower disc

and lower 'A of the vertical axle were submerged.

The remainder of the unit was above the water

level. The upper disc and the light and photocell

were encased in an opaque shield to prevent

artifact caused by extraneous light.

104 THE NAUTILUS

October

maintained under cool, white fluorescent liglit for

a 12-hour light, 12-hour dark cycle. Liglit intensity

at the tank surface measured 1.400 lux (130 foot

candles). Tanks were continuously aerated through

a charcoal and glass-wool filter.

Snail activity was monitored by a treadmill

illustrated in figure 1. A horizontally oriented

plastic disc, 1 cm thick and 16 cm in circum-

ference, was balanced on a needle bearing and

connected by an axle extending above the water

surface to a second disc. The circumference of the

upper disc was marked with two wedge-shaped

masks. A beam of liglit was directed througli the

edge of the upper disc to a sensitive photo cell on

the opposite side of the disc. Rotation of the disc

in either direction resulted in a change in the liglit

passing througli the wedge-shaped mask and falling

on the photo cell. The photo cell output was

measured on a strip chart recorder (Beckman,

Fullerton, Calif.). Calibration permitted monitoring

of both direction and rate of rotation of the unit.

Snails were held in a plastic clip with the shell

aperture 3 mm away from and facing the

circumference of the submerged lower disc. On

emerging from its shell the snail's foot fell on to

the disc rim. All snails were able to crawl easily in

either direction with the shell rigidly held and

such crawling resulted in rotation of the treadmill

unit. All observations were carried out in a

photographic darkroom. Tlie snails were clamped

in the unit and then followed for a 3-hour

observation interval. With the room liglits turned

off there was no incident light in the test area.

With a single incandescent bulb turned on the

light intensity at the treadmill level was 88 lux (8

foot candles). Four ligliting patterns were tested as

follows: 3 hours of continuous liglit (L), 3 hours

of continuous dark (D), liglit on for ninety

minutes, dark for ninety minutes (L-D) and liglit

off for ninety minutes then turned on for ninety

minutes (D-L). The initial 30 minutes were

considered as a stabilization period and were

discarded for the purpose of observation. Crawling

rates in cm per hour were calculated for the 30 to

90 minutes (interval 1), 90 to 120 minutes

(interval 2), and 120 to 180 minutes (interval 3).

Each snail was observed througli 7 trials of each

light pattern. Light intensities were measured with

a Gossen Luna-Pro Cadmium sulfide type light

meter (Woodside, N. Y.).

1973

60

50

40

30

u20

Vol. 87 (4)

10

D^rk

Light

Dark Lighl Lighl D.uk

FIG. 2. Crawling rates observed during the three

periods of each lighting pattern. Dotted lines

represent the means of 7 trials for each snail. The

heavy lines represent the overall means for the

four snails.

RESULTS

Results are summarized in figure 2. Mean

crawling rates are plotted for periods 1, 2 and 3

of each ligliting pattern for each of the four snails

and an overall mean is indicated by the heavy

line. Analysis of the pattern yields the following

results. Mean rates during period 1 were higher

than the following periods. Mean rates during

period 2 were usually equal to or higlier than

those of period 3 except for the L-D lighting.

Wlien the light was turned off there was a pause

in activity expressed as a lower mean rate for

period 2 of the L-D trial. The maximum crawling

rate for Biomphalaria glabrata. 1 cm in diameter,

measured over a 10 minute intei"val was 90 cm an

hour and the maximum distance traveled during a

1 hour period of observation was 89 cm.

DISCUSSION

Three characteristics of light may be of signifi-

cance in affecting snail behavior. These are

intensity, duration and wave length, or color. A

number of investigators have commented on the

effects of the light intensity on snail behavior.

Deschiens (1957) described withdrawal of B. glab-

rata into its shell on exposure to intense liglit. A

withdrawal response on exposure to intense light

has been documented for the marine mollusk.

Vol. 87 (4)

THE NAUTILUS

105

Pleurobranchaea califomica (Davis and Mpitsos,

1971). Harry and Aldridge (1958) could not

confirm a difference in distribution of B. glabrata

between light and shade sections of the environ-

ment. Deschiensand Bijan (1956) described normal

behavior and normal fertility, fecundity and devel-

opment in B. glabrata maintained in darkness of

periods of up to 90 days. Joy (1971, a, b) studied

growth and oviposition of B. glabrata in changing

patterns of light and dark. A prevalence of

nighttime oviposition was demonstrated though the

differences between oviposition rates in the dark

and in the light narrowed as the dark interval was

shortened. When a greater portion of the light

energy was delivered in the infrared spectrum,

oviposition in the dark interval was favored.

Changes in the day length of plus or minus 2

hours did not affect the growth. Nocturnal

oviposition has been documented for other species

of gastropods, including Planorbarius conieus and

Helisoma trivolvis (Cole, 1925), Gyraulus parvus

(Krull, 1931) and Oncomelania quadrasi (Pesigan,

et al., 1958) and for the limpet Ancylus fuscus

(Clapp, 1921).

Four points are clear from the current study.

First, the threshold for Ught perception in B.

glabrata is at least as low as 88 lux (8 foot

candles). Second, B. glabrata may respond to

changes in light intensity as small as 88 lux (8

foot candles) with a change in activity. Third,

with sustained activity B. glabrata will tire.

Finally, the maximum rate of travel for B.

glabrata based on a 10 minute measurement is 90

cm per hour and the maximum observed distance

traveled on a one hour period was 89 cm.

The low levels of Hght intensity were selected

for this study to represent a better simulation of

changes in intensity in natural environments. A

number of investigators (Abbott, 1948; Kawanoto,

1952; Komiya, et al, 1959; Deschiens, 1957;

Moose and Williams, 1961-62) have documented

withdrawal reactions, or correlated behavior, of a

variety of snails when exposed to intense Ught.

While abrupt transition from deep shade to

brightly sunlit water could approach the levels

used by most of these investigators, it is more

frequent to find environmental changes of a

smaller magnitude. The fact that this snail does

respond to such low light intensities as 88 lux (8

foot candles) sets the physiologic limit of its

perception at an even lower level. Several investiga-

tors (Joy, 1971, a, b; van der Schalie and Davis,

1968; Yasuraoka, 1955; Pesigan, et al, 1958;

Clapp, 1921; Cole, 1925; KruU, 1931) reported an

effect of hght on oviposition by a variety of

molluscan species.

This study would add locomotive activity to

the Ust of behavioral alterations affected by

changes in ambient light. Several questions are

suggested by these observations, particularly the

effect of changes in light intensity when they

occur against a background level of ambient light

rather than the light to dark changes utilized here.

Also, the response to changes in Ught wave-length

would also be of interest. Such studies were

beyond the scope of the present investigation and

would entail such prolonged periods of observation

that another protocol permitting simultaneous

observation of multiple snails would probably be

necessary. In any event, changes of ambient

illumination of 88 lux (8 foot candles) to darkness

do result in measurable transient decrease in

activity of Biomphalaria glabrata. Oviposition on

the treadmill was never noted.

Snails were not fed during the trial. In the

absence of substrate, continued activity is marked

by a decrease in the rate of activity. Whether this

is a result of fatigue or whether the faUing activity

is a result of a substrate deficiency is not clear.

The cliche that our research proceeded at a snail's

pace rapidly lost favor in the laboratory; however,

it is the limiting factor in any behavioral study.

When the snail must choose among several environ-

ments, the speed with which the snail can sample

the environments controls the observation time.

Mixing times in tanks 1 meter long must be weU

in excess of 1 hour to permit vaUd observations of

B. glabrata the size with which we experimented.

Critical observation of snail activity should be an

integral part of the planning of any behavioral

study that depends upon snail migration.

LITERATURE CITED

Abbott, R. T. 1948. Handbook of medicaUy

important moUusks of the Orient and the

Western Pacific. BuU. Mus. Comp. Zool. Har-

vard. 100: 245-328.

Clapp, W. F. 1921. Eggs and young of the river

Umpet, Ancylus fuscus C. B. Adams. Boston

Soc. Nat. Hist. Occ. Papers. 5: 5-10.

106 THE NAUTILUS

October 1973

Vol. 87 (4)

Cole, W. H. 1925. Egg-laying in two species of

Planorbis. Am. Nat. 59: 284-286.

Davis, W. J. and G. J. Mpitsos. 1971. Behavioral

Choice and Habituation in the Marine Mollusk

Pleurobranchaea califomica MacFarland (Gastro-

poda, Opisthobranchia). z. vergl. Physio. 75:

207-232.

Deschiens, R. 1957. Sur la perpetuation des

elevages des mollusques vecteurs des bilharzioses

a Tobscurite. Bull. Soc. Path. Exot. 50:

229-233.

Deschiens, R. and H. Bijan. 1956. Comportement

d'elevages de mollusques vecteurs des bilhar-

zioses a I'obscurite. Bull. Soc. Path. Exot. 49:

658-661.

Harry, H. W. and D. V. Aldrich. 1958. The

ecology of Australorbis glabratus in Puerto

Rico. BuU. WHO. 18: 819-832.

Joy, J. E. 1971. The Influence of Light Condi-

tions upon the Egg-Laying of the Planorbid

Snail, Biomphalaria glabrata. The Nautilus 85:

43-49.

Joy, J. E. 1971. Tlie influence of day length upon

the egg-laying of Biomphalaria glabrata, Ann.

Trop. Med. and Parasit. 65: 573-578.

Kawanoto, S. 1952. On the photophobotaxis of

Oncomelania nosophora. Med. Biol. 23: 76-79.

Komiya, Y., K. Kuniko, and C. koyama. 1959. A

simple breeding method for Oncomelania using

a Petri dish. Jap. Jour. Parasitol. 8: 721-724.

KruU, W. H. 1931. Importance of laboratory-raised

snails in helminthology with life history notes

on Gyraulus parvus. Mich. Univ. Mus. Zool.

Occ. Papers. 10(226): 1-10.

Moose, J. W. and J. E. WiUiams. 1961-62. Medical

General Laboratory (406), U, S. Army Medical

Command, Japan, Professional Reports.

Pesigan, T. P., N. G. Hariston, J. J. Jauregui, E.

G. Garcia, A. T. Santos, B. C. Santos, and A.

A. Gesa. 1958. Studies on Schistosoma japon-

icum infections in the Phillippines. 2. The

moUuscan host. Bull. WHO. 18: 481-578.

van der Schalie, H. and G. M. Davis. 1968.

Culturing Oncomelania snails (Prosobranchia:

Hydrobiidae) for studies of Oriental schisto-

somiasis. Malacologia 6: 321-367.

Yasuraoka, K. 1955. The behavior of Oncomelania

nosophora, the intermediate host of Schisto-

soma japonicum, to light in water. Jap. Jour.

Med. Sci. Biol. 8: 323.

INDO-PACIFIC

MOLLUSCA

MONOGRAPHS OF THE MARINE MOLLUSKS OF

THE WORLD WITH EMPHASIS ON THOSE OF

THE TROPICAL WESTERN PACIFIC

AND INDIAN OCEANS

The most technical and most beautifully illustrated

journal now being published on Recent and Tertiary

marine moUusks. Over 20 professional malacologists are

currently contributing. Edited by R. Tucker Abbott.

Among the groups treated are Strombidae, Cassidae,

Tridacnidae, Turridae, Littorinidae, Phasianellidae, and,

soon to come, Patellidae, Harpidae and Mitridae.

Issued to date in looseleaf form with three sturdy,

permanent binders — 1100 pages, 810 plates (31 in full

color). Limited number of complete sets left, $91.90 U.S.

(foreign: $94.00), postage paid. Any number of extra

binders available at $6.00.

Published by

The Delaware Museum of Natural History, Box 3937, Greenville, Delaware 19807 U.S.A.

Vol. 87 (4)

THE NAUTILUS

107

ON THE IDENTITY OF HELICINA GRANULUM PFEIFFER

(PROSOBRANCHIA)

Morris K. Jacobson

American Museum of Natural History

Central Park West at 79th Street

New York, N. Y. 10024

ABSTRACT

Helicina granulum Pfeiffer, 1864 is shown to be a valid species, differing in

important morphological respects from H. tantilla Pilsbry, 1902. The former taxon

has been taken in several localities in the south central part of Oriente Province,

Cuba. The use of subgenus Poenia Adams & Adams, 1856 for H. granulum is shown

to be erroneous.

The taxon Helicina granulum Pfeiffer (ex MS,

Gundlach), 1864, has not been clearly understood

in the Uterature. Pfeiffer described the species on

the basis of a single specimen sent him from

Monte Toro, Guantanamo, Oriente Province, Cuba,

by Gundlach. He was moved to do this by the

unusually small size of the shell, smaller than any

known helicinid at that time. The original Latin

description and my English translation follow:

Helicina granulum Pfeiffer (1864, Malak. Blatt.

11: 161): "T. minuta, depressa, tenuiscula, laevi-

gata, translucida, succinea; spira parum conoi-

deo-elevata; anfr. fere 4 convexiusculi, ultimus

depresso-rotundatus, subtus medio subexcavatus,

levissime callosus; columella brevissima, verticahs;

apertura obhqua, late semiovalis; perist, breviter

expansum, margine basali arcuatin in columellam

transuente -- Operc? -- Diam maj. 2 2/3, alt. vix 1

mill. Habitat Monte Toro in districto Guanta-

namensi insulae Cubae."

"Shell minute, depressed, quite thin, smooth,

translucent, amber-colored; spire little conic-ele-

vated; whorls almost 4, rather convex, the last one

depressed rounded, subexcavated centrally below,

Ughtly callused; columella exceedingly short, verti-

cal; aperture oblique, widely semi-oval; peristome

briefly expanded, basal margin inserting arc-like

into the columella. Operculum? Major diameter 2

2/3 mm, altitude barely 1 mm."

It might be noted that Pfeiffer did not mention

the presence of axial sculpture, but this omamert-

tation could have been overlooked in a single dead,

minute specimen. However, as wall be seen later,

the references to an expanded peristome and an

arched basal insertion clearly point to granulum,

Pfeiffer's holotypic specimen was never figured

and was destroyed during World War II (Clench &

Jacobson, 1971a: 101). There are few subsequent

references to the species in the hterature: Pfeiffer

(1865: 233) reprinted his original description

verbatim and later (1876: 271) merely listed the

name. Sowerby (1866: 296) listed it as an

unidentified species, and Reeve (1874) and Wagner

(1910) omitted it entirely; nor did Fulton (1915)

FIG. 1. Lucidella granulum (Pfeiffer) between

Nimfitas and La Victoria, Monte Toro, Guanta-

nomo, Oriente Province, Cuba; Ramsden, leg. 1914,

ANSP 110769, 10 X.

108 THE NAUTILUS

October 1973

Vol. 87 (4)

mention it among the names omitted by Wagner

in his extensive monograph of the Helicinidae.

Arango (1879), Crosse (1890), and Aguayo &

Jaume (1948) repeated only the locaUty cited

originally by Pfeiffer, thus leading to the supposi-

tion that Pfeiffer's record is the only one known,

and that the species had not been collected again.

Clench & Jacobson (1971b) tentatively included it

in the genus Troschelviana, although Pfeiffer had

made no reference to a mucronate protoconch, a

diagnostic feature of the helicinine tribe Vianini in

which the genus Troschelviana belongs. There is no

reason to doubt that granulum belongs to the

genus Lucidella Swainson, 1840, as was noted by

Aguayo & Jaume (1948). However, their sub-

generic placement in Poenia H. & A. Adams,

1856, is erroneous. As the type-species of Poenia

they cited the subsequent designation by Fischer,

1885 of H. unidentata Pfeiffer. But since Fischer

did not use the word "type" in his citation, this

must be regarded as an example and is not a valid

designation. Pilsbry (1911, p. 585) vahdly desig-

nated Helicina depressa Gray, 1825, as the type.

Furthermore, Poenia has been regarded as a

synonym of Lucidella by both Thiele (1929: 88)

and Wenz (1938: 445). Because of the axial

sculpture, granulum belongs in the subgenus

Poeniella H. B. Baker, 1923, type species Helicina

plicatula christophori Pilsbry, 1897, by original

designation.

The vagueness regarding the identity of granu-

lum increased when it was confused with another

species, Helicina tantilla Pilsbry, 1902. This species

was described on the basis of a single specimen

taken near Palm Beach, Florida in 1899 together

with other small land shells in a forest about

one-quarter mile from the beach. Pilsbry wrote

that the new species was distinct from any other

Antillean helicinid. He did not compare it with

the description of granulum, which it closely

resembles, nor, as we have seen, were any

typological specimens of that species available for

study. Thus when Ramsden in 1914 sent him

some small shells from Monte Toro, Pilsbry

identified them as H. tantilla, overlooking the fact

that Monte Toro is the type locality of H.

granulum I have been able to examine these

specimens in the ANSP and they clearly differ

from tantilla and are undoubtedly topotypes of H.

granulum The specimen here illustrated is, there-

fore, a potential neotype.

Tlie differences between tantilla and granulum

are the following: the peristome of tantilla is

rounded and faintly thickened, but barely expand-

ed and not reflected, and the basal angle of

insertion in the body whorl is barely perceptible.

In granulum the hp is well expanded and slightly

reflected and the angle at the point of insertion is

quite distinct. In addition, the spire of granulum is

somewhat higher and the suture deeper.

Ramsden reported that he found the shell "in

dirt at the root of trees." Ramsden's description

of the supposed "tantilla" (1914: 50) clearly

refers to granulum since reference is made to an

expanded and reflected Up and a shallow notch at

the basal insertion.

There are several lots of granulum in the

Ramsden Collection labelled as Lucidella rugosa

Pfeiffer and coming from the following locahties:

Florida Blanca, Alto Songo; Km. 54, Sierra Canas-

ta, Santiago de Cuba; La Cueva, Diego Cobas,

Majaguabo, all from Oriente Province. In addition

the USNM has one lot from Guaso River, Monte

Libano, Guantanamo, and both the USNM and

ANSP have one lot each from the type-locality,

Monte Toro, Guantanamo.

Lucidella tantilla has also been reported from

Cuba in Cayo GuUlermo, northern Camagiiey

Province (Aguayo & Jaume, 1948) and from Cayo

Las Brujas, Caibarien, Las Villas Province (Clench

1959: 35). In addition. Clench records it from

Mores, Great Abaco, Bimini, and Great and Little

Inagua Islands in the Bahamas, and Pilsbry (1948:

1085) reports it from several localities in the

Florida Keys, as well as erroneously from Monte

Toro.

It is thus clear that Lucidella granulum is

confined to south central Oriente Province, while

L. tantilla occurs in the more western provinces' of

Camagiiey and Las Villas. From here tantilla

appears to have dispersed to the Bahamas and the

Florida Keys, since Pilsbry was most likely correct

when he wrote that the Florida specimen was

"probably a hurricane-borne waif from Cuba"

(1948).

I wish to express my gratitude to Dr. William

K. Emerson of the American Museum of Natural

History who discussed the problem with me and

critically read the manuscript. Dr. Robert Robert-

son of the Academy of Natural Sciences and Dr.

Vol. 87 (4)

THE NAUTILUS

109

Joseph Rosewater of the National Museum of

Natural History, Smithsonian Institution, most

generously permitted me to examine material in

their respective collections. I am indebted to the

kindness of Dr. Kenneth J. Boss for the photo-

graphs. The abbreviations ANSP and USNM stand

for the Academy of Natural Sciences of Philadel-

phia and the National Museum of Natural History,

Smithsonian Institution (formerly the United

States National Museum) respectively.

LITERATURE CITED

Aguayo, C. G. and M. L. Jaume, 1948. Cat&logo

molluscos de Cuba, HeUcinidae, no. 495, 1 p.

mimeo.

Arango, R., 1879. Contribucion a la fauna mala-

col6gica cubana, Helicinidae, Havana, pp. 41-57.

Baker, H. B., 1923. The Mollusca collected by the

University of Michigan - Williamson Expedition

in Venezuela. Occ. Papers Mus. Zool. Univ. of

Michigan, no. 137, 58 pp., 5 pis., 21 figs.

Qench, W. J., 1959. Land and freshwater mollusks

of Great and Little Inagua, Bahama Islands.

Bull. Mus. Comp. Zool., Cambridge, Mass.,

121(2): 29-54, 1 pi.

Clench, W. J. and M. K. Jacobson, 1971a. Mono-

graph of the Cuban genera Emoda and Glyp-

temoda. Bull. Mus. Comp. Zool. 141(3):

99-130, 7 pis.

Clench, W. J. and M. K. Jacobson, 1971b. A

monograph of the genera Calidviana, Ustronia,

Troschelviana, and Semitrochatella in Cuba.

ibid., 141(7): 402-463, 8 pis., 1 text fig.

Crosse, H., 1890. Faune malacologique terrestre et

fluviatile de I'ile de Cuba. Jour. Conchyl. 38:

173-335, pis. 4-6.

Fischer, P., 1885. Manuel de Conchyliologie, Paris,

pp. 794-797, figs. 554-556 (Helicinidae).

Fulton, H. C, 1915. On Dr. Anton Wagner's

monograph of Helicinidae in the Conchy-

lien-Cabinet. Proc. Mai. Soc. London, 11:

237-241; 324-326.

Pfeiffer, L., 1864. Zur Molluskenfauna von Cuba.

Malak. Blatt. 11: 157-161.

Pfeiffer, L., 1865. Monographia Pneumono-

pomorum Viventium, suppl. 2, 284 pp.

Pfeiffer, L., 1876. ibid., suppl. 3, 10 + 479 pp.

Pilsbry, H. A., 1902. A new Floridian Helcina,

Nautilus 19: 53.

Pilsbry, H. A., 1948. Land Mollusca of North

America. Monograph 3, Acad. Nat. Sci. Phila-

delphia 2(2): 1078-1090, figs. 576-581 (HeUcini-

dae).

Pilsbry, H. A. and A. P. Brown, 1911. The Land

Mollusca of Montego Bay, Jamaica; with Notes

of the Land Mollusca of the Kingston Region.

Proc. Acad. Nat. Sci. Phila. for 1911, p. 585.

Ramsden, C. T., 1914. Notes on some land shells

of eastern Cuba. The NautUus 28: 49-51, pi. 2,

Reeve, L., 1874. Conchologia Iconica, 19: 34 pis.

-H text (Helicinidae).

Sowerby, G. B. II, 1866. Thesaurus Conchyliorum,

3: 277-302, pis. 266-278 (Helicinidae).

Thiele, J., 1929. Handbuch der systematischen

Weichtierkunde, Jena, 1: 80-91, figs. 57-67

(Helicinidae).

Wenz, W., 1938. Gastropoda [in] Handbuch der

Palaozoologie, BerUn, pp. 435-448, figs.

1061-1118 (Helicinidae).

Wagner, A. J., 1910. [in] Martini and Chemnitz,

Conchylien-Cabinet (2) 1: sect. 18, pt. 2, pp.

337-351, pis. 67-69. (Lucidella)

110 THE NAUTILUS

October 1973

Vol. 87 (4)

THE VALVULAR MEMBRANE IN YOUNG

MACTRID CLAMS, SPISULA SOLIDISSIMA

Wayne D. Cable

National Marine Fisheries Service

Middle Atlantic Coastal Fisheries Center

Milford Laboratory

Milford, Connecticut 06460

ABSTRACT

Juvenile surf clams, Spisula solidissima, were reared from fertilized eggs in the

laboratory. Observations were made of the appearance and activity of the valvular

membrane (primary exhalant siphon), which has not previously been reported in

recently-metamorphosed animals.

The siphonal system of the bivalve mollusks has

been considerably studied. Aspects of mantle

fusion in the Lamellibranchia and the subsequent

formation of siphons from the mantle folds were

described in detail by Yonge (1957). Development

of siphons in the evolution of the lamellibranchs

enabled this group of animals to live in the shelter

of a substrate while obtaining food and oxygen

from the water above the substrate (Yonge, 1957).

In discussing the types of siphon systems,

FIG. 1. Juvenile Spisula solidissima with the

valvular membrane, or exhalant siphon, (on the

left) fully extended to a length of 1.5 mm.

FIG. 2. Juvenile Spisula solidissima, valvular mem-

brane (on the right) collapsed but not inverted

within shell.

Vol. 87 (4)

THE NAUTILUS

111

Yonge ascribes to the Mactridae what he terms a

"type C" siphon system. In this type the opening

of the definitive exhalant siphon is usually bound-

ed by a valvular membrane (a narrow flange just

inside the ring of tentacles) which, when open,

controls and directs the flow of water from the

exhalant siphon. However, he does not illustrate

this structure in any of the Mactridae, nor does he

show how this structure is formed in the young of

the Mactridae. Prior to development of the

definitive siphonal system in many recently meta-

morphosed Lamellibranchia a precursor of the

definitive exhalant siphon, a valvular membrane or

primary exhalant siphon is formed (Quayle, 1952;

Carriker, 1961).

The presence of the valvular membrane in

newly metamorphosed Spisula solidissima has been

recenfly observed by the author. There was no

question as to the identity of these young surf

clams since they were reared from fertilized eggs

spawned in the laboratory. The presence of this

structure and its appearance may serve as an aid

to identification of young juvenile surf clams

gathered in the field.

Photographs were made approximately one

month after metamorphosis; the size range of the

young clams then ranged from 0.5 mm to 3.0

mm. Photograph #1 shows the siphons of a young

clam 1.5 mm long. Here the valvular membrane is

fully extended. Photograph #2 shows the valvular

membrane in a state of collapse and dangling

externally; this clam is also 1.5 mm long.

One should notice that the state illustrated in

these photographs is an advanced one, in that

juvenile Spisula solidissima of this size have

already developed the inhalant siphon with its ring

of tentacles, and the tentacles of the exhalant

siphon have also begun to form. These structures

are formed subsequent to the development of the

valvular membrane.

The valvular membrane was observed to be an

extremely flexible and active structure. It was

rapidly extended when the young clam was

pumping water through the mantle cavity, and

rapidly withdrawn or inverted within the mantle

cavity when the clam was disturbed.

The presence of such a "filmy membrane" on

the excurrent siphon on young clams, which

moved in and out with a folding motion and was

held out as a hose to direct flow of water from

the exhalant opening, was described in Mercenaria

mercenaria by Belding (1912). Carriker (1961) has

subsequently described in some detail the develop-

ment of this membrane in young Mercenaria

mercenaria ranging from 210 to 300 ^.

The valvular membrane of recently meta-

morphosed bivalve moUusks appears to function to

deflect the excurrent and incurrent streams during

the period of definitive siphon formation. The

same function would be served in later life by the

shght offset of the tips of the siphons and the

smaller exhalant siphon orifice (Carriker, 1961). In

describing a similar membrane in young Mya

arenaria and Venerupis pullastra, Quayle (1952)

reported the function of this structure to be one

of keeping the mantle and gills free of feces by

directing them away from the vicinity of the

young animal.

Since siphon formation takes place considerably

after metamorphosis and the loss of the velum,

there is a sedentary phase prior to the time when

the siphons are formed and the animal is able to

enter the substrate (Carriker calls this the byssal

plantigrade). He believes it probable that young

byssal plantigrade Mercenaria mercenaria cannot

burrow until the ring of tentacles is available on

the inhalant siphon to exclude sedimentary grains.

This is most likely true for other lamellibranchs

that also live in soft substrates, including the

Spisula solidissima described here.

LITERATURE CITED

Belding, D. L. 1912. Report on the Quahaug and

Oyster Fisheries of Massachusetts, Common-

wealth of Massachusetts, Boston.

Carriker, M. R. 1961. Interreladon of Functional

Morphology, Behavior, and Autecology in Early

States of the Bivalve Mercenaria mercenaria.

Journal of the Elisha Mitchell Scientific Society

77 (2): 168-241.

Quayle, D. B. 1952. Structure and Biology of the

Larva and Spat of Venerupis pullastra (Mon-

tagu). Trans. Roy. Soc. Edinburgh 62 (8);

255-297.

Yonge, C. M. 1957. Manfle Fusion in the

Lamellibranchia. Pubbl. Staz. Zool. NapoU 26:

151-171.

112 THE NAUTILUS

October 1973

Vol. 87 (4)

SIZE AND SEX RATIO DIFFERENCES IN UROSALPINX CINEREA

(SAY) (MURICIDAE) FROM GREAT BAY, NEW HAMPSHIRE'

Kenneth W. Turgeon

and

Richard A. Fralick

Jackson Estuarine Laboratory, University of New Hampshire

Adams Point, Durham, New Hampshire 03824

ABSTRACT

Three hundred and ninety-seven Urosalpinx cinerea from Great Bay, New

Hampshire, were sexed and measured for total shell length. Statistical comparison of

the mean shell lengths and chi-square analysis of sex ratios for several size classes

show that females are significantly larger than males. Hypotheses to account for this

sexual dirmorphism are presented.

INTRODUCTION

Previous investigators have noted in several

populations of the common oyster drill, Uro-

salpinx cinerea (Say), that females attain a greater

maximum shell length than males (Federighi,

1931a, 1931b; Cole, 1942; Hargis and MacKenzie,

1961). However, none of these workers demon-

strated statistically significant differences in mean

shell lengths between the sexes. Griffith and

Castagna (1962), working with Urosalpinx cinerea

folly ensis (Baker, 1951) from Chincoteague Bay,

Maryland, showed that females, in addition to

having a greater maximum shell length, had a

mean shell length significantly greater than males.

Our paper reports the results of a similar study

conducted on U. cinerea from Great Bay, New

Hampshire.

Great Bay is a shallow, highly turbid estuary

with a mud-silt bottom. Yearly salinities range

from approximately seven ppt during the spring

freshet to 30 ppt in late summer in the mid and

lower reaches of the Bay. The drills are limited in

their distribution to the few, remaining oyster

reefs and are present in very low abundance. Drill

density is approximately one to two individuals

per square meter of oyster reef. However, they are

often distributed in widely separated clusters of

four to seven individuals.

METHODS

Approximately 500 oyster drills were

hand-collected during the summer of 1972 by

'Jackson Estuarine Laboratory Scientific Contri-

bution No. 1.

scuba divers. The collection site was an oyster reef

located in the mid-portion of Great Bay encom-

passing an area of about 30,000 square meters.

The divers collected all drills sighted. Trapping

techniques were not employed since bias for a

particular size or sex might have resulted. Griffith

and Castagna (1962) found that male:female ratios

differed between trap-collected and hand-collected

drills. The low population density of the Great

Bay drills and the Irigh turbidity of the water

necessitated numerous dives in order to collect

sufficient numbers for statistical analyses. Only

individuals greater than 10 mm in length and

exhibiting a complete lack of a penis (females) or

presence of a well-defined, C-shaped penis (males)

were used. Smaller individuals were not used since

they are extremely difficult to sex and quite often

have not reached sexual maturity. Sexing was done

by the live method of Hargis (1957) using 5 X

and 10 X hand magnifiers. Shell length was

measured to the nearest 0.1 mm with vernier

calipers. Drills, after being sexed and measured,

were assigned to one of four, pre-determined size

classes: >\0 & < 15 mm (class 1), > 15 & < 20

mm (class 2), > 20 & < 25 mm (class 3) and >

25 mm (class 4). These size classes were arbitrarily

chosen and do not represent natural breaks in size

distribution. Statistical comparison of female and

male mean shell lengths was by Cochran's approxi-

mation of the Behrans-Fisher test (Snedecor,

1956). Chi-square analysis was used to determine

significant deviations from 50-50 sex ratios for the

individual size classes and the total sample. All

statistical analyses were evaluated at the 99

percent confidence level.

Vol. 87 (4)

THE NAUTILUS

113

RESULTS

The results of this study are presented in

figures 1 and 2. Females had a maximum shell

length of 38.3 mm and a mean shell length of

20.2 mm. The respective values for males were

30.0 mm and 17.8 mm. Statistical analysis showed

that the mean shell lengths were significantly

different. Two hundred and thirty-three drills

(58.69%) were females and 164 (41.31%) were

males. This is a female: male ratio of 1.42:1 and

represents a significant deviation from a 50-50 sex

ratio. Significant deviations in sex ratios occurred

in size classes 3 and 4, the female: male ratios

being 1.73:1 and 18:1 respectively. The sex ratios

for size classes 1 and 2 showed no significant

deviation from a 50-50 sex ratio (0.95: 1 and

1.09:1). One hundred females (42.9%) and 39

males (23.7%) were 20 mm or more in length,

while 36 females (15.4%) and only 2 males (1.2%)

were 25 mm or more in length. Eleven females

(4.7%) were greater than 30 mm in length.

DISCUSSION

The results of our study clearly indicate that

sexual dimorphism exists in oyster drills from

Great Bay, New Hampshire. In addition, our work

suggests that this dimorphism is due to the

significantly greater number of females than males

in the larger size classes. This preponderance of

large females was sufficient to yield a significant

deviation from a 50-50 sex ratio.

The sexual dimorphism in Great Bay drills may

be attributable to the following reasons: females

have a faster rate of growth than males, females

? 20-

LENGTH IN MM

FIG. 1. Length-frequency and size distribution of

Urosalpinx cinerea from Great Bay, New Hamp-

shire. Upper figure represents range, mean and ±

one standard deviation.

live longer than males and thus have a longer

period of growth, or both. Another possibility is

that sexually mature males undergo a sex trans-

formation into females. The occurrence of female

U. cinerea with structures resembling vestigal penises

suggests that this deserves investigation.

LITERATURE CITED

Baker, Bemadine B. 1951. Interesting shells from

the Delmarva Peninsula. Nautilus 64 (3): 73-77.

Cole, H. A. 1942. The American whelk tingle,

Urosalpinx cinerea (Say), on British oyster beds.

J. Mar. Biol. Assoc. U. K. 25: 477-508.

Federighi, H. '1931a. Further observafions on the

size of Urosalpinx cinerea Say. J. Conchol. 19:

171-176.

Federighi, H. 1931b. Studies on the oyster drill

{Urosalpinx cinerea Say). Bull. U. S. Bur. Fish.

47 (4): 83-115.

Griffith, G. W. and M. Castagna. 1962. Sexual

dimorphism in oyster drills of Chincoteague

Bay, Maryland-Virginia. Chesapeake Sci. 3(3):

215-217.

Hargis, W. J., Jr. 1957. A rapid live-sexing

technique for Urosalpinx cinerea and Eupleura

caudata with notes on previous methods. Lim-

nol. Oceanogr. 2: 41-42.

Hargis, W. J., Jr. and C. MacKenzie. 1961. Sexual

behavior of the oyster drills: Eupleura caudata

and Urosalpinx cinerea. Nautilus 75 (1): 7-16.

n?

80-

\-

z

Ld

^ 60H

u

Q.

40-

3 4

SIZE CLASSES

FIG. 2. Percent distribution of female and male

Urosalpinx cinerea from Great Bay, New Hamp-

shire in each size class. * = Significant deviation

from a 50-50 sex ratio.

1 14 THE NAUTILUS

October 1973

Vol. 87 (4)

THE MOLLUSCA OF CANADARAGO LAKE

AND A NEW RECORD FOR

LASMIGONA COMFRESSA (LEA)

Willard N. Harman

New York State University College at Oneonta

Oneonta, N. Y. 13820

Canadarago Lake is located in Otsego County,

New York (42°50'N, 75°00'W). The lake is 3

miles long, averages 1 mile in width and has a

surface area of 761 hectares. It is shallow and

turbid with the greatest depth being only a little

over 12 meters and secchi transparencies averaging

less than 2 meters. Summer epilimnion tempera-

tures attain 25°C. Oxygen is absent from the

hypolimnion during these periods and varies near

saturation at the surface. Hydrogen ion concentra-

tion is extremely erratic, varying from about 4.5

to 10 in both surface and profundal waters (Fuhs,

1972). Canadarago would be expected to be a

moderately eutrophic lake in its natural state but

is now highly eutrophic as a result of sewage

input from the village of Richfield Springs and

from summer camps. Blue-green algal blooms are

common and the undersides of cobbles are covered

with fungi and bacteria. Turbid waters prevent the

extensive growth of rooted macrophytes, although

Myriophyllum is abundant in the shallows. Com-

mon emergent plants are Scirpus, Nuphar,

Nymphaea, Pontederia, Typha, and Sparganium. A

eutrophication study of the lake was begun in

1968 by the New York State Department of

Health and the New York State Department of

Environmental Conservation. As a result, a tertiary

sewage treatment plant was installed at Richfield

Springs and was activated in February 1973.

This survey of the moUuscan fauna was under-

taken for the acquisition of base hne data that

can be used for comparison with studies in future

years to indicate possible improvements in water

quality. Samples were made at 9 stations in both

deep waters and Uttoral areas. It appears that

Canadarago once supported dense populations of

mollusks that are now severely depleted. We found

an abundance of empty shells of Lymnaea humilis,

L. palustris, Gyraulus parvus, Helisoma anceps,

Ferrissia parallela, Physa spp., Viviparus georgianus,

Amnicola limosa, A. Integra, Valvata tricarinata,

Lampsilis radiata, and Pisidium spp. Only a few

hving specimens of L. humilis, G. parvus, F.

parallela, Physa sp. and Pisidium spp. were col-

lected. All Physa, empty shells and living speci-

mens, were so small as to prevent confident

determination to the species level. One living

specimen of Lasmigona compressa (Lea) (Union-

idae) was found at the entrance of Herkimer

Creek into the lake. This is the first record of this

species in the Susquehaima River watershed.

After noting the condition of the molluscan

fauna in the Lake we made collections along 5

miles of Oaks Creek, the outlet of Canadarago, to

ascertain if water quality improved downstream

from the lake. The only snail found was an

immature Physa sp. A few living specimens of the

bivalves Lampsilis radiata, Elliptio complanatus and

Strophitus undulatus were found. Empty shells of

all the mollusks found in Canadarago Lake were

collected.

LITERATURE CITED

Fuhs, W. 1972. Canadarago Lake eutrophication

study. Lake and tributary surveys 1963-1970.

Methodology and Data. N.Y.S. Dept. of En-

viron. Cons., Research and Development Unit.

292 pp.

Vol. 87 (4)

THE NAUTILUS

115

OBITUARY

Elmer P. Cheatum

(1901-1973)

On May 1, 1973, science and education lost

one of their most notable contributors with the

death of Dr. Elmer P. Cheatum. Throughout his

48 years of service at Southern Methodist Uni-

versity he remained a leader of Southwestern

scientific investigation and educational inspiration.

Elmer Phillip Cheatum was born in Langdon,

Kansas, July 19, 1901. He grew up in rural

surroundings assisting in his father's general store.

As a young man, he washed dishes in a hotel near

Southwestern College where he earned his B.A. in

biology (1921-1924). He was also a good athlete,

setting the Kansas record for the sprint race that

stood for some time. Later, he earned his M.S. in

zoology at Kansas State College (1924-1925). At

this time, he married Edith Deck and left alone

two days later to assume a job as instructor at

Southern Methodist University. Five weeks later he

had earned enough money to bring his bride to

Dallas. During summers and for two winters he

and Edith traveled to the University of Michigan

where he gained his Ph.D. in aquatic biology

under such people as Dr. Paul Welch (1931-1933).

As a scientist and educator. Dr. Cheatum

worked in several fields, parasitology, entomology,

malacology and Umnology. Although he continued

to teach in these fields and in comparative anatomy,

his main research interest gravitated to the Mol-

lusca. He pubUshed over 30 scientific articles,

some in The Nautilus (1934-1972). Most were

concerned with the area he loved most, Texas.

His academic affiliations were numerous. He

was a member of Sigma XI, a fellow in the

A.A.A.S., a member of the American Microscopical

Society, a fellow and past President (1941-1942)

of the Texas Journal of Science, a member of the

Texas Ornithological Society and the Limnologjcal

Society of America. In Dallas, he belonged to the

S.M.U. Men's Faculty Club, Town & Gown Qub,

and he was a director of the Junior Leader

Training, Circle Ten Council, Boy Scouts of

America. For his leadership in Boy Scout work, he

was given the Silver Beaver Award.

Although Dr. Cheatum was a moving force in

the scientific worid, I think his true greatness lay

in his teaching. He had the unique ability to

inspire student thought and activity far beyond

the classroom. He led students to almost every

type of environment that Texas had to offer.

These field trips with Dr. Cheatum were truly

experiences that few students ever forget. Dr.

Cheatum's life work and collections have been

transferred to the Dallas Museum of Natural

History.

by Richard W. Fkllington

Dallas Museum of Natural History

P. O. Box 26193, Fair Park Sta., Texas 75226

JUST PUBLISHED

Third Edition, enlarged and

completely rewritten,

now includes shells

of the West Indies,

This enlarged edition of the Atlantic Coast

shell guide takes its place as an up-to-date

counterpart of the author's Field Guide to

Shells of the Pacific Coast and Hawaii.

More species and new material on individual

shells have been included, and information in

earlier editions has been brought up to date

throughout. Redesigned and entirely reset, the

new edition covers 1035 shells, 1033 of which

are illustrated with photographs. All the plates

are new, and there are 31 additional plates.

Changes in typography and arrangement en-

hance the ease of use. Boldfaced subheadings

point up facts about range, habitat, and physi-

cal characteristics. Illustrations are conven-

iently located in the center of the book for

quick reference. And the area of coverage has

been expanded to include the more common

shells of the West Indies— a boon to those who

visit the Caribbean.

Introductory sections contain all the help and

advice a beginner needs to start him on a fasci-

nating hobby. For the experienced collector

there exists no equally satisfactory one-volume

guide to identification.

A Field Guide to Shells

OF THE ATLANTIC AND GULF

COASTS AND THE WEST INDIES

By Percy A. Morris

EDITED BY WILLiAM J. CLENCH

At your bookstore. Cloth, $7.95. Paper, $4.95

HOUGHTON MIFFLIN COMPANY • 2 Park Street, Boston, Massachusetts 02107

INFORMATION FOR SUBSCRIBERS

The annual subscription rate for The Nautilus is

$7.00 for individuals and $12.00 for institutions

(domestic or foreign). Subscriptions may begin in

January. Send check or money order to "The

Nautilus" to Mrs. Horace B. Baker, Business

Manager, 11 Chelten Road, Havertown, Pa. 19083.

Back issues from volume 72 to date are

obtainable from the Business Manager. Volumes 1

througli 71 (if available) may be obtained in

reprint or original form from Kraus Reprint Co.,

Route 100, Millwood, New York 10546.

Advertising rates may be obtained from the

Business Manager or Editor.

CONTRIBUTORS

Manuscripts: Authors are requested to follow the

recommendations of the Style Manual for Biological

Journals, which may be purchased from the American

Institute of Biological Sciences, 2000 "P" Street,

N.W. Washington, D.C. 20036. Manuscripts should be

typewritten and doublespaced; original and one copy

are required, to facilitate reviews. Tables, numbered

in arable, should be on separate pages, with the title

at the top. Legends to photographs should be typed

on separate sheets. Explanatory terms and symbols

within a drawing should be neatly printed, or they

may be pencilled in on a translucent overlay, so that

the printer may set them in 8 pt. type. There is a

charge of 50 cents per word for this extra service. All

authors or their institutions will be charged 50 cents

per line of tabular material and taxonomic keys. The

pubhshers reserve the right, seldom exercised, to

charge S32 per printed page.

An abstract should accompany each paper.

Reprints and covers are available at cost to

authors. When proof is returned to authors,

information about ordering reprints will be given.

They are obtained from the Economy Printing Co.,

Inc., R. D. 3, Box 169, Easton, Maryland 21601.

MOLLUSK VOUCHER SPECIMENS

It is becoming increasingly important for future

research purposes that an identified sampling of

species mentioned in pubHcations be deposited in

a permanent, accessible museum speciahzing in

mollusks. This is particularly true of mollusks used

in physiological, medical, parasitologjcal, ecological,

and experimental projects.

The Delaware Museum of Natural History has

extensive, modern facilities and equipment for the

housing and curating of voucher specimens. Mater-

ial should be accompanied by the identification,

locahty data and its bibliographic reference. There

is no charge for this permanent curating service,

and catalog numbers, if desired, will be sent to

authors prior to pubUcation.

MBl. WHOI I IHKARY

liJH 17XV Z