ANUARY 25. 1982

THE

NAUTILUS

1^«N ()()28-i:i44

Vol. 96

No. 1

A quarterly

devoted to

malacology and

the interests of

conchologists

Founded 1889 by Henn' A. Pilsbry. Continued by H. Burrington Baker.

Editor-in-Chief; R. Tucker Abbott

EDITORIAL COMMITTEE

CONSULTING EDITORS

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

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

103 West 8th Avenue

Cudjoe Gardens

Summerland Key, Florida 33043

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

Dr. Charles B. Wurtz

3220 Penn Street

Philadelphia, Pennsylvania 19129

EDITOR-IN-CHIEF

Dr. R. Tucker Abbott

American Malacologists, Inc.

Box 2255, Melbourne, Florida 32901

Mrs. Cecelia W. Abbott

Business and Subscription Manager

P.O. Box 22.55

Melbourne, Florida 32901

Tlie Nautilus (USPS 374-980)

ISSN 0028-1344

OFFICE OF PUBLICATION

American Malacologists, Inc. (United Parcel Address:

2208 South Colonial Drive, Melbcjurne, FL 32901)

Mail: Box 2255, Melbourne, FL 32901

Second Qaas Postage paid at Melbourne, Florida

and other post offices

Subscription Price: $12.00 (see inside back cover)

$14.00 (foreign); institutions $18.00

THE

NAUTILUS

Volume 96, number 1 - January 25, 1982

ISSN 0028-1344

CONTENTS

Joseph Rosewater

A New Species oiHipjiopu^- (Bivalvia: Tridaciiidae)

Francesco B. Trama

Occurrence of the Asia^c Clam Corbicula Jluininea in the Raritan River,

New Jersey 6

William D. Shepard

Rediscovery of a Portion of the Isely Unionid Collections 8

Philippe Bouchet and Francoise Danrigal

Napoleon's Egyptian Campaign (1798-1801) and the Savigny Collection of Shells 9

Clement L. Counts, III and Robert S. Prezant

Shell Microstructure of Corbicula flmninea (Bivalvia: Corbiculidae) 25

Paul W. Parmalee and Walter E. Klippel

A Relic Population oi Obuvaria retussa in the Middle Cumberland River,

Tennessee 30

Publications Received 2 Recent Deaths 2

THE NAUTILUS

January 25, 1982

Vol. 96(1)

PUBLICATIONS RECEIVED

Brooks, Stanley Truman (1902-1960), Biobibliosraphy by

R. I. .lohnson in Ocr. fnpers MoUuaka. va]. 4, no. 59,

pp. 266-268(1981).

Clarke, Arthur H. 1981. The Tribe Alasmidontini (Unioni-

dae: Anodontinae), Part 1: Pegias, Alasmidon.la, and

A rcidenx. Smithsonian Contributions to Zoology, no. 326,

pp. iii + 1-101, 32 figs. Ecologic and taxonomic study with

illustrations of shell, soft parts, gloehidia, descriptions,

distributional maps, synonymies, and simple anatomical

test for sex determination. Alusmidontn mliusta n. sp..

extinct mussel from (jharlotte, N(" (not Vnio mlntatif

Sowerby, 1840).

Habe, T. and Iwao Taki. 1981. A Catalogue of Molluscs of

Wakayama Prefecture, the Province of Kii. 1. Bivalvia,

Scaphopoda and Cephalopoda. F'ubl. Seto Marine Biol.

Lab.. Special Series, vol. 7, no. 1, pp. xx 304, 13 pis. (One

in color). Based on Kuroda's Manuscript. Eleven new

species of .Japanese Bivalvia are described. Synonymies

very complete. 51 cephalopods treated by I. Taki (pp.

234-264).

.Johnson, Richard I. 1981. Recent and Fossil Unionacea and

Mutelacea (Freshwater Bivalves) of the Caribbean Islands.

Occasional Papers on Mollii.<!ks. vol. 4, no. 60. pp. 269-288.

•SI. 60 from Dept. Mollusks. Museum ('omparative Zool-

ogy, Cambridge, MA 02138.

Martins, Antonio M. Frias. 1981. Oxyrliilus. (Drouetia)

agosl.inhm New Species (Stylommatophora: Zonitidae)

from the Azores Islands, its Anatomy and Phylogenetic

Relationships. Occ. Papers on Mollwsks, vol. 4. no. 59.

pp. 245.

Rehder, Harald A. 1981. The Audubon Society Field Guide

to North .American Seashells. 894 pp., 705 color pis. An

excellent pocket guide of the common shells, with espe-

cially good photographs by James H. Carmichael, Jr.

Gives descriptions, habitats, range and both popular and

Latin names, but unfortunately omits all authors and

dates.

.Sharabati, Doreen. 1981. Saudi Arabian Seashells. 119 pp.

VNU Books International. A popular account with numer-

ous beautiful photographs, many of living mollusks.

T}iais. 1981. Publicaciones Diversas de la Sociedad Pana-

mena de Malacologia. No. 1 (La Fauna Malacologica

Panamena), 28 pp.; No. 2 (Lista Prelimininar de Referen-

cias Bibliograficas Sobre la Fauna Malacologica

Panamena), 59 pp. Both by Miguel C. Aviles E. Annual

subscription Balboa 10.00 (U.S. $10.00), Soc. Panamena de

Malacologia, Estafeta Universitaria, Panama, Rep. of

Panama.

RECENT DEATHS

Grab, Robert McKay, former President of the Naples Shell

Club, bom New York City on April 19, 1904, graduate

University of Michigan in 1926, died in North Carolina

September 1981.

Lyman, P>ank, long-time shell dealer, editor of "Shell

Notes" (1942-51), born in Lantana, Florida, June 10,

1895, died Oct. 11, 1981, in Ferandina Beach, Florida.

Sexauer, Howard T. 1981-82 President of the Sanibel

Captiva Shell Club, well-known poet, former resident of

Montpelier, Vermont, died at age 82 in Florida on Dec. 1,

1981.

Zeigler, Rowland Franklin, born .lune 1, 1915, in Greely-

ville, SC, formerly chief of obstetrics and gynecology at

McLeod Hospital in P'lorence, SC, author (with Porreca)

of "Olive Shells of the World" and articles in medical

journals, died at age 66, on Sept. 21, 1981, in Durham,

NC. His widow, Edna Jo continues his interests in olive

shells.

I'ln' I'irsI a>vrhfdogiral wtirk liij mi .■\mmcaii

jiuhHuhed in A mtn'icit.

THOMAS Savs CONCHOLOGY

1819

A beaut iful facsimile copy of the third

edition of Say's fiindeinental umrk from

Nicholson's British Encyclopedia

Philadelphia 1819

This rare work, never before reprotluced in full, con-

tains 20 pages and four lithograph plates of Say's new

land and freshwater species of mollusks. This is the

expanded version of the first (1816) and second (1818)

edition. 32 pages, paperbound.

The [ireface, by R. Tucker .Abbott, tells the history of

this article and gives brief biographies of the author,

the editor, the publisher and the printer. Limited edi-

tion. $5.00, including postage. F'oreign postage,

plea.se add $1.00.

Amki;i('an MALACOUHiisTS, Inc.

P.O. Box 2255

Mi-llMiurne. FL 32»(I1-I«2S

Vol. 96(1)

January 25, 1982

THE NAUTILUS

A NEW SPECIES OF HIPPOPUS (BIVALVIA: TRIDACNIDAE)

Joseph Rosewater

Department of Invertebrate Zoology

National Museum of Natural History

Washington, D.C. 20560

ABSTRACT

Hippopus porcellanus n. sp. is described from Sibutu Island, Sulu Archipelago,

Philippines, bringing the known members of the genus Hippopus to 3, including

H. hippopus (Linne, 1758), Recent, I ndo- Pacific, and H. gimteri Mansfield, 1937,

Fossil. Lower Miocene of Florida.

For several years shell dealers have received

large numbers of a rather distinctive giant clam

belonging to the genus Hippopus which they

refer to as the "China Clam", and which they

consider to be distinct from the "Horse's Hoof

or "Bear Paw", Hippopus hippopus (Linne,

1758). It is much thinner and smoother than the

usually elaborately sculptured H. hippopus, and

lacks most of the characteristic strawberry color

of the latter. When I reviewed the classification

of Tridacnidae this phenotype was believed to

intergrade completely with H. hippopus (Rose-

water, 1965, p. 361). Recent examination of

nearly two dozen specimens of the "China Clam"

and comparison with H. hippopus persuades me

that they are separate species.

Hippopus porcellanus new species

(Figs. 1-4)

Description: Shell reaching 216 mm (about 8

1/2 inches) in length, semicircular in outline and

globose in shape; usually only moderately in-

flated; with valves closed byssal orifice is very

narrowly gaping. Valves not excessively heavy,

markedly translucent, colored occasionally with

weak strawberry blotches arranged concentri-

cally or scattered; color of interior porcel-

laneous, of exterior grayish white. Surface of

valves remarkably clean, except for scattered

coralline algae and debris. Primary radial sculp-

ture consisting of 13 or 14 low rib-like folds

distributed over surface of valve, extending on-

to ventral slope where they become obsolete.

Secondary radial sculpture consisting of low rib-

lets which are nearly obsolete on primary folds

but are more prominent in their interstices.

Riblets varying somewhat in width, usually lack-

ing spines or evidencing only microscopic con-

centric imbrications. A few low tubular spines

present on primary fold bordering ventral slope

and occasionally on ventral portions of other

primary folds; folds usually smooth dorsally.

Concentric sculpture consisting of microscopic,

wavy, imbricate lines of growth. Dorsal margin

undulate, with series of 8-9 rounded to squarish,

medially projecting, interdigitating processes

representing extremities of rib interstices.

Hinge line usually longer than half the length of

valve. One oblong cardinal tooth in each valve; 2

elongate posterior laterals in right and a single

moderately sharp one in the left valve. Liga-

ment secondarily prosodetic. Umbos directed

postero-medially. Edge of byssal orifice with a

series of 8-12 light-yellow, rather poorly

developed plicae which remain fairly constant in

size or become only slightly larger posteriorly.

Ventral slope moderately concave. Hinge plate

suffused with orange. Pallial line entire, mod-

erately narrow. Muscle scars central, medium

sized; the posterior adductor scar round in both

valves, the posterior pedal retractor scar is

smaller and elongate, the two extending over

portions of two fold interstices in each valve.

Area within pallial line, excluding muscle scars,

dull; pallial line, muscle scars and areas to edge

of shell shiny. Prodissoconch unknown.

The anatomy and life history of this species

are unknown. Following my studies (Rosewater,

1965) several persons have achieved success in

studying the spawning and development of Hip-

THE NAUTILUS

January 25, 1982

Vol. 96(1)

Vol. 96(1)

January 25, 1982

THE NAUTILUS

popus hippopus (Jameson, 1976; Gwyther and

Munro, 1981).

Measurements (mm): Tridacnidae are indis-

criminately inequivalved, and the figures given

under "length" and "height" are always the max-

imum measurement. "Width" is the greatest

distance through opposed valves.

Lticality Sihiitu MaMmte

(Philippines) Zmtiliiiiniiia hi Id. SnIii Sra

Types: Holotype ANSP (The Academy of

Natural Sciences of Philadelphia) 246600, 157

mm length, 113 mm height; 3 paratypes ANSP

354770, 155 mm length, 115 mm height, 101 mm

length, 81 mm height, and 75 mm length, 53 mm

height; 1 paratype USNM (National Museum of

Natural History, Washington, D.C.) 807720, 133

mm length, 98 mm height.

Type locality: Sibutu Island, Tawi Tawi

Group, Sulu Archipelago, Philippines (4°46'N;

119°29'E), du Pont- Academy Expedition, 1958.

Other Material Examined: ANSP 209699

Zamboanga, Mindanao Island, Philippines, ex.

A. B. Bronson, 1956 (10 specimens, see Meas-

urements); ANSP 228977, Masbate Island,

Philippines, du Pont-Academy Expedition, 1958

(1 specimen, see Measurements); USNM, "Sulu

Sea", ex. A. D'Attilio and John Root (6 speci-

mens, see Measurements).

Distribution: Philippines, mostly known from

the Sulu Archipelago; 1 specimen from Masbate

Island, central Philippines.

Etymology: "porcellanus", an adjectival name

referring to the porcelain-like appearance of the

shell of this species.

Remarks: Shells of this new species are con-

sistently smoother, more semicircular in outline,

proportionately lighter in weight and thinner

than H. hippopus (see figs. 1-4). When viewed

apart from the latter they present a rather ro-

tund appearance although measurements show

they are no more obese, but, in fact, are often

less wide. Compared with H. hippopus. H.

porcellanus has a very narrow byssal orifice,

with the plicae less well developed and lighter in

color (fig. 4). The single posterior lateral tooth

of the left valve is moderately sharp in H.

porcellanus while in H. hippopus it is blunt.

The only other large bivalve species with

which H. porcellanus is likely to be confused is

Tridacna derasa (Roding, 1798). Specimens of

T. derasa, of the same length as H. porcellanus

(200 mm-f ), are smooth, may be similarly semi-

circular in outline and exhibit similar obesity. It

is quite likely that living T. derasa has a con-

siderably more colorful mantle, that of H. hip-

popus, at any rate, being rather sombre olive.

The shell of T. derasa is even smoother than

that of H. porcellanus, lacking in development

of tubular spines and having low primary and

secondary folds, although sometimes developing

strong, continuous, undulate concentric ridges

(Rosewater, 1965, pi. 281, fig. 1). Shells of T.

derasa lack the orange and yellow coloration

present in Hippopus in the areas of hinge and

byssal orifice. Byssal plicae number 6-7, are low

and elongate to nearly obsolete in T. derasa

while in H. pjorcellanus they are shorter and

number from 8-12. The posterior adductor

muscle/posterior pedal retractor scar complex is

comparatively larger in T. derasa and its umbos

tend to be less distinctly convoluted than in H.

porcellanus. although size for size the former

tends to have a heavier shell.

I originally thought that H. porcellanus dif-

fered from H. hippopus subspecifically rather

than specifically. As subspecies the two would

normally have more or less discreet geographic

ranges. While it appears that H. porcellanus

lives mostly in the southern Philippines, more

precisely the southern Sulu Sea, a specimen has

been reported from Masbate Id. in the central

Philippines. Furthermore, H. hippopus is

distributed in the same area besides having a

more extensive range in the western Pacific.

Since the ranges of the two species seem not to

be in any real way mutually exclusive, the sub-

specific status seems doubtful, and they are here

considered to be separate species within the

THE NAUTILUS

January 25. 1982

Vol. 96(1)

genus HippoptLs. It is quite obvious that the

known geographic range of H. porcellanus is

considerably more restricted than that of//, hip-

popus (see Rosewater, 1965, pi. 272).

ACKNOWLEDGMENTS

Special thanks are expressed to those who

assisted in gathering information and donating

specimens: Anthony D'Attilio, John Root,

Robert W. Morrison, R. T. Abbott, and Betty

and Robert Lipe. I am grateful to R. Robertson,

ANSP, for loaning specimens with accurate

locality data. R. D. Turner, and K. J. Boss,

Museum of Comparative Zoology, Harvard

University, and H. A. Rehder, and R. S.

Houbrick, USNM, offered helpful criticism.

LITERATURE CITED

Gwyther, J. and J. L. Munro. 1981. Spawning induction and

rearing of larvae of Tridacnid Clams (Bivalvia: Tridacni-

dae). AquaeuHurc 24:197-217.

Jameson, Stephen C. 1976. Early life history of the Giant

Clams Tridarna crocea Lamarck, Tridarna maxima

(Roding). and Hippopus hippopus (Linnaeus). Pacific

Science 30(3):219-233.

Rosewater, Joseph. 1965. The Family Tridacnidae in the

Indo-Pacific. Indo-Pacifie Mollusca l(6):347-396.

Shell Factory. 1963. Shells of the Winid. Catalog no. 109,

160 pp. Fort Myers. Florida. "China Clam" illustrated

on p. 45.

OCCURRENCE OF THE ASIATIC CLAM CORBICULA FLUMINEA IN THE

RARITAN RIVER, NEW JERSEY^

Francesco B. Trama

Department of Biological Sciences

Rutgers LTniversity

New Brunswick, New Jersey

ABSTRACT

The mid-Atlantic range of the Asiatic clam, Corbicula fluminea (Muller. 1771^)

is extended to the Raritan River in central New Jersey. A successfully breeding

population was found in the nontidal region near a water supply intake. These

clams have also colonized far upstream into the South Branch of the Raritan but

not the North. Branch. No reason for this difference is known. Living specimens

were found downstream in the tidal portion of the river which is freshwater but

polluted. There was no evidence of reproduction in this region of the river.

Since it was first observed in this country in

the Columbia River, Washington (Dundee and

Dundee, 1958) the Asiatic clam, Corbicula

fluminea (Muller, 1774; (alias manilensis Philip-

pi) has spread rapidly into many river systems

across the United States (Sinclair, 1971). Exten-

sion of its range to the mid- Atlantic region was

documented by Diaz (1974) in the James River,

'This study was supported iiy the Center for Coastal and En-

vironmental Studies. Rutgers University. Assistance of Joy

Bergelson in pierforming field collections is acknowledged.

Virginia, and by Fuller and Powell (1973) in the

Delaware River between Philadelphia, Pennsyl-

vania and Trenton, New Jersey. Diaz estimated

the year of introduction in the James River to be

1968 and Fuller and Powell concluded that Cor-

bicula was present in the Delaware River since

at least 1971 or 1970. Crumb (1977) later re-

ported it in the Delaware River between Tren-

ton and Burlington in September 1971.

On March 26, 1981 many empty shells and a

few living Corbicula were collected from a tidal

(Init freshwater) region of the Raritan River

Vol. 96(1)

January 25, 1982

THE NAUTILUS

near New Brunswick, New Jersey (Sta. 1, Fig.

1). Collections of macrobenthos taken from this

same region in September 1980 did not contain

any specimens of Corbicula. The largest shell

collected in March 1981 was 25 mm in length

and was estimated to be between 3 and 4 years

of age at the time of death. It was concluded

that the year of invasion was not later than 1978

and that the invasion site had to have been

upstream from this point of initial discovery.

During July and August 1981 the occurrence

and range of Corbicula in the Raritan River was

studied. Samplmg extended upstream from Sta.

1 (tidal but freshwater) into the North and South

Branches of the Raritan River (Fig. 1).

At Sta. 1 many empty shells were again

found. Many live specimens, however, were also

collected from current-swept bottoms covered

by gravel and small pebbles The livmg clams

fell into two size classes; largest specimen was

25 mm in length and had a grossly distorted

shell growth that appeared to commence from a

shell length of 20 mm. The smallest specimen

was 15 mm in length and appeared nt)rmal.

The greatest population density and evidence

of reproduction were found at Sta. 2 which is in

the vicinity of the Elizabethtown Water Com-

pany's intake on the Raritan River which is non-

tidal at this point. Largest living clams ranged

from 15 to 17 mm in length and a first year class

ranged from 2 to 6 mm in length. No Corbicula

were found in the Millstone River (Sta. 3) im-

mediately above its confluence with the Raritan

River. The substrate in this portion of the

Millstone River was soft muck and, therefore,

was not a preferred habitat for these clams.

Very few living Corbicula were taken at Sta-

tions 4 and 5, and these ranged from 7 to 10 mm

in length. Many empty shells were found at Sta.

4. Even though a suitable type substrate existed

at Stations 6, 7 and 8 only one specimen of Cor-

bicula (10 mm) was found at Sta. 6. None was

found at Stations 7 or 8, nor was there evidence

of dead Corbicula shells anywhere in the North

Branch of the Raritan River. On the other hand,

tliere were significant populations in the South

Branch. Specimens ranged from 2.5 to 10 mm in

length at Stations 9, 10 and 11. No dead

no. 1. Samjile files ,„, the Ranta,, Hirer. Netr Jersey. Samplvngfor Corbicula look place belween July 2:1 and

August 2ti. 1981.

8 THE NAUTILUS

January 25, 1982

Vol. 96(1)

specimens or empty shells were found in those

locations.

It seems that the Asiatic clam has established

itself in the tidal and nontidai, freshwater (even

polluted) regions of the Raritan River. The

population center (initial colonization?) is in the

nontidai waters of the main stem in the vicinity

of the intake of the Eiizabethtown Water Com-

pany. Furthermore, this foreign species has suc-

cessfully colonized the South Branch of the

Raritan River but for some reason has not as yet

moved very far into the North Branch of the

Raritan River. This discovery extends the north-

ern range of Corbicula in the mid-Atlantic

region.

LITERATURE CITED

Crumb, S. E. 1977. Macrobenthos in the tidal Delaware

River between Trenton and Burlington, New .Jersey.

Chesapeake Science 18:253-265.

Diaz, R. J. 1974. Asiatic clam, Corbicula manilensis (Philip-

pi), in the tidal James River, Virginia. Chesapeake Science

15:118-120.

IXindee, D. S. and H. A. Dundee. 1958. Extension of known

ranges of four mollusks. The Nautilus 72:51-53.

Fuller, S. L. H. and C. E. Powell. 1973. Range extensions

(jf Corbicula manilensis (Philippi) in the Atlantic drainage

of the United States. The Nautilus 87:59.

Sinclair. R. M. 1971. Annotated bibliography on the exotic

bivalve Carbicula in North America, 1900-1971. Sterki-

anaNo. 43:11-18.

REDISCOVERY OF A PORTION OF THE

ISELY UNIONID COLLECTIONS

William D. Shepard

Oklahoma Biological Survey

Norman, Oklahoma 73019

During a recent renovation of the Inverte-

brate Range at Stovall Museum of Science and

History (University of Oklahoma), a collection of

unionids was found carefully wrapped in old

newspapers and stored away. Part of these were

labeled as collected by F. B. Isely in Oklahoma

between 1908 and 1912. The remaining unionids

were also from Isely's collections. Upon inquiry

with several malacologists, it appears that this

is the largest extant portion of Isely's collec-

tions. These are part of the specimens collected

by Isely for his study of the clams of Eastern

Oklahoma, which appeared in 1925.

After rearrangement of the above specimens

and merging of several other collections, a

series of unlabeled unionids were found. These

differed from all the other specimens, however,

in that they had small, circular, numbered, cop-

per tags wired to the shells. On further examina-

tion it was determined that these were a portion

of the specimens used by Isely in his classical

migration study (Isely 1914). They were also col-

lected in Oklahoma.

The rediscovery of these specimens is impor-

tant for several reasons. They were used by Ise-

ly and therefore are now voucher specimens for

liis works. Their presence also allows some in-

sight into the taxonomy of his time or at least

Isely's interpretation of it. Secondly, the

specimens have a historical importance in that

some of the streams Isely collected have now

been altered so as to deplete the original unionid

fauna. Finally, some of these specimens repre-

sent the only collections from various streams in

Oklahoma.

Isely's specimens plus others donated to

Stovall Museum form a nucleus for a growing

collection of Oklahoma unionids. The Stovall col-

lection combined with that located at OU's

Biological Station on Lake Texoma (Texas-

Oklahoma border) represents perhaps the major

collections of Oklahoma unionids.

LITERATURE CITED

Isely, F. B. 1914. Experimental study of the growth and

migration of freshwater mussels. Bureau of Fisheries

Document, No. 792.

1925. The freshwater mussel fauna of Eastern

Oklahoma. I'rur. OkUi. Acad. Sci. 4:43-118.

Vol. 96(1)

January 25, 1982

THE NAUTILUS

NAPOLEON'S EGYPTIAN CAMPAIGN (1798-1801)

AND THE SAVIGNY COLLECTION OF SHELLS

Philippe Bouchet and Francoise Danrigal

Museum National d'Histoire Naturelle

55, Rue de Buffon

75 Paris (5°), France

ABSTRACT

During the Napoleonic campaign to Egypt a collection of Red Sea, Mediter-

ranean and continental niollut^k^ ivas brought together by Savigny who published

upon them in 1817 as folio engravings. These served as type figures for a number

of species described by 19th ceyitury malacologists, including Audouin,

Ehrenberg, Deshayes, Philippi, Hupe, Landrin, Jonas, Issel, Morlet, Vaillant, P.

Fischer, Tapparone-Canefri. von Martens, Weinkauff, Monterosato, Jousseaume,

H. Fischer and finally Pallary. A checklist and illustrations of 86 of these

original types frmn the Savigyiy collection, now in the Paris Museum, are

presented, supplemented by reproductions of the unpublished color vellums of

opisthobranchs and cephalopods.

Jules Cesar Savigny was born at Provins,

France, on April 5, 1777. In 1793, in the middle

of the Revolution, the Convention transformed

the Royal Gardens into a "Museum d'Histoire

naturelle". Savigny was then 16 years old and

was sent to Paris to study at the "Ecole de

sante" and the Museum. His professors were

Lamarck, Cuvier, Daubenton; they noted the

young student's assiduity and also took note

that he was always wearing the same Nankin

costume.

Lamarck was then writing the "Nouvelle En-

cyclopedie" and assigned Savigny his first work,

the drafting of the part on the sorrel plant. This

paper showed the wealth of his knowledge and

soon he was named professor of botany at the

"Ecole Centrale" in Rouen. Cuvier advised him

to wear a wig to look older and more serious!

But Savigny was never to go to Rouen. Napo-

leon Bonaparte was already preparing the

Egyptian Campaign.

Bonaparte not only wanted to conquer the

country but also make it a model of French

culture. He added to the expedition troops a

"Commission des sciences et des arts", con-

sisting of scholars, painters, architects and

others. Bonaparte asked his friend Cuvier to

name the naturalists who would join the expedi-

tion. Savigny and Geoffroy Saint Hilaire were

selected for the zoological part. Savigny was

then 21 years old; he had been trained as a

botanist, but Cuvier's opinion was that "he

would become a zoologist when he would decide

to".

The Expedition

After the Battle of Campoformio in 1797,

Bonaparte realized that he could not invade

England, so he decided to make war on Egypt.

In 1798, 38,000 men gathered at Toulon on 335

ships; among them were the members of the

"Commission des Sciences et des Arts". On

Floreal 30th, year VI, (i.e. May 19th, 1798)

Geoffroy Saint Hilaire and Savigny left Toulon.

They were to reach Alexandria 41 days later.

Savigny was placed in the 4th class of the Com-

mission, with a very low salary, but thanks to

Geoffroy Saint Hilaire, he was soon admitted to

the first class.

The Nile was flooding and the expeditionary

forces were directed to Cairo which they en-

tered on July 23 after the Battle of the Pyra-

mids. During this time the Commission settled

in Rosetta for a month. Savigny and Geoffroy

Saint Hilaire stayed for several weeks in the

islands of Lake Menzale; in September they

visited Salahied and Damiette. They worked all

during the autumn in the delta and left Damiette

for Cairo in December. Geoffroy Saint Hilaire

was exhausted, but Savigny was in very good

condition. A current joke of the time in Cairo

was to call scholars, the donkeys, then used as

"taxis", because the members of the Commission

were always travelling on donkeys.

Reprints of this article are available for U.S. $7.00

(postage paid) from American Malacologists. Inc..

P.O. Box 2255, Melbourne. FL. ;32901, U.S., A.

10 THE NAUTILUS

January 25, 1982

Vol. 96(1)

Between August 1 and 2, Nelson defeated the

French Naw in Aboukir and cut all contacts

with F" ranee. In Cairo, Bonaparte organized the

country under his rule and founded the "Institut

d'Egypte". He then prepared the expedition to

Syria 13,000 infantrymen and the cavalry left

in the beginning of February, 1799. After sev-

eral victorious battles, Bonaparte was finally

stopped at Saint Jean d'Arce, where 4,000 were

killed. On June 14 the army was back in Cairo,

Savigny was the only naturalist to follow the

troops.

In August, Bonaparte turned over the com-

mand to Kleber and returned to Paris, where

the "Directoire" was weakened. The members of

the "Commission" were sent to Suez at the end

of 1 799. Savigny was very e.xcited by the wealth

and beauty of the Red sea fauna. But Kleber had

to face the Turks and the British and the

scholars were sent back to Cairo. Kleber was

murdered shortly after he won the Battle of

Heliopolis; France was then finally defeated at

Canope in 1801.

The journey of the 45 members of the "Com-

mission" back to France was rather unfortu-

nate. On April 6, 1801, they left Cairo for Alex-

andria. On their arrival they were quarantined

by the French general, Menou. They were trans-

ferred to a brig, and waited 21 days for permis-

sion to leave the bay. The British troops stopped

them, took them to Abukir, then back to Alexan-

dria. Menou had surrendered on terms stipulat-

ing that the collections gathered by the "Com-

mission" in Egypt should be given to the British.

The scholars rebelled and threatened to destroy

the collections. Finally, on September 26, 1801,

they were allowed to go back to France with

their collections.

The Original Plates

Savigny returned to Paris in February, 1802.

He did a great amount of work on the material

he took back with him and published various

papers and memoirs between 1801 and 1810.

Among them were the study of 1,200 insects

and his famous "Histoire naturelle et mytholo-

gique de I'Ibis" (1805). In 1817, he published

125 plates for the atlas of the "Expedition

d'Egypte", in "grand-aigle" size (71.5 x 52 cm);

in 1826 they were reprinted by Panckoucke in

reduced size (68.5 x 52 cm). These engraved

plates were prepared from the original colored

vellums now preserved in this Museum's library.

They had beon drawn very accurately, with a

wealth of details, by the best artists of the time.

Because the "Commission" judged that they

were too difficult to reproduce in color, only

black and white copies were printed.

Savigny suffered progressively failing eye-

sight, approaching blindness after 1815, for

which reason he never published any text to ac-

company the plates. But Cuvier found that this

iconography was unique and Audouin was cho-

sen to write an explanatory part. Audouin was a

professor of entomology at the Museum, and

was not prepared for this work. But he was the

son-in-law of Brongniart, a member of the "In-

stitut", director of the Sevres manufacture, and

a friend of the powerful Cuvier.

There is considerable difficulty in determining

the exact dates of publication of the "Explica-

tion sommaire des planches", of which two edi-

tions were printed.

The first edition bears on the title page "public

par les ordres de sa majeste I'Empereur

Napoleon Le Grand" and is dated 1809 on the

cover page, but was evidently published much

later (Laissus, 1973); the Mollusca are treated in

tome 1, part 4, and begin with a copy of a letter

dated Nov. 1, 1825! The register of the letters

and declarations of the "Commission d'Egypte",

kept in the Manuscript Department of the Na-

tional Library, Paris, shows that the Natural

History section belongs to the third issue of

reports. On April 13, 1826 the last manuscript

by Audouin concerning shells was sent to the

"Imprimerie Royale", but we could not deter-

mine whether the first edition, third issue, was

actually printed before the end of 1826.

However it seems wise to assume that Sherborn

(1897:287) was right and that this part "may be

safely regarded as dated 1826."

Almost simultaneously a second edition,

"dediee au roi", was being printed by

Panckouke. This bears 1827 on the title page but

it appears also that this date is a fake. Pallary

accepted this date (1827) in a bibliography, but

mentioned 1829 in the text (Pallary 1926). In

fact the earliest date we can ascertain is July 19,

1828 (Bibliographie de la France, 1828).

The text was published while Savigny was still

alive. When he became aware of the contents, he

became very irritated and sent a letter to the

Academy of Sciences. Later on, L. Pfeiffer pro-

nounced similarly .severe judgement, and re-

proached Audouin with the fact that he had

taken all the benefit of Savigny's work. P.

Fischer and Pallary had the same opinion. In the

checklist, we have given references to both edi-

tions of Audouin's text because the first one is

apparently very rare in libraries.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 1 1

Savigny's work, despite its unfinished condi-

tion, was to interest naturalists for a long time.

The very short explanations by Audouin were

later completed, first by Issel (1869), then by

Pallary (1926). But many other authors referred

to Savigny, among them, Jonas (1846), Pfeiffer

(1846), Vaillant (1865), P. Fischer (1865, 1870,

1871), von Martens (1866), Tapparone-Carnefri

(1875), Jousseaume (1888). Many of the speci-

mens depicted by Savigny have therefore be-

come the types of new species, which are

enumerated below.

The Collection

From Egypt, Savigny brought back a rich col-

lection of birds, fishes, mammals, insects and

shells. Included were also mollusks in alcohol,

given by Savigny to Cuvier in 1802; later on

these were said to be deposited in the Gallery of

Anatomy but we have been unable to trace

them.

In 1853, the manuscripts, notes and collec-

tions of Savigny were offered to the town of

Versailles. Curation was entrusted to the

"Societe des sciences naturelles et medicales de

Seine et Oise". Deshayes once planned to publish

a report on the Mollusca collection but never did

(Landrin, 1865). In 1864, the Society realized its

inability to curate the material properly and

asked for help from the Museum. Valenciennes

and Gratiolet were contacted, but the Society

and the Museum lacked the sum of 2,500 francs

to fund the naturalist who intended to work at

Versailles. The collection then sank into obli-

vion. In 1919, the council of the "Societe des

Sciences naturelles" was disbanded because of

World War I. The librarian of the Versailles

library took this opportunity to transfer the col-

lection in a very rough fashion to a cellar in

order to gain some space.

In 1926, when Pallary wrote his "Explication

des Planches", the vellums and collections of

Savigny were considered to be lost. In April

1927, he visited his friend the bryozoologist,

Canu, in Versailles and realized that some shells

in the cellar of the municipal library matched

exactly some depicted on Savigny's plates. Be-

tween August and September, he carefully

studied the shells which were finally to join the

national collections in 1930, together with the 5

volumes of vellums.

Pallary (1931, 1932, 1934) produced a very

detailed history of the expedition and the collec-

tions together with a biography of Savigny. He

evidently had planned to publish a report on the

collection after he had rediscovered it, but only a

small part was published (1932); this concerned

only the shells not depicted in the plates.

We are therefore producing here the latest

chapter of this two centuries-old story with the

realization of the nomenclatorial importance of

this material.

Acknowledgements

We are grateful to the librarians who helped

us during this work and to A. Foubert for

Figures 1-51, Mrs. Guillaumin, Centre de

Microscopie du C.N.R.S. for Figures 52-84 and

P. Lafaite for the colorslides of the vellums.

Figures 93-101. We wish to thank Mr. Richard

E. Petit of South Myrtle Beach, South Carolina,

for making possible the publication of the color

plate.

Checklist of the names based partly or entirely on

Savigny's plates.

It has been our intention to simply list the

material on which a name is based, when the

original description refers to the 1817 plates.

Lectotype designation is the affair of the mala-

cologist actually engaged in a process of revi-

sion of a group and should not be done for the

mere sake of it, as is too often the case in such

checklists.

We are aware of the fact that, strictly, only

the very specimen depicted on Savigny's plate

should be considered the tyyje of later authors. It

is, indeed, possible to recognize the ty^ie when it

is a large and distinct shell, but it is not so when

dealing with the many microgastropods, of

which Savigny appears to have been the earliest

collector in the Indo-Pacific.

(legyptmcii Ehrenberg, 1831 (Siuxinen) Fig. (51

Savigny 1817: pi. 2, fig. 24

Ehrenberg 18.31: (no page number). Signature E.

Name based on an unknown number of animals from

Damiette and Savigny's figure. One shell in coll. Savigny.

Ehrenberg collection in the Berlin Mu.seum may contain

specimens.

iwgyptidcd Chenu, \8A5 (Tridacna)

Savignv 1817: pi. 10, fig. 1-2

Chenu '1845: 2, pi. 7. fig. 1-2

Name based on Savigny's plate and a number of subfossil

shells from the Suez area. .Judging from the illustrations,

however. Chenu's shells have nothing to do with Savigny's,

and are tyjjical Tridacna mfuimd (Rbding).

affinis Issel, 1869 (Chiton) Fig. .') I

Savignv 1817: pi. 3, fig. 8

lsseri869: 234

Name based on Savigny's figure and seven specimens from

the Gulf of Suez, now in MGD (Genoa, Italy). One specimen

in coll. Savigny. See also sariynyi Pilsbry 1892.

(irsMomsin Issel, 1S69 (Turbo) Fig. 2.5

Savigny 1817: pi. 5, fig. 28

12 THE NAUTILUS

January 25, 1982

Vol. 96(1)

Issel 1869: 220

Name based on Savignv's figure and a subfossil shell which

could not be traced in MGU. Five shells in coll. Savigny.

audouini Jonas, 1846 (Fasciolaria) Fig. 42

Savignv 1817: pi. 4, fig. 17

Jonas i846a: 63

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

audouini Jousseaume, in Lamy, 19lH(Pristis) Fig. 86

Savignv 1817: pi. 8, fig. 11

Lamy 1918: 30

Name based on Savigny's figure and two specimens from

Suez, now in MNHN. No Savigny material left.

bacillum Issel, 1869 (Cerithium) Fig. 77

Savignv 1817: pi. 4, fig. 28

Issel 1§69: 340

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

bertholleti Issel, 1869 (Rissoina) Fig. 38

Savignv 1817: pi. 4. fig. 2

Issel 1869: 208

Name based on Savigny's figure, named by Audouin 'Rissoa

de Bertholett'. Two shells in coll. Savigny.

bourguignati Landrin, lS65(Unio) Fig. 4

Savigny 1817: pi. 2, fig. 3

Landri'n 1865: 5, fig. 1-3

Name based on Savigny's figure and specimens collected by

Savigny in Damiette (Landrin actually saw the collection).

Three syntypes in coll. Savigny.

brongnartii Audouin, \S26 (Tricoiia) Fig. 26

Savignv 1817: pi. 5, fig. 23

Audouin 1826: 41; 1828: 181

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

aiillaudi von Martens, 1866 iSpatka) Fig. 1

Savignv 1817: pi. 7, fig. 1

von Martens 1866: 9

Name based on Savigny's figure and specimens in the collec-

tions of Caillaud, Mousson, Leiden Museum and British

Museum. Two bivalve specimens and one valve in coll.

Savigny.

calloiia P. Fischer, 1 871 (Gen.a) Fig. 32

Savigny 1817: pi. 5, fig. 10

Fischer 1871: 218

Name based on Savigny's figure and specimens collected at

Suez by Gaudry, which could not be traced. Two shells in

coll. Savigny; the small one (3.2 mm) is here depicted. The

larger one (5.8 mm) is chipped and smaller than the natural

size shell depicted fig. 10,5 in Savigny.

carinata Pallarv, 1926 (Risella isseli var.)

Savigny 1817: pi. 5, fig. 34

Pallary 1926: 84

Name based on Savigny's figure and an unknown number of

shells from Suez. See under isseli.

ringulata Issel, 1869 (Eulimella) Fig. 73

Savigny 1817: pi. 3, fig. 25

Issel 1869: 182

Name based on Savignv's figure and 3 specimens from Suez,

two of which are in MGO (Genoa). One shell in coll. Savignv.

The name was changed to TurboniUa isseli by Trvoii

<1886:339) because it is supposedly congeneric with fur-

bonilla nngutatn Dunkcr 1860.

Issel 1869: 177

Name ba.sed on Savigny's figure and a specimen from Suez.

There are two shells with tnis locality m the Issel coll. in

MGD. One specimen in coll. Savigny.

c/j/ppomoru-s Jousseaume. \SSS {Clypeomorus) Fig. 40-41

Savigny 1817: pi. 4, fig. 10

.Jousseaume 1888: 171

Name based on Savigny's figure and an unknown number of

specimens from the southern Red Sea. In the Jousseaume

coll. (MNHN), there are 33 shells from Massawa and 12

shells from Obock. Two shells in coll. Savigny.

eoenobita VaiWant, IS65 (Mytilus) Fig. 12

Savigny 1817: pi. 11, fig. 3

Vaillant 1865: 115, 122

Name based on Savigny's figure and several specimens from

Suez, now in MNHN. One specimen in coll. Savigny.

mncentrica Audouin, 1826 (Doris)

Savigny 1817: Gasteropodes pi. 1, fig. 5

Audouin 1826: 14; 1828: 128

Name based on Savigny's figure. No material left.

Fig. 97

Fig. 18

corbieri JoTtus, lSi&(Chama)

Savigny 1817: pi. 14, fig. 8

Jonas 1846c: 126

Name based on Savigny's figure. Two complete specimens in

coll. Savigny.

craticulata Issel, 1SG9 (Odontostomia) Fig. 75

Savigny 1817: pi. 3, fig. 39

Issel 1869: 180

Name based on Savigny's figure and subfossil specimens,

which could not be traced in MGD (Genoa). One shell in coll.

Savigny.

ruvieri Audouin, l&2G(Emargmula)

Savigny 1817: pi. 1, fig. 9

Audouin 1826: 27; 1828: 152

Name based on Savigny's figure. No material left.

Fig. 89

Fig. 37

dautzenbergi Pallary, 1926 (Donovania)

Savigny 1817: pi. 4, fig. 20

Pallary 1926: 71

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

desgenettii "Risso" Audouin, \S2G (Bul.lal Fig. 55

Savignv 1817: pi. 5, fig. 6

Audouin 1826: 39; 1828: 178

Name based on Savigny's figure. Five shells in coll. Savigny.

desmarestii Audouin, 1826 (/Sissoaj

Savigny 1817: pi. 3, fig. 21

Audouin 1826: 36; 1828: 171

Name based on Savigny's figure. No material left.

Fig. 91

Fig. 74

dysmatica Issoi. \St'^>lOdontostomia)

Savigny 1817: pi. 3, fig. 36

Fig. 72

doliiformis Pallary, l92(ifPyrgulina)

Savigny 1817: pi. 3, fig. 42-43

Pallary 1926: 63

Name based on Savigny's figure and several specimens from

Suez, which could not be traced. Five shells in coll. Savigny.

dnrbignii Audouin, \S26IRissoa) Fig. 68

Savignv 1817: pi. 3, fig. 22

Audouin 1826: 36; 1828: 171

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

dftrbigvii Audouin, \?<2(>(Snssurella) Fig. 63

Siivignv 1817: pi. 5, fig. ,30

Audouin 1826: 42; 1828: 183

Name based on Savigny's figure. Five shells in coll. Savigny.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 13

doriae Issel, 1869 (Stomatella) Fig. 48

Savigny 1817: pi. 5. fig. 8

Issel 1869; 228

Name based un Savigiiy's figure and 10 specimens from

Suez, which could not tje traced in MGD. Seven shells in coll.

Savigny.

draparnaudi Audomn, IS26 (Tricolia) Fig. 46

Savigny 1817: pi. 5, fig. 19

Audouin 1826: 41; 1828: 181

Name based on Savigny's figure. Two shells in coll. Savigny.

elata Semper, in Issel, 1869(Sfa/io/a^ Fig. 84

Savigny 1817: pi. 3. fig. 15

Issel 1869: 330

Name based on Savigny's figure. Fifteen shells in coll.

Savigny.

elegans Audouin, 1S2G (Tritonia)

Savigny 1817: Gasteropodes pi. 2, fig. 1

Audouin 182?: 15; 1828: 130

Name based on Savigny's figure. No material left.

Fig. 94

Fig. 24

eroopotitanus Issel, 1&Q9 (Turbo)

Savigny 1817: pi. 5, fig. 27

Issel 1§69: 219

Name based on Savigny's figure and one specimen from Suez

which could not be traced in MGD. Six shells in coll. Savigny.

erythraea Hupe, 1SS4 (Blanivillia) Fig. 94

Savigny 1817: pi. 8, fig. 6

Hupe 1854: 223

Name based on Savigny's figure. Two valves in coll. Savigny.

erythraea Issel, 1869 (Nassa costulata var.) Fig. 44

Savigny 1817: pi. 6, fig. 4

Issel 1869: 126

Name based on Savigny's figure and specimens from Suez

which could not be traced in MGD. One shell in coll. Savigny.

erythraealsse], 1SQ9( Lticina) Fig. 20

Savigny 1817: pi. 8, fig. 8

Issel 1869: 84, pi. 1, fig. 9

Name based on Savigny's figure and 3 specimens from Suez

which could not be traced in MGD. One shell in coll. Savigny.

erythrofa Issel, 1869 (Area lactea var.)

Savigny 1817: pi. 10, fig. 7

Issel 1869: 89

Name based on Savigny's figure and 6 specimens from Suez.

Six paired valves in coll. Savigny.

favrei Landrin, 1865 (Helix) Fig. 21

Landrin 1865: 2, fig. 1-3

Name based on a shell from the Savigny coll., not depicted in

the atlas. Holotype in coll. Savigny.

ferussacii Audouin, 1826 (Scalaria) Fig. 81

Savigny 1817: pi. 3, fig. 13

Audouin 1826: 35; 1828: 169

Name based on Savigny's figure. One shell in coll. Savigny is

considered to be the holotype.

feuilletii Audouin, 1826 (Neritina) Fig. 28

"Savigny 1817: pi. 5, fig. 11

Audouin 1826: 40; 1828: 179

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

fourierii Audouin, 1S2& (Bulla) Fig. 54

Savigny 1817: pi. 5, fig. 5

Audouin 1826: 39; 1828: 178

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

Ireminvillii Audomn, lS2(\(Kist;ua) Fig 67

Savigny 1817: pi. 3, fig. 20

Audouin 1826: 36; 1828: 170

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

gennesi H. Fischer, 1901 (Claneulus) Fig 49

Savigny 1817: pi. 3, fig. 3

Fischer 1901: 123, pi. 4, fig. 11-12

Name based on Savigny's figure and one specimen from

Djibouti, now in MNHN. Five shells in coll. Savigny.

gevtiluomiana Issel, lS69(Eulima) Fig. 78

Savigny 1817: pi. 3, fig. 32

Issel 1869: 183

Name based on Savigny's figure and four shells from Suez.

There are 6 shells witfi this locality in MGD. Three .speci-

mens in coll. Savigny.

quenni Audouin. 1826 fTrico/ia^

Savigny 1817: pi. 5, fig. 24

Audouin 1826: 41; 1828: 181

Name based on Savigny's figure. Pallary argued that this is

only a color form of T. brongnarti. No material left.

girardi Audouin, 182& (Bulla) Fig. 52

Savigny 1817: pi. 5, fig. 3

Audouin 1826: 39; 1828: 178

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

hemprichi Ehrenberg, 1831 (Helix) Fig. 27

Savigny 1817: pi. 2, fig. 12

Ehrenberg 1831: Helix no. 4 (no page number)

Name based on a number of snails collected near Alexandria

and on Savigny's figure. 24 shells in coll. Savigny. The

Ehrenberg coll. is supposedly in Berlin.

hemprichi \&se\, \i>%9 (Trochus) Fig. 22

Savigny 1817: pi. 3, fig. 6

Issel 1869: 329 _

Name based on Savigny's figure. Six shells in coll. Savigny.

/wjrn'dMS Orbigny, li,2() (Octopus) Fig. 100

Savigny 1817: Cephalopodes pi. 1, fig. 2

Orbignv 1826: 144

Name based on Savigny's figure. No material left.

kumboldti Audouin, \B2() (Anatola)

Savigny 1817: pi. 5. fig. 1

Audouin 1826: 39; 1828: 177

Name based on Savigny's figure. No material left.

immaculata Audouin, 1826 (Doris)

Savigny 1817: Gasteropodes pi. 1, fig. 2

Audouin 182?: 13; 1828: 126

Name based on Savigny's figure. No material left.

Fig. 85

Fig. 93

infracostata Issel, \SS9(Riselln) Fig. 64

Savigny 1817: pi. 5, fig. 40

Issel 1869: 195

Name based on Savigny's figure and three specimens from

Suez, which could not be traced in MGD. Three shells in coll.

Savigny.

i'sseli Semper, in Issel, 1869 (Risella) Fig. 65

Savigny 1817: pi. 5, fig. 35

Issel 1869: 194

Name based on Savigny's figure and several specimens from

Suez and Zanzibar. There are 2 shells from Zanzibar with a

label in Semper's handwriting in MGD. Five shells in coll.

Savigny.

isseli Nevill & Nevill, 1875 (Marginella)

Nevill & Nevill 1875: 95

Fig. 57

14 THE NAUTILUS

January 25, 1982

Vol. 96(1)

New name for Marginelta pygmaea Issel 1869, non Sowerby

1846. See pygmaea.

isseli Trvon, XSSQ (TurhoniUa)

New name for Eulimella cingulata Issel 1869. See this

name.

tstfemicuTO Issel, 1869 ^Cardwmj F'g- 1^

Savignv 1817: pi. 9, fig. 11

Issel 1§69: 74 , , • f

Name based on Savigny's figure and several sp?cirneris from

Attaka, near Suez, which could not be traced in M(,t). One

valve in coll. Savigny.

jmnardi Audouin, 1 826 (Scalaria) Fig' '^'^

Savignv 1817: pi. 3, fig. 4

Audouin 1826: 3.5; 1828: 169 . . , u „ ■ ii

Name ba.sed on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

hi n th a Audouin , 1 826 (Cypraea) Fig. 33

Savigny 1817: pi. 6, fig. 27

Audouin 182?: 4.5; 1828: 190 . . , u „ ■ u

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

lamarckii Audouin, 1826(PMpa^ Fig. 83

Siivignv 1817: pi. 2, fig. 1

Audoufn 1826: 31; 1828: 161 . , . „ ■

Name ba.sed on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

teroii Landrin, 1 865 (Helix) Fig. 23

Landrin 1865: 4, fig. 1-4 , , ^ . . .. „

Name based on a shell collected by Savigny at the Fyra-

mides, not depicted in the atlas. Holotype in coll. Savigny.

lessepsianus Vaillant, 1865 iLUhodomus) Fig. 1 1

Savignv 1817: pi. 11, fig. 1

Vaillant 1865: 123 , • .

Name based on Savigny's figure and several specimens trom

the gulf of Suez, two of which are in MNHN. One specimen

in coll. Savigny.

Fig. 17

mareoticum Pallarv, 192S (Cardiiim)

Savigny 1817: pi. ?), fig. 10

PallarVl926: 109

Name ba.sed on Savigny's figure. One specimen in coll.

&ivigny is considered to be the holotype.

marmorata Audouin, 1 826 (Doris) Fig. 95

Savignv 1817: Gasteropodes pi. 1, fig. 7

Audoutn 1826: 14-15; 1828: 129

Name based on Savigny's figure. No material left.

marmorata Pallary, 1926fGCTta; Fig. 31

Savigny 1817: pi. 5, fig. 9

Pallarv 1926: 76

Nameoased on Savigny's figure. Four shells in coll. Savigny.

marfen-si Issel, 1869 (Alaba) Fig. 71

Savignv 1817: pi. 3, fig. 26

Issel lJi69: 206

Name based on Savigny's figure and a subfossil specimen

from the Red sea, which could not be traced in MGI). Four

shells in coll. Savigny.

rtwngii Audouin, 1826rB»/W Fig. 58

&vi'gny 1817: pi. 5, fig. 7

Audouin 1826: 39; 1828: 178

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

oblongun Audouin. \&2i\ I FJ nun ibrnnchus) Fig. 98

Savigny 1817: Gasteropodes pi. 3. fig. 1

Audouin 1826: 20-21; 1828: 140

Name based on Savigny's figure. No material left.

olivaefonni'i Issel, 1869 (Tomatina) Fig. 56

Savigny 1817: pi. 6, fig. 25

Issel 1869: 171 , , .

Name based on Savignv's figure and three specimens trom

Suez, one of which is in MGD. One shell in coll. Savigny.

perlntus Issel , 1869 (Triforis) Fig. 79

Savigny 1817: pi. 4, fig. 4

Issel 1869: 152 ,,,.., ■ ,

Name based on Savigny's figure and subfossil specimens, ot

which one is in MGD with the data 'Red sea'. One shell in coll.

Savigny. See also savignyanus.

pharaonis P. Fischer, 1871 (Area) Fig. 7

Savignv 1817: pi. 10, fig. 9

Fischer 1871: 213 , , • ,

Name based on Savigny's figure and several specimens trom

Suez which could not Be traced. One valve in coll. Savigny.

iMraonis P. Fischer, 1870 (Mytilus) Fig. 10

Savigny 1817: pi. 11, fig. 5

Fischer 1870: 169 , ■ r

Name based on Savignv's figure and several specimens trom

Suez, of which six bivalve specimens and one valve are m

MNHN. One specimen in coll. Savigny.

philippii Issel, 1869 (Cydostremn) Fig. 66

Savigny 1817: pi. 5, fig. 33

Issel 1869: 189 , . • ^ c

Name based on Savignv's figure and 6 specimens trom Suez

which could not be traced in MGD. Two shells in coll.

Savigny.

pulvis Issel, 1869 (Cerithium) ' Fig. 35

Savigny 1817: pi. 4, fig. 5

Isseri869: 1.50 , ,

Name based on Savigny's figure and one or several

specimens from Suez of which one is in MGD in Genoa. One

shell in coll. Savigny.

miqmaea Issel, 1869 (Marginella) Fig. 57

Savigny 1817: pi. 6, fig. 26

Issel 1869: 150 , c, ■ ■ r ,

Name based (with question mark) on Savigny s figure anti

one shell from Suez, now in MGD. Four shells in coll.

Savigny.

reticulata Philippi, 1853 (Scissurella) Fig. 62

Savignv 1817: pi. 5, fig. 29

Philippi 18.53: 38, pi. 6, fig. 11 ^,. , , . ,.

New name fi)r "Scissurella decussatn Orbigny. .-^ucJouin

(1826:42; 1828:183), not Orbigny, 1824. Therefore the name

is based on Savignv's figure; Phihppi also refers to

specimens from thp Red Sea collected by Hempnch and

Ehrenberg. Four shells in coll. Savigny.

richardi Audouin, 182& (Cardiuvi) Fig. 14

Savigny 1817: pi. 9, fig. 14

Audouin 1826: 51; 1828: 201

Name based on Savigny's figure. One specimen m coll.

Savigny is considered to be the holotype.

rissoi Audouin, 1826 (Tricolia) Fig. 47

Savignv 1817: pi. 5, fig. 18

Audouin 1826: 41; 1828: 181

Name based on Savigny's figure. Two shells in coll. Savigny.

rissoi Weinkauff, 18S5 (Rissoina)

Savignv 1817: pi. 4, fig. 1

Weinkauff 1885: 63. pi. 15d

Fig. 76

,. „„, ^ fig. 13

Name based on the name 'Manzelia de Risso' Audouin (1828:

171) and a shell from Mauritius. Two shells in coll. Savigny.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 15

roemeriana Issel, IS&d (Verms) Fig. 62

Savigny 1817: pi. 8, fig. 3

Issel 1869: 64

Name based on Savigny's figure and 7 valves from Suez, now

in MGD. Seven valves in coll. Savigny.

savigniana Audouin. IS26 (Bursatella) Fig. 99

Savigny 1817: Gasteropodes pi. 2, fig. 2

Audouin 1826: 17-18; 1828: 134

Name based on Savigny's figure. No material left.

saifignyana Ehrenberg, 1831 (Helix)

Savigny 1817: pi. 2, fig. 20

Ehrenberg 1831: Helix no. 9 (no page number)

Name based on three specimens collected near Alexandria

and Savigny's figure. Pallary however (1926: 47) argues that

Ehrenberg's species is different from the one on Savigny's

figure, which indeed depicts Zonites algirus (^Linne),

represented by 4 shells in coll. Savigny. If Ehrenberg s types

proved to be lost (they are supposedly in the Berlin

Museum),. the present material appears to be formally avail-

able for lectotype designation if it proves necessary for

nomenclature stability.

savignyanus delle Chiaje, 1828 (Murex)

Savigny 1817: pi. 4, fig. 4

delle C"hiaje 1828: 222. pi. 49, fig. 32-34

Name based on Savigny's figure and additional material

from southern Italy, presumably lost. One shell in coll.

Savigny. See also perlatus.

smngnyi Blainville, 1827 (Sepi'a^

Savigny 1817: Cephalopodes pi. 1, fig. 3

Blainville 1827: 285

Name based on Savigny's figure. No material left.

Fig. 101

Fig. 45

sarngnyi Deshayes, \S44 (PlarKucis)

Savigny 1817: pi. 4, fig. 29

Deshayes 1844d: pi. 109 (with two unnumbered pages of

text)

Name based on Savigny's figure and an unknown number of

specimens from Madagascar, which could not be traced.

Three shells in coll. Savigny.

savignyi Deshayes, 1844 (Purpura) Fig. 30

Savigny 1817: pi. 6, fig. 1

Deshayes 1844a: 112

Name based on Savigny's figure. A single specimen in coll.

Savigny is considered to be the holotype.

sarignyi Jor\2LS, 1S46 (Cytherea) Fig. 15

Savigny 1817: pi. 8, tig. 17

Name based on Savigny's figure. Eight bivalve specimens

and three valves in coll. Savigny.

savignyi P. Fischer, \S€i5 (Cerithium) Fig. 39

Savigny 1817: pi. 4, fig. 8

Fischer 1865: 244

Name based on Savigny's figure and several specimens from

Suez which could not be traced. Two shells in coll. Savigny.

savignyi Vaillant, 1865 (Diplodonta) Fig. 9

Savigny 1817: pi. 8, fig. 7

Vaillant 1865: 124

Name based on Savigny's figure and several specimens from

El Toueneb bank. Red sea, two of which are in MNHN. Two

specimens in coll. Savigny.

savignyi Issel, 18&9 (Litiopa) Fig. TO

Sa\'ignv 1817: pi. 3, fig. 19

Issel 1§69: 197

Name based on Savigny's figure and ten shells from Suez, of

which seven are in MGD. Ten shells in coll. Savigny.

savignyi Issel, \9i%9 (Marginella)

Savigny 1817: pi. 6, fig. 18

Fig. 88

Issel 1869: 115

Name based on Savigny's figure and a number of specimens

from Suez, of which two are in MGD. No Savigny material

left.

sanqnyi P. Fischer, 1871 iPeclunculus) Fig. 3

Savigny 1817: pi. 10, fig. 14

Fischer 1871: 219

Name based on Savigny's figure and one sjiecimen from

Suez, which could not be traced. Two shells in coll. Savigny.

savignyi Tapparone-Canefri, IHlh (Fasciolana) Fig. 43

Savigny 1817: pi. 4, fig. 14

Tapparone-Canefri 1875: 612

Name based on Savigny's figure. A single shell in coll.

Savigny is considered to be the holotype.

samgiiyi Morlet, \S7S (RingieuLa) Fig. 59

Savigny 1817: pi. 6, fig. 7"

Morlet"l878: 117, pi. 5, fig. 1

Name based on Savigny's figure and one shell from the gulf

of Suez, now in MNHlSl. Two shells in coll. Savigny.

savignyi Jousseaume, 1888 (Mesodesma) Fig. 5

Savigny 1817: pi. 8, fig. 5

Jousseaume 1888: 206

Name based on Savigny's figure and two left valves from

Cameron island. Red sea, apparently lost. One specimen in

coll. Savigny.

savignyi Pilsbry, 1892 (CaUistochiton keterodon var.)

Savngny 1817: pi. 3, fig. 8

Pilsbry 1892: 277, pi. 60, fig. 16 (copied from Savigny)

Name based on Savigny's figure. It is therefore an objective

synonym of Chiton affinis Issel 1869. which is based on the

same figure.

savignyi Monterosato, 1899 (Meleagrina) Fig. 2

Savigny 1817: pi. 11. fig. 8-9

Monterosato 1899: 392

Name based on Savigiiy's figure and many specimens from

Cyprus and Alexandria, proliably in the Museo Comunale,

Roma. Two specimens in coll. Savigny.

savignyi Pallary, 191% (Fissurella) Fig. 50

Savigny 1817: pi. 1, fig. 5; Pallary 1926: 34

Name based on Savigny's figure and .several specimens from

Suez, which could not be traced. Two shells in coll. Savigny.

.•OTr/'gwV? Pallary, l'i2f>(Gastrochaenal Fig. 13

SavignV 1817: "pi. 1, fig. 15; Pallary 1926: 39

Name based on Savigny's figure and one specimen from

Suez, which could not be traced. One bivalve specimen and 5

valves, some being fragmentary, in coll. Savigny.

savignyi ? aWaxy , 1926 (Lioconcha) Fig. 6

Savigny 1817: pi. 9, fig. 5; PallaiT 1926: 107

Name based on Savigny's figure. One shell in coll. Savigny is

considered to be the holotype.

savignyi Pallary, 1926 (Nassa) Fig. 90

Savigny 1817: pi. 6, fig. 3; Pallary 1926: 87

Name based on Savigny's figure. No material left.

savignyi Pallary, 1926 (Sculus) Fig. 92

Savigny 1817: pi. 1, fig. 10; Pallary 1926: 35

Name based on Savigny's figure. No material left.

savigniji "?\\\\vpp\" Krauss, \^i?, (Siphonaria)

Savigny, 1817: pi. 1, fig. 1; Pallary 1926:28

Krauss, 1848:61; Reeve, vol. 9, pi. 5, sp. 20.

3 syntjpes in MNHN,

seguenziana Issel, 1869 (Rissoina) Fig. 82

Savigny 1817: pi. 4, fig. 3

16 THE NAUTILUS

January 25, 1982

Vol. 96(1)

Issel 1869: 209

Name based on Savig7iy's fig:ure. A single sl^ell in coll.

Savigny is considered to be the holotype.

seynperiana Issel, lS&9(Lucina) Fig. 19

Savignv 1817: pi. 8, fig. 12

Issel lS69: 82

Name based on Savigny 's figure and one specimen from

Suez, which could not be traced in MCID. A single shell in

coll. Savigny.

sismondiana Issel, 1869 (Rissoa) Fig. 87

Savigny 1817: pi. 3, fig. ,33

Issel 1^69; 205

Name based on Savigny's figure and three shells from Suez,

two of which are now in M(jD. No Savigny material left.

sue2irasi'.s' Issel. 1869(A/(iryjnW;o; Fig. 60

Savigny 1817: pi. 6, fig. 17

Issel ito: 115

Name based on Savigny's figure and several shells from

Suez, six of which are in MOD. Si.x shells in coll. Savigny.

siigillata Jonas, 1846 (CythereaJ Fig. 8

Savigny 1817: pi. 9, fig. 3

Jonas i846a: 64

Name based on Savigny's figure and two additional shells

which could not be traced. Two bivalve specimens and one

valve in coll. Savigny.

tiberiana Issel, 1869 (Cingula) Fig. 69

Savigny 1817: pi. 3, fig. 16

Issel 1^69: 199 ^ ^

Name based on Savigny's figure and several specimens from

Suez, of which si.x are in MGD. About three dozen shells in

coll. Savigny.

tigrina .Audouin, 1826(Dorrs^

Savigny 1817; Gasteropodes pi. 1. fig. 3

Fig. 96

Audouin 1826: 13; 1828: 127

Name based on Savigny's figure. No material left.

tninrala (Ferussac m,s) Audouin, "[S'l^lPhysa) Fig. 29

Savigny 1817: pi. 2, fig. 27

Audoum 1826: 34; 1828: 166

Name based on Savigny's figure. Two shells in coll. Savigny.

There are also several shells from Syria, collectea by

Bruguiere and Olivier, in the Ferussac coll. (MNHN).

undata Pallary, 1926 (Risella isseii var.)

Savigny 1817: pi. 5. fig. 35.3

Pallary" 1926: 84 ^

NameDased on Savigny's figure and a number of shells from

Suez, which could not De traced. See under isseii.

unicolor Dautzenberg, in Pallary, 1926 (Cassis turgida var.)

Savignv 1817: pi. 6, fig. 6

Pallary 1926: 88

Name based on Savigny's figure and an additional shell in

Dautzenberg's private collection, which could not be traced

by Abbott (1968: 199). One shell in coll. Savigny.

venusta Issel, 1869 (Turbonilla) Fig. 34

Savigny 1817: pi. 3, fig. 34

Issel 1869: 175

Name based on Savigny's figure and three shells from Suez,

now in MGD. Nine specimens in coll. Savigny.

rillae Issel, 1869 (Cingula) Fig. 80

Savigny 1817: pi. 3, fig. 17

Issel 1869: 198

Name based on Savigny's figure and six shells from Suez,

four of which are in MGD. Seven shells in coll. Savigny.

villersii Audouin, IS'ZG (Bulla) Fig. 53

Savigny 1817: pi. 5, fig. 4

Audouin 1826: 39; 1828: 178

Name based on Savigny's figure. Two shells in coll. Savigny.

FIGS. 1-84. Specimens from the Snvinny Egyptian collec-

tion yww in the Musinim National il'Hi^toire Naturelle in

Paris. France.

1, Spatha caillaudi van Martens, 68 mm.

2, Melcagrina savi^nyi Monterosato, 58 mm.

3, Pectunculus savigiivi P. Fischer, 18 mm.

4, Unio bourguignati Landrin, 2.i mm.

5, Mesodesma savignyi Jousseaume, 20 mm.

6, Lioconcha savignyi Pallary, ,W mm.

7, Area pharaonis P. Fischer, J,8 mm.

8, Cytherea sugillata Jona^, 3J, mm.

9, Diplodonlii savignyi Vnillant, 28 mm.

10, Mylilus pharaonis P. Fischer, .12 mm.

11, Lithodomus lessepsianus Vaiilant, 10 mm.

12, Mylilus coenobita Vaiilant, 7 mm.

13, Ga.sirochaena savignyi Pallary, 11 mm.

14, Cardium richardi Audouin. 10 mm.

15, Gytherea savignyi Jona.s, 35 mm.

16, Cardium isthmic'um Issel, SJ, mm.

17, Cardium mareoticum Pallary, 2J, mm.

18, Chama corliierei ,)onas, .H7 mm.

19, Lucina semiieriana Issel, 4 mm.

20, Lucina erythruea Issel, 16 mm.

21, Helix favroi Landrin, 12 mm.

22, Trochus liemprielii Issel, 2 mm.

23, Helix leroii Landrin. 7 mm.

24, Turbo cr(<opolitanus Issel, ;l mm.

25, Turbo tiisinoensis Issel. J, ynm.

26, Tricoiia brongnartii .4Mrfo»(ii, I, mnL

27, Helix hemprichi Ehri-nlierg. IJ, mm.

28, Neritina feuilietii ,4 «(/.<»)«. /, mm.

29, Physa truncata Audauin, 5 mm.

30, Purpura savignyi Deshayes, iO mm.

31, Gena marmorata Pallary, Jf.7 mm.

32, Gena callosa P. Fischer, 3.2 mm.

33, Cypraea kunthii .Audouin, 22 mm.

34, Turbonilla venusta Issel. 6 mm.

35, Cerithium pulvis Issel, 3 mm.

36, Scalaria jomardi Audouin, 3 mm.

37, Donovania dautzenbergi Pallary. 5 mm.

38, Rissoina bertholleti /,s.sr/. 6' mm.

39, Cerithium savignyi P. Fischer, iO mm.

40, Clypeomorus clypeumorus jDM.s'.-JcaMme. 18 mm.

41, Clyi)eomorus clypeomorus ,/()M.s'.-;c(j»mc. 9 mm.

42, Fasciolaria audouini Jonaii. 135 mm.

43, Fasciolaria savignyi Tapparone-Canefri, 30 mm.

44, Nassa erythraea Issel, 9 mm.

45, Planaxis savignyi Deshayes, 17 mm.

46, Tricoiia draparnaudii j4M(foui>i, 6 mm.

47, Tricoiia rissoi Ai/rfouin, 7 mm.

48, Stomatella doriae Issel, 1, mm.

49, Clanculus gennesi H. Fischer, 8 mm.

50, Fissurella sasngnyi Pallary, 13 mm.

51, Chiton affinis Issel. 9 mm.

52, Bulla girardi .4Miy<min. 2.5 mm.

Xi, Bulla villersii /IwftoMin, 1.25 mm.

54, Bulla fourieri .Audouin. 2.6 mm.

55, Bulla desgenettii .Audouin, 1.8 mm.

56, Tornatina olivaeformis /.s.s-e/. 3.4 mm.

57, Marginella isseii Nerill & Nevill, H mm.

58, Bulla niongii Audouin, 1.6 mm.

59, Ringicula savignyi Morlet, 2.8 mm.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 17

18 THE NAUTILUS

January 25, 1982

Vol. 96(1)

Vol. 96(1)

January 25, 1982

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20 THE NAUTILUS

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Vol. 96(1)

Vol. 96(1)

January 25, 1982

THE NAUTILUS 21

22 THE NAUTILUS

January 25, 1982

Vol. 96(1)

60, Marginella sueziensis Isse.l, 2.95 tmn.

61, Succinea aeg>'ptiaca Ehrenberg, 6.5 mm.

62, Scissurella reticulata Philippi, 1.75 mm.

63, Scissurella dorbignii Audomn, 1.75 mm,.

64, Risella infracostata hsel, 1.15 mm.

65, Risella isseii St-niper in Issel, 1.15 mm.

66, Cyclostrenia philipii /sse/, 1.6 mm..

67, Rissoa freminvillii Audouin, 3.0 mm.

68, Rissoa dorbignii Audouin, 2.5 mm.

69, Cingula tiberiana /sse^, 1.25 mm.

70, Litiopa savignyi Issel, 2.55 mm.

71, Alaba martens"! Issel, 2.3 mm.

72, Odontostomia clysmatica Issel, 2.7 mm.

73, Eulimella ciiijjiilata Issel. 2.6 mm..

74, Pyrgulina doliiformis Paltary. 2.1 mm.

75, Odontostomia craticulata I.'isel. 2.05 mm.

76, Rissoina rissoi Weinkauff, 3.Jt mm.

77, Cerithium bacillum Issel, 3.75 mm.

78, Eulima gentiluomiana Issel. 2.6 mm.

79, Triforis perlatus Issel, i.7 mm.

80, Cingula villae Issel, 2.0 mm.

81, Scalaria {erassa.ci\ Audouin. 1.95 mm.

82, Rissoina seguenziana Issel. 2.8 mm.

83, Pupa lamarckii Audouin, 2.2. mm.

84, Scaliola elata Semper in Issel, 2.05 mm.

87

''%\^

89

■^i.^

90

FIGS. 8.5-92. Types fiffun's copied from Savigny (no type

m.aterial found in SurK/nii inlltrtion).

85, Anatola humboldti .-1 udoum, 35 mm (?).

86, Pristis audouini Jousseaume, 31 mm..

87, Rissoa sismondiana Issel, 2.5 mm.

92

88, Marginella savignyi Issel. h mm..

89, Emarginula cuvieri Audauin, 9.5 mm.

90, Nassa savignyi Pallary, 9 mm.

91, Rissoa desmarestii Audouin, 3 mm..

92, Scutus savignyi Pallary, 7 mm.

<■'■'■' ■" ■ ■ ;. i.-.m, 3.9 mm.

FIGS. 96-103. Savigny's unpublished color vellum.s of

opisthobranchs and cephalopods in Bibliotheque Centrale,

MNHN. Only black and white prints were previously

published.

>^^--^3

t

V -vi?^

96, Doris immaculata Audouin. i6 mm.

*.#

%^ ■ r f. -^

%.

97, Tritonia elegans Audouin, 50 mm.

98, Doris marmorata /I Mrfoitiw, 30 mm.

99, Doris tigrina Audouin. 22 mm.

100, Doris concentrka Audouin. 21 mm.

' )-^**&:::

^

■•i-i^'

,1 ■■ ' ap*";

102, Bursatella savignyana Audouin. 130 mm.

101, Fleurobranchus obiongus .Audouin, 30 ?«/«,

\y^

103, Octopus horridus Orhigny. mantle length 30 mm.

24 THE NAUTILUS

January 25, 1982

Vol. 96(1)

FIG. 94. Blainvillia erythraea Hnpe. 58 mm.

FIG. 95. Sepia savignyi BlainviUe, mantle length 85 mm.

From an unpublished color vellum in the Bibliotheque Cen-

trale, MNHN.

LITERATURE CITED

Abbott. R. T. 1968. The Helmet shells of the World (Cassi-

dae). Part 1. Indv-Paafic Mollusca 2(9):7-202, pi. 1-187.

Audouin, V. 1826. Explication sommaire des planches de

Mollusques de I'Egypte et de la Syrie publiees par J. C.

Savigny. Description de I'Egypte ou recueil des observa-

tions et des recherches qui ont ete faites en Egypte pend-

ant I'expedition de I'armee fran^aise, publie par les ordres

de sa majeste I'empereur Napoleon le grand. Histoire

Naturelle, Animaux invertebres l(4):7-56. Imprimerie

imperiale, Paris.

1828. (same title), 2 erne edition, dediee au roi,

22:117-212. Panckouke, Paris. [1827 on title page is in-

accurate].

BlainviUe, H. M. Ducrotay de 1827. Dictionnaire des sciences

naturelles, 48. Levrauft, Strasbourg. 572 pp.

Chenu, J. C. 1845. Illustrations Conchyliologiques, vol. 2:

Bivalves marins. Genus Tridaena. 2 pp., 8 pi. Paris.

Chiaje, S. delle 1828. Memorie sulla storia e anatomia degli

animali senza vertebre del regno di Napoli, 3:1-232, pi.

31-49. Napoli.

Deshayes, G. P. 1844a. Histoire naturelle des animaux sans

vertebres, 2eme edition, 10. Baillere, Paris. 638 pp.

, 1844b. (no title). Mag. Zool.. Mollusques: pi. 109

(with two unnumbered pages of text).

Ehrenberg, C. G. 1831. Animalia Mollusca, in: Hemprich &

Ehrenberg: Symbolae Physicae, IV: Animalia Everte-

brata. Unnumbered pages. 3 plates.

Fischer, P. 1865. Note sur les launes conchyliologiques des

deux rivages de I'isthme de Suez. Jour. Conchyt. 13:

240-248.

1870. Sur la faune conchyiiologique marine des

baies de Suez et de I'Akabah. Jour. Conchyl. 18:161-179.

1871. Sur la faune conchyiiologique marine de

la bale de Suez. Joar. Conchyl. 19:209-226.

Fischer, II. 1901. Liste des coquilles recueillies par M. de

Gennes a Djibouti et Ali-Sabieh, avec la description de

plusieurs formes nouvelles. Jour. Conchyl. 49:96-130,

pi. 4.

Hup^, H. 1854. Note sur un nouveau genre de Mollu.sque

acephale, genre Blainvillie (Blainvillia). Re^>. Mag. Zool.,

(2)6:219-224.

Issel, A. 1869. Malacologia del Mar Rosso. Ed. della Biblio-

teca Malacologica, Pisa. 387 pp., 5 pi.

Jonas, J. H. 1846a. Beitrag zur Erklarung der in der Des-

cription de I'Egypte abgebildeten, nebst Beschreibung

einiger anderer in rothen Meere und den angrenzenden

Landem lebender Mollusken. Z. Malak. 3:59-64.

1846b. Unbeschriebene Konchylien des rothen

Meeres. Z. Malak. 3:65-67.

1846c. Beitrag zur Erklarung der in der Des-

cription de I'Egypte abgebildeten . . . Z. Malak.

3:120-127.

Jousseaume, F. 1888. Description des Mollusques recueillis

par Mr. le Dr. Faurot dans la mer Rouge et le golfe d'Aden.

Mem. Soc. Zool. France 1:165-223.

Krauss, F. 1848. Die sildafrikanischen Mollusken. 140 pp.

Stuttgart.

Laissus, Y. 1973. Napoleon et rimprimerie. Description de

I'Egypte; bilan scientifique d'une expedition militaire, iyi

L'art du Livre a I'lmprimerie Nationale: 190-205. Im-

primerie nationale, Paris. 295 pp.

Lamy, E. 1918. Les Tellines de la mer Rouge (d'apres les

materiaux recueillis par M. le Dr. Jousseaume). Bull.

M2(.s. natn. hist. nat. 24:26-33.

Landrin, A. 1865. Coquilles nouvelles. Aubert, Versailles.

6 pp., 1 pi.

Martens, E. von 1866. Uebersicht der Land- und Siisswasser

Mollusken des Nil-Gebietes Ma/.aA-. Bl. 7:1-21.

Monterosato, A. di 1899. Coquilles marines de Chypre. Jour.

Conchyl. 47:392-401.

Morlet, L. 1878. Monographie du genre Ringicula Deshayes

et descriptions de quelques especes nouvelles. Jour.

Conchyl. 26:113-133, pi. 5.

Nevill, G. and H. Nevill 1875. Descriptions of new marine

Mollusca from the Indian Ocean. Jour. Asiatic Soc. Bengal

44(2):83-104, pi. 7-8.

Orbigny, A. d' 1826. Tableau methodique de la classe des

Cephalopodes. Ann. Sci. Nat. 7( separate reprint).

Pallarv, P. 1926. Explication des planches de J. C. Savigny.

Mm. Inst. Egypte ll:Viii -h 138 pp., pi. 1-18.

1931. Marie Jules-Cesar Savigny. Sa vie et son

oeuvre. Premiere partie. Mem. Inst. Egypte 17:1-106.

Deuxieme partie (1932). Ibid. 20:1-112. 'Troisieme partie

(1934). Ibid. 23:1-202.

1932. Inventaire de la collection malacologique

de Savigny. Bull. Mus. natn. hist. nat.. 2nd ser., 4(3):

313-321. ■

1933. Les sources d'information concernant les

savants et artistes de I'Expedition d'Egypte. Bull. Inst.

Egypte 15:221-228.

Philippi, R. A. 1853. Die Gattungen Delphinula, Scissurella

und Globulus. Systematisches Conenylien-Cabinet von

Martini und Chemnitz ("second" edition), 2(4):l-57, pi.

1-8. Von Bauer & Raspe, Niirnberg.

Pilsbry, H. A. 1892. Manual of Conchology 14: Polyplaco-

phofa. 350 pp., 68 pi. Philadelphia.

Savigny, J. C. 1817. Description de I'Egypte, ou recueil des

observations et des recherches qui ont ete faites en Eg^'pte

pendant I'Expedition de I'armee fran^aise, publiee par

ordre du gouvernement. Histoire naturelle. Planches, vol.

II. Imprimerie royaie, Paris.

1826. (same title), 2 eme edition, dediee au roi.

Pjinf rCouf kp P urm

Sherborn, C. D. 1897. On the Dates of the Natural History

portion of Savigny's 'Description de I'Egypte'. Proc. Zool.

Soc. Lond.. 1897:285-288.

Ta|iparone-Canefri, C. 1875. Studio monografico sopra i

Muricidi del Mar Rosso. .Ann. Mus. Civ. St. nat. Genova

7:564-640, pi. 19.

Tr>'on, G. W. 1886. Manual of Conchology 8. Philadelphia.

Vaillant, L. 1865. Recherches sur la faune malacologique

de la bale de Suez. Jour. Conchyl. 13:97-127, pi. 6.

Weinkauff, H. C. 1885. Die Gattungen Ris.'ioina und Rts.wa.

Systematisches Conchylien-Cabinet, 2nd ed., l(22):l-205,

pi. 1-25. Bauer & Raspe, Nuremberg.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 25

SHELL MICROSTRUCTURE OF CORBICULA FLUMINEA

(BIVALVIA: CORBICULIDAE)i

Clement L. Counts, III and Robert S. Prezant

College of Marine Studies

University of Delaware

Lewes, DE 19958 U.S.A.

ABSTRACT

The shell microstructure ofCorbicula tluminea was examined using scanning

electron microscopy. The interface between the periostracum and calcareous shell.

and a marginal periostracal loop, are described with a discussion of possible func-

tions. Surface morphology of shells with periostracum chemically removed re-

waled periodic concentric surface ridges. The complex crossed-lamellar

crystalline arrangement of the shell is confirmed.

Shell structure of bivalves in the genus Cor-

bicula (Miihlfeld, 1811) has been examined by

several investigators (Tsujii, I960; Toots and

Fox, 1972; Taylor et al., 1973; Mackie, 1978).

Tsujii (1960) reported that three shell layers

(periostracum, prismatic and pearl [nacreous of

other authors]) are present in C. japonica Prime,

1864, collected from brackish waters of the Tose

River at Tsu, Japan.

Taylor et al. (1973) reported that shells of Cor-

bicula flumin-ea (Miiller, 1774), C. occidens

(Deshayes, 1854), C. cuneiformis (Sowerby,

1817) and the fossil species C. cordata (Morris,

1854) are composed totally of aragonitic calcium

carbonate. Toots and Fox (1972) found that

aragonite, as is typical of this mineral through

geologic time, had metamorphosed to calcite in

fossil shells of Corbicula (Leptesthes) fracta

collected in the Upper Cretaceous strata of

Wyoming.

The outer calcareous shell layers of Corbicula

fluminea and C. occidens are arranged in a finely

crossed lamellar pattern, the inner layers are

complex crossed lamellar and an indistinct myo-

stracum occurs at the pallial line (Taylor et al.,

1973). The complex crossed lamellar layer of C.

fluminea and C. occidens consists of laths ar-

ranged in regular columns as in the Limopsacea

(Taylor et al., 1969).

'University of Delaware College of Marine Studies Contribu-

tion No. 155.

Mackie (1978) examined the shell structure

and mineralogy of Corbicula fluminea in a com-

prehensive ultrastructural study of the fresh-

water Sphaeriacea. Corbicula fluminea has the

thickest shell of the 23 species of sphaeriaceans

studied. Mackie reported that the individual

layers of the complex crossed lamellae, which

are found in all Sphaeriacea, are inclined in op-

jx)site directions. No pallial myostracum, nacre-

ous or outer prismatic layer is present. Mackie

(1978) postulated that thicker-shelled species of

sphaeriaceans have correspondingly thicker

periostraca that protect the shell from dissolu-

tion in acid waters of organically enriched

habitats in which they are found (Ingram et al.,

1953; Mackie and Qadri, 1973).

No single study has carefully examined the

ultrastructure of the interior and exterior sur-

face features of the shell of the Asiatic clam. The

present paper presents such an analysis of the

shell of Corbicula fluminea. using scanning elec-

tron microscopy. The outer surface morphology

is emphasized.

MATERIALS AND METHODS

Specimens of Corbicula fluminea were col-

lected, through the kindness of Dr. Ralph W.

Taylor, from Tygarts Creek, Carter County,

Kentucky, in October 1978. The bivalves were

first relaxed in 7% magnesium chloride or pro-

pylene phenoxytol solutions. Periostracum was

tlien removed with a 5% sodium hjTJOchlorite

26 THE NAUTILUS

January 25, 1982

Vol. 96(1)

solution. Shells were washed in distilled water,

dehydrated through 70% ethanol and placed in a

hot air oven (73°C) for 48 hours to dry. Shells

were then fractured and shell fragments were

selected from the posterior and anterior adduc-

tor muscle scars, the hinge and ligament, the

growing edge, the outer surface near the grow-

ing edge, the pallial line, and the purple inner

surface. Dried shell fragments were mounted on

aluminum SEM pin stubs with double adhesive

tape and silver paint and stored in the hot air

oven (73°C) until coated with metal.

Fractured shells with intact periostracum

were dehydrated in a series of increasing con-

centrations of ethanol solutions to 100% in

which they were stored. Periostracum was also

prepared on whole shells by fixation in 3%i

gluteraldehyde for 7 days. Shells were fractured

and fragments placed in 50% ethanol (5 min.)

and 100% ethanol (28 hrs.). All specimens were

dried in a Denton DCP-1 critical point drying ap-

paratus using carbon dioxide as a transfer

agent. Specimens were then mounted on pin

stubs.

Mounted specimens were coated with a thin

layer of gold in a Denton Vacuum Evaporator

and examined in a Philips PSEM 501 SEM.

RESULTS

A well-developed periostracum covers the

calcerous shell oi Corhicula jlmrdnea. This layer

consists of an extremely smooth external sur-

face, is internally homogenous, and averages 10

ym in thickness. It is firmly conjoined with the

underlying aragonitic crystals.

The periostracum terminates along the ven-

tral shell margin in a reflective loop before in-

FIG. 1. Pi'notitnicat loop al vetitral shell margin. Uirectiun

of growth to the top. Hortzonlal field width = i!55 piw.

gressing to the secretory mantle (Fig. 1). Thus,

upon emergence from the mantle, the periostra-

cum remains disattached from internal calcer-

ous shell and extends up to 580 \j.m beyond the

shell edge in a loop that often bends back over

the outer shell margin. Where the periostracum

closely overrides the outer shell surface, the at-

tachment is very tight and rarely is this organic

layer seen to peel away from the calcareous ex-

oskeleton.

Ultrastructural examination of fractured shell

fragments confirms the complex crossed lamel-

lar structure reported by Taylor et al. (1973) and

Mackie (1978) (Fig. 2). Bands of crystalline laths

are arranged approximately normal to each

other. Individual layers or bands have an

average thickness of 10 ^im (Fig. 3) while in-

dividual crystalline laths are approximately 0.7

\jin in diameter.

A cursory examination of the outer surface oi

shells from which periostacum has been re-

moved with sodium hj'pochlorite reveals annula-

tions or major ridges visible to the unaided eye

that are separated by fine, concentric striations

FIGS. 2-7. 2, Radially fractured shell o/ Corbicula fluminea demo iist rat iri.y complex crossed lamellar structure. Direc-

tion of growth lo the right. Horizontal field width = 900 lim. 3, Normally arranged crystalline laths composiyig the com-

plex crossed lamellar shell. Direction of growth to the right. Horizontal field width = 8 (jto- 4, Outer shell surface.

periostracum removed, showing concentric major annulationis (arrows), unequally spaced, and finer, subannular concen-

tric striae helween major annulae. Direction of growth lo the right. Horizontal field width = 5.3 mm. 5, Shell surface,

perkmlnwum removed, in fractured specivwn. Two growth zones are found in association with the major annulae (ar-

rows). Direction of growth lo the right. Horizontal field width = 2.J, mm. 6, Growth zones demonstrating rugose, perforate

zones terminating in a thick band of shell, the periphery of which apparently marks the end of a growth peri.od (A), a

smooth hand oj shell that becoines rugose, perforate, and grooved and then smooth, terminating in a small ridge (B) and a

wide band of nigi.se. perforate shell (C). Direction of growth to the left. Horizontal field uidth = .'ISO urn.. 7, Rugose, per-

forate bond oj oilier shell surface sculpture organized into radial ridges and grooves with surj'ace perforatiuris. Direction

of growth In the nght. U'jrlzonhil field width = 82 \im.

Vol. 96(1)

January 25, 1982

THE NAUTILUS 27

(Fig. 4). Under higher magnification (Fig. 5)

three major striations are often found in

association with each large concentric annula-

tion. The surface morphology of a single major

ridge system (Fig. 6) changes with growth,

showing different patterns of rugosity that ap-

Vol. 96(1)

January 25, 1982

THE NAUTILUS 28

pear to be associated with cessation of growth

and that alternate with smooth areas at the ap-

parent beginning of renewed shell growth. The

largest region of rugose shell was found in areas

preceeding an apparent growth halt (Fig. 7).

This region demonstrated an overall pattern of

perforations and grooves that are radially

oriented. The grooves were either interrupted

or anastomosing and had an average width of

1-2 fjim. Perforations within this zone (Fig. 8)

were of various sizes, ranging from the ir-

regularly ovoid largest holes measuring 0.75 x

1.00 nm at their greatest dimensions, to the

smallest, more circular holes with an average

diameter of 0.5 j^m. The largest perforations

usually occurred at the bottom of the radial

grooves. The small perforations were usually

found on walls bordering grooves. No perfora-

tions penetrated very deeply into the interior of

the shell (Fig. 9). This rugosity is a natural oc-

currence on the calcareous shell of Corbicula

fiwminea and is not an artifact caused by

chemical removal of the periostracum. This is

evidenced by the alternating smooth areas.

The zone of perforated grooves terminates in

a narrow (approximately 20-30 f.im) band of

thick, radially arranged microstructures (Fig.

10). This calcareous band is continuous with the

FK.S. 8-10. o, hiny.-ijroove sculplnre ikrminstrntiHn varying Kizes of perforations. Direction of yrowth to the right.

Horhotit'it fielil u-idlh = li) nm. 9, FrnHitrc section of shell with periostracum removed. Note .luperficinl perforations.

Direction of ijrowlh to the right. Horizontal field width = S2 tim. 10. Thick, concentric hand (D) terwinatimj the ridge-

groon sralplurc. \,w shell (E) appears to be laid down from beneath this band. Direction of growth to the lejl. Horizontal

field ic'dth = I,,-; ii7,i.

Vol. 96(1)

January 25, 1982

THE NAUTILirS 29

FIG. 11. Thr 5 fjiiii ridge (B) xcpdnilutg smoolh bam! (F)

from rugose band (C) of shell. Perforation's of the. rugose band

are r}ot organized into ridge-groove patterns at this stage.

Tile sntiMith hand has few. scattered perfdrolnois. Dinclidii

of groiiili III Hie right. Hnrizniiliil field ivnllli = to jj/h.

radial-perforate zones, and has few perfora-

tions. The leading edge of this zone apparently

represents the end of a growing period.

A concentric ridge averaging 5 ytm in width

was found 110 ^m from the band of apparent

growth cessation (Fig. 6). Between these two

points, three transitional surface patterns were

found; a band of smooth surfaced shell averag-

ing 25 [im in width with diffuse perforations, a

25 fim-wide band of grooved, perforate surface

similar to that preceeding the thickened calcium

band before the apparent growth pause, and a

60 yim wide band of smooth surfaced shell that

terminates in the 5 ^^m-wide ridge.

Immediately following the 5 \xm ridge, the

shell surface becomes more rugose with perfora-

tions averaging 2-4 \xm in diameter (Fig. 11).

These perforations are heterogeneous but

gradually gain organization as they meld with

the ridge-groove pattern found later in the

growth period (Figs. 6, 7, and 8) and terminate

in the thickened surface sculpture (Fig. 10).

DISCUSSION

The periostracum tightly fills the irregular

contours of the underlying calcareous shell sur-

face as shown in Figin-e 10. Presumably the

periostracum has fully polymerized prior to the

initiation of calcium carbonate deposition on this

organic template (Waite and Wilbur, 1976), so

that this intimate association is more likely a

result of aragonitic nuclei being deposited in

rugosities of a preformed periostracal mold. In

any event, the tight fit between periostracum

and calcareous shell typically leaves an intact

outer organic layer over the aragonitic shell and

the only evidence of periostracal wear occurs

around the umbones.

The marginal periostracal looj) is an interest-

ing morphological feature whose function is

unclear. The periostracal extension might allow

free range of motion of the mantle so that upon

withdrawal of the mantle the periostracum

would easily follow. Thus upon adduction perio-

stracum would form an effective seal at the shell

edge.

The curvature of the periostracal loop, if

maintained during mineralization might offer a

mold for formation of calcareous concentric

shell ridges. Loop size corresponds well with

that of major concentric ridge size, lending sup-

port to this hypothesis. Extensive periostracal

folding along the mantle margin occurs com-

monly in many marine bivalves [for example.

Area zebni (Waller, 1980)]. "Excess" marginal

periostracum is probably a reflection of perio-

stracal growth exceeding development of cal-

careous shell. This phenomenon still requires

careful analysis.

Morton (1977) found that populations of Cor-

bicula Jluminea in Plover Cover Reservoir,

Hong Kong, spawn twice a year. He pointed out

that shell growth appeared to stop during these

episodes as energy is redirected to gonadal

development. The first spawning in the spring

of the Plover Cover population was the most in-

tensive, followed by a less intensive one in the

fall or late summer (Morton, 1977). Although

Morton (1977) reported no shell surface changes

that correspond to spawning seasons, if metabo-

lism is redirected so that only gonads are

developed and shell growth ceases, this halt

should be reflected in the surface morphology of

the shell. Fuziwara (1975, 1978) reported Cor-

hirula leana spawns biannually in response to

thermal stimuli and while portions of the gonad

ai'e mature throughout the year, the overall rate

of gametogenesis is unknown. Jones (1981)

recently reported variations in shell growth in-

crements that correspond to thermal variation

30 THE NAUTILUS

January 25, 1982

Vol. 96(1)

in the marine environment. The pattern of .sur-

face shell sculpture seen in the Tygarts Creek,

Kentucky, population may be the result of

redirected energetics during gametogenesis and

change in surface sculpture seen in Figure 10

represents the terminal period of shell growth

before gonadal development or a response of the

bivalve to fluctuations in thermal regime.

ACKNOWLEDGMENTS

We thank Dr. Ralph W. Taylor, Department

of Biological Sciences, Marshall University,

Huntington, West Virginia, for his assistance in

the collection of the bivalves used in this study.

We are also indebted to Dr. Melbourne R. Car-

riker, College of Marine Studies, University of

Delaware, Lewes, for his review of the manu-

script and Dr. Gerald L. Mackie, Department of

Zoology, University of Guelph, Ontario, for his

comments and discussion. Appreciation is also

extended to Mr. Walter H. Denny, Department

of Mechanical and Aerospace Engineering, Uni-

versity of Delaware, Newark, for his technical

assistance, Ms. Pam Palinski for her photo-

graphic help, and Ms. Linda Leidy for typing the

final versions of the manuscript.

LITERATURE CITED

Fuziwara, T. 197.S. On the reproduction of Carhiculii Icana

Prime. Venus 34(l):54-56.

1978. On the ovulation n{ Ciirhiruin lennn Prime.

VV?!«.s 37(l):22-28.

Ingram, W. M., U. G. Rallinger and A. K. Gaul'in. 19.'):i. Ke-

latitmship of Sphaerium nolidum Prime to organic pollu-

tants. Ohio J. Sci. 53(4):230-235.

Jones, D. S. 1981. Annual growth increments in shell.s of

Sjiisula soLidissima record marine temperature varia-

bility. Science 211(4778):165-166.

Mackie, G. L. 1978. Shell .structure in freshwater Sphae-

reacea (Bivalvia: Heterodonta). C'nuidian Jour. ZooL

56(l):l-6.

Mackie, G. L. and S. V . Qadri. 1973. Abundance and dis-

tribution of Mollusca in an industrialized portion of the

Ottawa River near Ottawa - Hull, Canada. Jour. Fish.

Res. Bd. Canada. 30:167-172.

Morton, B. 1977. The population dynamics of Corbicula

fluminea (Bivalvia: Corbiculidae) in Plover Cove Reser-

voir, Hong Kong. Jour. ZooL. London, 181(l):21-42.

Taylor, .J. D., W. I. Kennedy and A. Hall. 1969. The shell

structure and mineralogy of the Bivalvia. Introduction.

Nuculacea - Trigonacea. Bull. British Mus. (Nat. Hist.).

ZooL Suppl. 3. pp. 1-125.

1973. The shell structure and mineralogy of the

Bivalvia. II. Lucinacea - Clavagellacea. Conclusions. Bull.

British Mus. (Nat Hist.) ZooL. 22(9):2,55-294.

Toots, H. and J. E. Fox. 1972. Inversion of aragonite to

calcite in Corbicula (Leptesthes) fracta from upper Creta-

ceous strata of Wyoming. Contrib. Geol. 12:11-14.

Tsujii, T. 1960. Studies of the mechanism of shell and pearl

formation in molluscs. Jour. Fac. Fish.. Prefect. Univ.

M(V 5:1-70.

Waite, J. H. and K. M. Wilbur. 1976. Phenol oxidase in the

periostracum of the marine bivalve Modiolus demissus.

Jour. Exper. ZooL. 195:359-367.

Waller, T. R. 1980. Scanning electron microscopy of shell

and mantle in the order Arcoida (Mollusca: Bivalvia).

Smithsonian Cuntr. ZooL 313: 58 pp., 46 figs., 1 table.

A RELIC POPULATION OF OBOVARIA RETUSA IN THE

MIDDLE CUMBERLAND RIVER, TENNESSEE

Paul W. Parmalee and Walter E. Klippel

Department of Anthropology

University of Tennessee

Knoxville, TN 37916 '

ABSTRACT

Recovery in 1980-1981 of three specimens o/Obovaria retusa (Lamarck, 1819)

from cull piles left by commercial shelters provides additional evidence of an ex-

tant relic population of this naiad in the middle Cumberland River, Trousdale

County, Tennessee.

The fornior dir-triliution of Obovaria retusa

(Lamarck, 1819) included the Ohio, Cumber-

land, and Tennessee River systems. This mus-

sel, called by the common name Golf Stick by

Vol. 96(1)

January 25, 1982

THE NAUTILUS 31

Boepple and Coker (1912) and known variously

as Pink, Ring Pink, Ram's Horn Pink, Pink

Pigtoe or Rosebud by commercial shellers,

typically inhabits the deep stretches of a river

having swift current and a substrate composed

of coarse sand and gravel. Obovaria retusa has

been extirpated throughout most of its former

range. Stansbery (1970) stated that "A popula-

tion still living in the impounded lower Ten-

nessee [below Pickwick Dam] had apparently

not reproduced since impoundment and is ex-

pected to die out. The only known breeding

population of this once widespread species is a

small one in the Green River near Munfordville,

Kentucky." Stansbery has since commented

(Personal Communication, October 1980) that

the Green River population appears to be no

longer viable and very possibly has been entirely

eradicated. Commercial shellers still occasion-

ally take a large old relic individual from

stretches of the Tennessee River below Pick-

wick Dam, but the animal is extremely rare

judging by the limited numbers encountered.

Prior to impoundment, 0. return occurred

throughout the Cumberland River "although by

no means abundant anywhere" (Wilson and

Clark, 1914). However, Neel and Allen (1964). in

their study of the mussel fauna of the upper

Cumberland River basin before impoundment,

found it to be fairly common on the big stream

bars below the falls. 0. rptui^n was not found

during a collection made by Stansbery (1969)

after impoundment of the same general area

surveyed by Neel and Allen (1964). Tennessee

Valley Authority biologists (TVA, 1976), how-

ever, reported finding the Ring Pink (numbers

and location not given) at a sheller's cook-out

area during their mussel survey of the middle

Cumberland River in Wilson, Trousdale, and

Smith counties (CRM 270.0-305.0). Although a

few valves of 0. retusa were recovered from two

prehistoric Indian rock shelter middens (Wood-

land Period, c. 1000 B.C. - A.D. 1000) in Smith

County by Parmalee, Klippel and Bogan (1980),

no fresh specimens were taken by brailing or

found in sheller's cull and stock piles during

their three year middle Cumberland River naiad

survey in Smith County.

Examination of a series of cull and stock pile

belonging to a commercial sheller operating

from a location along the Cumberland River

(CRM 275.2), c. 6.5 km south of Hartsville,

Trousdale County, Tennessee were made No-

vember 14, 1980 and January 8 and September

17, 1981. In addition to the various piles of shell,

sorted on the basis of species, size, and/or nacre

color, several hundred naiads had been dis-

carded at the water's edge. It was in this area

that the first specimen of 0. retusa was found; it

was a very large female (in mm: length, 83;

height, 79, breadth, 50) and, by all appearances,

old with a heavily eroded umbo and slightly

deformed posterior margin (Figure 1). It is very

similar in size and apparent age to those speci-

mens taken in the Tennessee River below Pick-

wick Dam. The second specimen, another fe-

male and represented by the right valve, was

recovered January 8, 1981 from a large cull pile

of "pinks" (£'?h'pfio spp., Cyclonaias tuberculata,

Lampsilis orhiculata, Epioblasma sulcata).

Measurements (mm) of this valve are as follows:

length, 64, height, 60, estimated breadth of

paired valves, 40.

During re-examination of these abandoned

cull piles (no commercial shelling in this area

since summer 1980) on September 11, 1981, the

/ •

VU'i. 1. Sf)ecirnens o/ Obovaria retusa taken hy commercial

shelters. .>iummer 1980. in the Cumberland River. Trouxdale

County, Tennessee. Top shell 8.3 mm in length.

32 THE NAUTILUS

January 25, 1982

Vol. 96(1)

left valve of another female was found. This

specimen was slightly larger than the one repre-

sented by a right valve and measured 71 mm in

length, 63 in height, and estimated 44 mm in

breadth of paired valves. Diligent searches

failed to locate the other valves of these two in-

dividuals (Figure 1).

Admittedly three specimens or even several

do not represent a viable population, but they do

provide evidence for the continued existence of

0. retusa in the middle Cumberland River.

Although the two smallest of the three speci-

mens appear to be considerably younger than

the large individual, the outer rest-lines are

crowded and too obscure to establish even an

estimated age with any degree of certainty.

Their smaller size and the lesser extent of umbo

erosion suggests individuals somewhat younger

than the larger female, but it has been found

that in the case of certain species (e.g.

Epiobbtsma brevidens. Cyprogenia irrorafa) or

individuals that impoundment has the effect on

naiads of causing the production of extremely

heavy and thick but stunted shells (Parmalee.

Klippel, and Bogan, 1980).

The assemblage of naiad species from this sec-

tion of the river (CRM c. 273.0-278.0) where the

specimens of 0. retusa were taken exhibits some

interesting differences from the one reported by

Parmalee. Klippel, and Bogan (1980) at CRM

291.0-296.8. Although many species such as

Lampsilis orbiculata appear to occur in about

the same frequency in the stretch of river below

Hartsville as they do upstream some 16-18 km,

others, including Actinonaias ligamentina and

Ptychobranchvisfasciolare, are much rarer in oc-

currence. In contrast, Quadrula quadruLa ap-

pears in greater numbers below Hartsville. It is

of interest to note that a relic population of

Epioblasma sulcata continues to survive in

the Cumberland River below Hartsville; we sal-

vaged 38 specimens (all males) of this endan-

gered naiad from the cull piles. It is evident

from the paucity of specimens taken during the

past five years that 0. retusa continues to exist

only as a relic population in the middle Cumber-

land River and that, as Stansbery (1970) com-

mented regarding the population of this naiad in

the lower Tennessee River, it can be expected to

die out.

ACKNOWLEDGMENTS

We thank Miles Wright, Frank H. McClung

Museum, University of Tennessee, Knoxville,

for photographing the specimens in Figure 1

and Betty W. Creech for typing the manuscript.

LITERATURE CITED

Hiiepple, J. V. and R. E. Cuker. 1912. Mussel resources ol

ihe Holston and Clinch rivers of eastern Tennessei-.

Bureau of Fisheries Document No. ?65.'3-13.

Neel, Joe K. and William R. Allen. 1964. The mussel fauna

of the upper Cumberland Basin before its impoundment.

Malacologia l(.3):427-4.59.

Parmalee, Paul W., Walter E. Klippel and Arthur E. Bogan.

1980. Notes on the prehistoric and preserit status of the

naiad fauna of the middle Cumberland River, Smith Coun-

ty, Tennessee. The Nautilus 94(3):93-in5.

Stansbery, David H. 1969. Changes in the naiad fauna of

the Cumberland River at Cumberland Falls in eastern

Kentucky. The Ann^r-ican Malacol'igical l.'nion Annual

Reports for 1969. pp. 16-17.

1970. Eastern freshwater mollusks (1) the Mis-

sissippi and St. Lawrence River systems. In; Proceedings

of the Ajnerican Malafological Union Sympostum on Rare

and Endangered Mollusks. Malacologia 10(l):9-22.

Tennessee Valley Authority. 1976. Mussel fauna of the

Cumberland River in Tennessee. Unpub. Report: Div.

Environmental Planning and Div. Forestry, Fisheries, and

Wildl. Develop.

Wil.son, Charles B. and H. Walton Clark. 1914. The mussels

of the Cumberland River and its tributaries. Department

of Commerce, Bureau <if Fisheries Document No. 781.

63 pp.

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1 LIBRARY i NAUTILUS

II /\Y 1 iqfl? t Volume 96, number 2 - April 21, 1982

MAY 6 WOd j ISSN 0028-1344

!

CONTENTS Woods_Ho!MiaH:---.'

Joseph Charles Bequaert (1886-1982)

An Obituary by William J. Clench 35

Terrence J. Frest and R. Sanders Rhodes II '

Oxychilus drapamaldi in Iowa 36 I

Piet Kaas !

Ischnochitcm. dilatosculptiis, A New Species from Florida

(Polyplacophora: Ischnochitonidae) 40

Hans Bertsch and Loyal J. Bibbey I

A New Tropical Eastern Pacific Ovulidae (Gastropoda) A'andarovu^a hammesi 42

Jesus Ortea

A New Favorinus (Nudibranchia: Aeolidoidea) from the Canary Islands 45

Ralph W. Taylor and Beverly D. Spurlock

The Changing Ohio River Naiad Fauna: A Comparison of Early Indian I

Middens with Today 49 I

Richard J. Neves and Alexander V. Zale

Freshwater Mussels (Unionidae) of Big Moccasin Creek, Southwestern Virginia 52 ]

George N. Wiley, Ronald C. Circe and John W. Tunnell, Jr.

Mollusca of the Rocky Shores of East Central Veracruz State, Mexico 55

Twila Bratcher and Walter 0. Cernohorsky

Six New Species of Indo-Pacific Terebridae (Gastropoda) 61

Ralph W. Taylor '

The Freshwater Mussels (Naiads) of Big Indian Creek, a Small Southern

Indiana Tributary of the Ohio River (Bivalvia: Unionidae) 66

Jack T. Moyer, William K. Emerson and Michael Ross I

Massive Destruction of Scleractinian Corals by the Muricid Gastropod,

Drupella, in Japan and the Philippines 69

Dorothea S. Franzen

Succinea avara Say from the Southern Great Plains of the United States 82

I

Recent Deaths 68 Meetings and Shell Shows 68

34 THE NAUTILUS

April 21, 1982

Vol. 96(2)

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Vol. 96(2)

April 21, 1982

THE NAUTILUS 35

JOSEPH CHARLES BEQUAERT

1886-1982

Dr. Joseph C. Bequaert, Agassiz Professor of

Zoology emeritus of the Museum of Compara-

tive Zoology, Harvard University, died in

Amherst, Massachusetts, at the age of 95, after

a lingering illness, on January 12, 1982.

Dr. Bequaert was born in the small town of

Thourout, near Bruges, Belgium, and as a

schoolboy was interested in plants and mollusks.

In 1908 he received his Ph.D. in botany from the

University of Ghent, and he spent the rest of his

long and productive life studying many facets of

the natural world.

His first entomological field work was in the

Belgian Congo where he was Entomologist for

the Belgian Sleeping Sickness Commission from

1910 to 1912. He stayed on in Africa after the

Commission gained control of the disease by

isolating its victims, and between 1913 and 1915

he continued his botanical work, being in charge

of botanical explorations for the Belgian Colo-

nial Government. During his time in Africa he

collected mollusks, many of them new, from the

Congo River, throughout the jungles to the

mountainous Ugandan border. This material

was studied by Dr. Henry A. Pilsbry and formed

the basis of his two-volume work on the mol-

lusks of the Belgian Congo, the second volume

of which was written in collaboration with Dr.

Bequaert.

In 1916 Joe Bequaert immigrated to the

United States, and by 1921 he was a naturalized

citizen. His first position in the U.S. was at the

American Museum of Natural History as Re-

search Associate in Congo Zoology. He left the

American Museum in 1922 for Boston, where he

was first Instructor (1923-1925) and then Assis-

tant Professor in the Department of Tropical

Medicine at the Harvard Medical School,

1925-1945. In 1927 he married Frances Brown

of Ohio; they had two children, Helen and

Frank.

After his retirement from Harvard in 1956, he

sought warmer climes and went to the Univer-

sity of Texas as a Visiting Professor in biology.

In 1960 he moved to Tucson, Arizona as he was

Visiting Entomologist and Curatorial Assistant

at the University of Arizona, where he orga-

nized the mollusk collection and continued to

collect and study the little known land and fresh-

water snails of the area. His last paper was on

the "Mollusks of the Arid Southwest" with

Walter B. Miller, published in 1973.

In 1980 he and his wife Frances moved to

Amherst, Massachusetts, to be near their

daughter. Dr. Helen Holmes. He died there at

the age of 95, on January 12, 1982.

Dr. Bequaert's prolific bibliography (over 250

papers, more than 50 of them on mollusks) at-

tests to his broad background and interests. His

work includes medical research on sleeping sick-

ness and schistosomiasis, extensive molluscan

studies, particularly of the Achatinidae and

Strophochelidae, many works on wasps and

other insects, botanical papers, and even a

paper on the use of the fauna of putrefaction to

determine time of death. He was a member of

many scientific societies, president of the

American Malacological Union in 1954, and an

Honorary Associate in Entomology and Mala-

cology at the Museum of Comparative Zoology

from the time of his retirement. Dr. Bequaert,

affectionately called Uncle Joe by the many

young students then at the MCZ, was available

to advise and help at all times. He was an in-

credible linguist (said he grew up on the streets

of Belgium speaking four languages) and was

always willing to help translate difficult

passages in any of the European languages.

Dr. Bequaert's contribution to malacology and

to the Mollusk Department of the MCZ will be

appreciated by students for many generations.

Memorial contributions to support research in

land molluscan systematics may be given to

either the Friends of the Department of Mol-

lusks, Museum of Comparative Zoology, Har-

vard University, Cambridge, MA 02138 or the

American Malacological Union, 3706 Rice

Boulevard, Houston, Texas 77005.

William J. Clench, Curator Emeritus

Department of Mollusks

Museum of Comparative Zoology

36 THE NAUTILUS April 21, 1982

OXYCHILUS DRAPARNALDI IN IOWA

Vol. 96(2)

Terrence J. Frest and R. Sanders Rhodes II

Department of Geology

University of Iowa

Iowa City, lA 52242

ABSTRACT

Feral and greenhouse colonies o/Oxychilus draparnaldi (Beck, 1837) in Iowa

City, Johnson Co., Iowa constitute the first Iowa record for an introduced land

snail. The species has now been reported as feral in 13 states and in greenhouses

in 13 states and the District of Columbia. The feral colony was established in or

prior to 1966, occupies approximately 2,U00 sq. ft., and contains several hundred

individuals. Differences in land snail faunas in adjacent areas with and without

0. draparnaldi, plus feeding experiments, suggests that 0. draparnaldi reduces

local populations of some land snail species.

Records of introduced mollusks in Iowa are

extremely rare. Barnhard (1978) lists two slugs

and one fresh-water snail: as far as we know this

paper constitutes the first record for an intro-

duced land snail in Iowa. In addition, the Iowa

City 0. draparnaldi occurrence is of signifi-

cance because it is the only Midwest feral colony

and because some information as to date of in-

troduction (and hence rate of spread) is

available.

In the course of a systematic survey of the

land snails of Johnson County, Iowa in 1976-78

a single dead specimen of 0. drapanaldi was

recovered from a drift pile along a fence in the

northwest corner of Hickory Hill Park, a city

facility in northeastern Iowa City. Subsequent

intensive searches for the species resulted in the

discovery of living specimens in July 1978 on the

west side of a ravine just southwest of the south

terminus of St. Clements Street in Iowa City

(SEV4 NWV4 SWV4 SWV4 sec. 2,T. 79 N., R. 6

W., Iowa City West IVz' topographic quadran-

gle: Fig. 1). Repeated visits to the site were

made in March - August 1979 and 1980 to collect

0. draparnaldi and define the limits of the col-

ony. Litter and drift samples were taken from

the colony and adjoining areas to assess the ef-

fects, if any, of the introduction. A laboratory

colony was maintained for one year to check

feeding preferences. Also a visit made to one

local greenhouse (Pleasant Valley Orchards and

rv-^'

'■■:'■• OAK LAN P

fMfT^Y \

FIG. 1. Portion uflow<i ( 'itij W'l si ?'/■>' topoyrtiiiliic iptadran-

<jle fihoiving location of Oxychilus draparnaldi (Beck, 1837)

colony (black dot). Scale bar (lower left) length: 0.3 km (ap-

proximately 0.2 mi.).

Nursery, Inc., 1302 S. Gilbert) turned up addi-

tional 0. dra;panialdi.

Three species of Oxychilus {allarius, cellarius

and draparnaldi) are common introductions in-

to the United States. Hanna (1966, p. 19) was of

the opinion that the three "cannot be separated

consistently from shell characters", but Burch

(1960, 1962) and Ellis (1969) suggest distin-

guishing shell features as well as soft body

characters enabling relatively easy discrimina-

tion of adults of all three. Adult allarius are

small (6-7 mm diameter), the spire is nearly flat,

and the animal is very dark. The larger cellarius

Vol. 96(2)

April 21, 1982

THE NAUTILUS 37

(pa. 9 mm diameter) has a more lunate aperture,

higher spire, and the animal is pale gray. 0.

drapamaldi is the largest of the three (diameter

12-16.5 mm), has a flattened spire and aperture

as in allarius but the animal is blue-gray. Our

specimens (Fig. 2) fit most readily into drapar-

naldi: mean diameter of the 10 largest speci-

mens is 11.8 mm, the largest (Fig. 2) is 13.2 mm

in diameter, and the body is blue-gray. As a fur-

ther check the Iowa City Oxychilus were com-

pared with authenticated European allarius

and cellarius provided by C. R. C. Paul (Liver-

pool University). Available material (43 speci-

mens) shows little variation in shell, animal col-

or, or spire height.

Occurrence and Ecology

The feral colony occupies a 200 ft. stretch of

overgrown steep slope (height 10-15 ft.: area

about 2400 sq. ft.) on the west side of a small

ravine (Fig. 1). Abundance seems to fluctuate

yearly even under seemingly identical weather

and moisture conditions: our best estimate is

that a minimum of 400 individuals are normally

present. Abundance also varies widely on the

slope with individuals most common under

debris near its top. Because the area formerly

was a brickyard the present slope configuration

is in part artificial. It was still in operation as

late as 1935, when the site was collected for

Pleistocene (Peoria Loess) land snails

(Cameron, 1935). After abandonment the site

was largely filled with mixed junk and earth and

housing constructed over it. According to the

present landowner, filling ceased in 1966.

Among the last materials dumped were a con-

siderable quantity of discarded greenhouse

stock, including "spoiled" tulip bulbs. It is very

likely that the snails were introduced with this

material. Following cessation of dumping the

slope has been little disturbed, and much of the

trash is now overgrown by weeds and fast-grow-

ing trees, e.g. elms and cotton woods. At pre-

sent, the site is moist and shaded into late sum-

mer, but leaf litter and ground cover are sparse.

A 2-liter litter sample and hand collecting shows

that the site provides adequate habitat for a

small land snail fauna of 11 species (Table 1).

Eggs and young have been observed in the

TABLE 1. Comparison of land snail faunas in ecologically

similar areas with and mthout Oxychilus draparnaidi (Beck,

1837) at Iowa City, Iowa.

With iMthout

draparnaidi draparnaidi

FIG. 2. Oxychilus draparnaidi iSU 1 1,9535) from Iowa City,

Iowa: respectively top, apertural, and bottom views, x3.

Scale bar length is 5 mm. Actual diameter 13.2 mm.

Al logona profunda (Say , 1321)

Angu ispira alternata (Say, 1916)

Tnodopsis multilineata (Say, IB21)

Mesodon clausus (Say, 1821)

Stenotrema f raternum (Say, 1824)

Stenotreina barbatum (Clapp, 1904'

Cat inella avara (Say. 1824)

Gastrocopta contracta (Say . 1822)

Gastrocopta pentodon (Say, 1821)

Hawai la minuscula (Bmney, 1840)

Hel icodiscus parallelus (Say, 1821)

Vertigo tndentata Wolf. 18 70

Vallonia parvula Sterki, 189J

Zcnitoides arboreus (Say, 18161

Ret inei la indentata (Say, 1823)

38 THE NAUTILUS

April 21, 1982

Vol. 96(2)

wild from April to August. Captives maintained

in a temperature-controlled room, however, laid

eggs as early as March. The species is reported

to be carnivorous by preference (Ellis, 1969, p.

247), but captive specimens would readily eat a

variety of plant and vegetable matter, as well as

other land snails. Litter sampling (2 liters) and

hand collecting of adjacent dump areas without

drapamaldi yielded 15 species. Comparison of

the two faunas (Table 1) shows considerable

overlap, but areas without drapamaldi have

several additional large species.

Feeding tests, while incomplete, indicate that

the Iowa drapamaldi are snail carnivores, and

that absence (more likely, reduced numbers: the

small sample size precludes certainty as to real

abundance of the larger species.) of some

species in areas with drapamaldi is due to selec-

tive carnivory, rather than competition for food.

Three species available in large numbers, Trio-

dopsis multilineata (Say), Anguispira altemata

(Say), and Stenotrema fratemum (Say), were

presented to drapamaldi alone, with a favorite

vegetable food (cucumber slices), in pairs, and

all together. 0. drapamaldi would eat all three

snails, but showed a marked preference for S.

fratemum. A. altemata was least favored, be-

ing often neglected in favor of cucumbers. T.

jnullilineata was eaten frequently, but when

paired with S. fratemum the latter was prefer-

red. Attempts to induce 0. drapamaldi to eat

smaller species (Z. arboreus and G. contracta)

were unsuccessful. In summary, Iowa 0. dra-

pamaldi can subsist on a variety of plant and

animal foods, and shows a moderate preference

for animal matter, particularly relishing S.

fratemum. It seems probable that absence (or

rarity) of some large native species from areas

inhabited by 0. drapamaldi is due to predation

by the introduced snail.

Because of the proximity of a city park, the

Iowa City colony will probably persist. Although

some intervening areas are presently unsuitable

for snails, maintenance of Hickory Hill Park in a

relatively undeveloped state makes available a

large area suitable for colonization. The nearest

good habitat is 0.3 mi. (0.3 km) southeast in the

same ravine (Fig. 1), and finds of occasional

drift specimens in the intervening stretch make

continued spread highly probable. The colony is

within 0.7 mi. (1.1 km) of the Iowa River valley:

the ravine in which it is situated drains into

Ralston Creek 0.6 mi. (0.9 km) down valley.

Though both waterways are locally heavily ur-

banized, both will provide adequate habitat for

0. drapamaldi. If the present rate of spread

{i.e. 200 ft. in 14 years: 15 ft/yr.) is now main-

tained), 0. drapamaldi would reach Ralston

Creek in 211 years and would require around

1000 years to reach the Iowa River via Ralston

Creek, a distance of approximately 6 mi. (13.2

km). These figures are probably conservative: to

cite a well-known example, 50 years after the

deliberate introduction of Triodopsis fosteri

(Baker) into Burlington, New Jersey by W. G.

Binney, the species had progressed a minimum

of 8 miles to Plum Point, New Jersey via the

Delaware River valley (Pilsbry, 1940), a rate of

845 ft/yr.

U. S. Distribution

Counting the Iowa records, introductions of

0. drapamaldi are reported from 13 states as

feral colonies and 13 states and the District of

Columbia as greenhouse colonies (Table 2): the

scattered literature is well summarized in

Pilsbry (1946), Hanna (1966) and Dundee (1974).

Judging from the number of greenhouse cita-

tions in states which have been surveyed in

detail {e.g. New York, 25 counties: Wurzinger,

1975, p. 37), many more feral colonies probably

exist, and the potential for continued accidental

introduction is great. The Iowa colony is signifi-

cant in that it is the first Midwest feral colony to

be reported: others are concentrated on the east

and west coasts or in long-settled areas of the

U.S. (Fig. 3). Burch (1960, p. 25-26) reports the

original distribution of 0. draparnaldi to in-

clude much of Europe, the Orkney and Shetland

Islands, the Outer Hebrides, Asia Minor, North

Africa, and Madeira. Both U. S. and foreign

distributions suggest a synanthromorph with

broad environmental tolerances making much of

the continental U. S. potential habitat. The

species does not appear to be a major agricul-

tural pest. The greatest potential for ecological

disruption seems to be in its possible tendency to

prey on native snails, combined with its poten-

Vol. 96(2)

April 21, 1982

THE NAUTILUS 39

TABLE 2. U. S. occurrences o/Oxychilus draparnaldi (Beck.

tially wide distribution. It is somewhat

analogous to the native Haplotrema concavum

in its environmental and feeding preferences;

thus the possibility of its being directly com-

petitive with the latter should be investigated.

LITERATURE CITED

Barnhart, M. C. 1978. Three introduced gastropods in Iowa.

The Nautilus 92:106-107.

Burch, J. B. 1960. Some snails and slugs of quarantine

significance to the U. S. Sterkiana 2:13-53.

1962. How to know the Eastern land snails.

Wm. C. Brown Co., Dubuque, Iowa. 214 p.

Cameron, C. C. 1935. A comparative study of the fossils of

two loess sections at North Liberty and Iowa City, Iowa.

Unpublished M.S. thesis. University of Iowa, 46 p.

Dundee, D. S. 1974. Catalog of introduced MoUusca of east-

ern North America (north of Me.xico). Sterkiana 55:1-36.

Ellis, A. E. 1969. British snails. A guide to the non-marine

Gastropoda of Great Britain and Ireland. O.xford Uni-

versity Press, London. 298 p.

Hanna, G. D. 1966. Introduced mollusks of western North

America. Occ. Papers California Acad. Sci. 48, 108 p.

Pilsbry, H. A. 1940. Land Mollusca of North America (North

of Mexico), V. 1, pt. 2. Acad. Nat. Se. Phil., Mon. 3(1):

575-994.

1946. Land Mollusca of North America (North

of Mexico), V. 2, pt. 1. Acad. Nat. Sc. Phil. Mon. 3(2):

1-520.

Wurzinger, K-Hans. 1975. The land snails of New York-

preliminary report. Sterkiana 57:33-39.

FIG. 3. County map of U. S. shou'ing localities unth feral Oxychilus draparnaldi iBeck.

1837). Note concentration on west and east coasts.

40 THE NAUTILUS

April 21, 1982

Vol. 96(2)

ISCHNOCHITON DILATOSCULPTUS, A NEW SPECIES FROM FLORIDA

(POLYPLACOPHORA: ISCHNOCHITONIDAE)

Piet Kaas

Steenvoordelaan 104, 2284 CZ Rijswijk

The Netherlands

My friend, Mr. Richard A. Van Belle, Sint-

Niklaas, Belgium, showed to me two Ischno-

chiton specimens which he had received from

collectors in Florida. One of them proved to be

correctly identified by Van Belle as Ischnochiton

(I.) pseudovirgatus Kaas, 1972. It is a fine,

7-valved, dried and flattened specimen, col-

lected by Mr. A. Crovo at Long Reef, off Elliott

Key, Florida, 18th August 1968, in the base of a

"sea-fan", and sent to Van Belle by Mr. D.

Steinke. The specimen, measuring 3.2 mm long,

1.8 mm wide, 0.6 mm high, is no. 2601 in the col-

lection of Van Belle. In all respects it is identical

to the holotype from Curasao, and the single

paratype from Aruba, Lesser Antilles.

Seven-valved specimens of Polyplacophora

are rather rare, but well-known. Acanthochitona

crinita (Pennant, 1777), and the holotype of

Callochiton septemvalvis (Montagu, 1803) are

abnormally seven-valved (vide Kaas, 1978).

The other specimen could not be identified as

it differs from all known species of Ischnochiton.

It was collected by Mr. R. Lyles in 1968, off Fort

Lauderdale Beach, Florida, in a depth of 12 m.

Although only one specimen was procured, its

characteristics are decisive enough to attach a

new name to it:

Ischnochiton dilatosculptus n. sp.

(Figs. 1-9)

Material: 1 specimen, dry. Off Fort Lauder-

dale Beach, Florida, U.S.A., 12 m. R. Lyles,

leg., now in the Rijks Museum van Natuurlijke

Historie, Leiden, Holotype 55382.

Diagnosis: Animal elongate oval, moderately

elevated, hardly carinated, angle of divergence

±110°, the valves only slightly beaked, side

slopes a little convex (Fig. 1). Head valve semi-

circular, quincuncially granulated, the anterior

margin somewhat dentated by a row of large

granules. Total length of animal: 14.5 mm;

width 8.5 mm.

Intermediate valves (Fig. 2) not sharply di-

vided into central and lateral areas as the latter

are not raised, only marked by a sculptural divi-

sion. The dorsal part of the central area is ir-

regularly granulose, more or less reticulate.

Towards the pleurae the granules rapidly in-

crease in size, forming curved longitudinal

chains, strongly converging close to the division

between central and longitudinal areas, where

they meet the chains of granules sculpturing

these parts of the valves, which are diverging

towards the anterior and side margins, thus giv-

ing the valves a fan-like, or feather-like, ap-

pearance. On the lateral areas the granules

make the sutures dentate.

Mucro of the posterior valve subcentral, not

very prominent, the back slope slightly concave.

Antemucrona! area sculptured like the central

areas of the intermediate valves, postmucronal

area like the head valve. Both areas are divided

by a rib formed by larger granules.

The articulamentum is well-developed, por-

celaneous, somewhat transparent; the color of

the tegmentum showing through; apophyses

evenly rounded, separated by a wide, bay-like

sinus; insertion plates smooth, with 8 inequidis-

tant slits in the anterior valve; 1-1 slits in the in-

termediate valves, and 11 slits in the posterior

valve; the slit-rays well-marked, the eaves solid.

The tegmentum is cream-colored, with irregu-

lar light greenish gray spots, especially towards

the side margins of the valves.

Girdle dorsally clothed with imbricating

scales, 66 ^^ wide, 48 ^i high, the base narrowly

diamond-shaped; the strongly convex dorsal

side with 8-10 strong, narrow riblets, converg-

ing towards the broadly rounded top of the

scale; interstices twice as wide as the ribs (Fig.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 41

I

Js" '-■/ .''^' .* ■(!

I -3 '""■■? '■"■■J '"? -'^ I '^$ ''* %•

0kl.- -7 -J '.4 ^4 4 I'

CRT" , . y" ■cf^^ jf' . *-'^. ,.;■■' .jv" ■ - ,

T

2_

r _. J

FIGS. 1-9. Ischnochiton dilatosculptus new species. 1, OffFort Lauderdale Beach, Fliirida. Holotype

(12 mm): 2, Left half of valve VI ( x 20): 3, Dorsal girdle scale, 66 ^ broad: 4, Ventral girdle scales (the

largest Jfg jj x 16 jj): 5, Major lateral radula tooth (side view): 6, Cusip of major lateral radula tooth: 7,

Central iimlh of radula: 8, First lateral tooth of radula: 9, Minor lateral tooth.

3). There is no marginal fringe. Ventral side of

girdle paved with radiating rows of rectangular

flat scales, about 48 x 16 f.i (Fig. 4).

Central tooth of the radula (Fig. 7) twice as

long as wide, reversed pear-shaped at the base,

distally sharply pointed, the second (major)

lateral tooth with a strongly developed triden-

tate cusp (Figs. 5, 6) and an interior projection

just below the cusp. Minor lateral tooth strongly

curved, gradually widening distally, ending in a

blunt point, non-cuspidate (Fig. 9).

Discussion: Ischnochiton (I.) dilatosculptus un-

doubtedly belongs to the group of /. (I.) striola-

fus (Gray, 1828), erythronotus (C. B. Adams,

1845), and papillosus (C. B. Adams, 1848), well

known from the Caribbean region (including

Florida and Bermuda), from which it markedly

differs in its quite peculiar sculpture, the dif-

42 THE NAUTILUS

April 21, 1982

Vol. 96(2)

ferently, more sparsely ribbed dorsal girdle

scales, and the differently shaped central

radular tooth.

It is to be hoped that more specimens will turn

up in due course, although I have the impression

that it is rather rare in this well-investigated

area.

LITERATURE CITED

Kaas, P. 1972. Polyplacophora of the Caribbean Region.

Stud. Fauna CMrafao41(137):l-165, figs. 1-247,' pis. 1-9.

(The Hague, M. Nijhoff)

1978. Notes on Loricata, 10. On the European

Callochiton species. Basteria 42:73-75.

A NEW TROPICAL EASTERN PACIFIC OVULIDAE (GASTROPODA):

XANDAROVULA HAMMESI

Hans Bertsch and Loyal J. Bibbey

Ciencias Marinas

Universidad Autonoma de Baja California

Ensenada, B.C., Mexico

In a recent review of the Ovulidae, Cate (1973)

listed 7 species from the Panamic province

(tropical west America). Since then, although

more than 25 additional new taxa of ovulids

have been proposed from I ndo- Pacific and

Caribbean waters (e.g., Azuma, 1972; Cate,

1974b, 1975, 1976a, and 1978; and Petuch,

1979), only one new species has been named

from the Panamic region (Cate, 1976b). Emer-

son and Old (1965) had previously reported the

Galapagan occurrence of Pseudocypraea adam-

sonii, an Indo-Pacific ovulid. To these tropical

eastern Pacific ovulid species we add the follow-

ing: 2 species from Panama, one known only

from the type locality and the other a widely

ranging taxon in the western Pacific:

Xandarovula hammesi

Bertsch & Bibbey, sp. nov.

(Figs. 1-6)

Description: Thin, white glossy shell, quite

large for the genus; ovalish, with both ends

pointed; bulbous in the middle; adapical terminal

protrudes sharply, completely on right-half of

shell, distinctly set off from the left side of the

bulbous body whorl; abapical terminal much

more gently narrowing along the left side;

smooth except for extremely fine longitudinal

growth striae, and faint, fine transverse line

more prominent terminally and on the columel-

la; outer lip evenly rounded, circular rather than

oval; no lip callus; apertural opening comprises

nearly 1/2 the total area of the ventral side;

aperture terminals open slightly to the side

adapically, but straight abapically (anteriorly);

anterior columellar region thin and narrow,

forming a fragile edge to the anterior gutter-

shaped siphonal canal; posterior axis delicately

tortuous, twisting a full 180° from its proximal

juncture with the body whorl to its distal ter-

mination.

Type locality: All three specimens examined

were collected in shrimp nets from about 1000

feet, off Cebaco Island (approx. 7°30'N;

81°30'W), Pacific coast of Panama, approxi-

mately in September of 1979. Holotj'pe: San

Diego Natural History Museum, Marine Inver-

Vol. 96(2)

April 21, 1982

THE NAUTILUS 43

FIGS. 1-4. Xandarovula haiiiniesi. 1 a ml 2, Holotype (SDNHM Marine Invertebrates T.S. 513). 3? mm in length. 3 and 4,

Paratype (SDNHM Marine Invertebrates T.S. 517). (Photon by Bertsch.)

have a slight callus and sulphur yellow colora-

tion along the margin of the outer lip; by con-

trast, the outer lip of X. hammesi has no

thickening and is the same white as the rest of

the shell. This new species most resembles X

figgisae Gate, 1973, but can be immediately

distinguished by the shape of the adapical

(posterior) terminal. In X. figgisae the col-

umellar portion of the terminal extension is

relatively broad and short (width greater than

1/2 the length). Comparative illustrations of

European and western VaQ\i.\c Xandarovula can

be found in Gate (1973: figs. 66 to 71) and Azuma

(1972: figs. 2 and 3; radula, fig. 7) and 1976: pit.

1, figs. 14 and 15).

Etymology: This new species honors Mr. Terry

Hammes, of Panama.

FIGS. 5 and 6. Xandarovula hammesi n. sp. Paratype in tlte

Hammes' collection. Length: 33 mm. (Photos by Bertsch)

tebrates, Type Series 513.

S.D.N.H.M., Type Series 517.

Paratype:

Disciission: This new species is separated

from the other 5 known species oi Xandarovula

by geographic location (X. patula occurs in

European waters, and the other 4 species are

western Pacific, from Japan to Australia) and

morphology. Xandarovula hammesi has an

evenly rounded outer lip, but the lip flares

adapically inX pagoda Gate, 1973, and flares (is

broader, wider) adapically in X. patula (Pen-

nant, 1777). Xandarovula xanthochila (Kuroda,

1928) and X. formosana (Azuma, 1972) both

Phenacovolva brevirostris (Schumacher, 1817)

(Figs. 7-8)

Description: Smooth pale-apricot shell, with 3

transverse brownish bands; spindle-shaped;

thick callus on outer lip, orange-colored,

straight anteriorly, angles sharply at each end;

funiculum faintly crenate.

Measurement: 20 mm x 9 mm; in the collec-

tion of the American Museum of Natural

History, Department of Invertebrates, AMNH

198612.

Locality: This specimen was live-collected

near Los Zurrones, west Panama, by Royce E.

Hubert, in 1979. Mr. William E. Old, Jr., first

44 THE NAUTILUS

April 21, 1982

Vol. 96(2)

FIC.S. 7 and 8. Fhenacovolva brevirostris from western

Panama (AMNH 198612). (Photos by Barbara Myers)

identified and recognized the significance of this

specimen.

Discussion: Two recent papers have sum-

marized the known Indo-Pacific gastropod

mollusks that occur in tropical west America

(Emerson, 1978; Bertsch, 1979). The related

family Cypraeidae has about 10 Indo-Pacific

species in the Panamic province, but this is only

the second ovulid with pan-Pacific distribution.

Previous records of the present species include

the east Asian mainland, the Japanese Ryukyus,

Philippine and Cook Islands, Celebes-Sulu Sea

(Gate, 1969:364-365), Queensland and Sydney,

Australia (Allan, 1956:132) and Hawaii (Kay,

1979:204). This, however, is the first record of

the species from the Pacific coast of the

Americas.

Detailed comparison of internal and external

(shell) anatomy will be necessary to determine

the relationship between Phenacovola

brevirostris and P. lenoreae Cardin & Walls,

1980.

ACKNOWLEDGMENTS

We thank Barbara Myers (SDNHM) for the

photographs; Dr. William K. Emerson and

William E. Old, Jr. (AMNH) for advice and the

loan of the Phenacovolva specimen, and Terry

Hammes for allowing us to examine his Xan-

daroimla specimens.

TABLE 1. Species of Ovulidae in the eastern Pacific.

Distribution Key: P signifies a Panamic species (tropical

west America); C a Califomian species, and IP indicates

Indo-Pacific distribution. Generic tisuages after Cate

(I'njal. McLean (1978). and Schilder and Sehilder (1971).

Cymbomda Cate, 1973

Cymbovula bratcherae (Cate, 1973) P

Cyphoma Roding, 1798

Cyphoma eynarginatum (Sowerby, 1830) P

Delonovolva Cate, 1973

Delonovolva aequalis aequalis (Sowerby, 1832) P

Delonovolva aequalis vidleri (Sowerby. 1881) C

Delonovolva macleani Cate. 1976 P

Jenneria Jousseaume, 1884

Jenneria pustulata (Lightfoot, 1786) P

Neosimnia Fischer, 1884

Neosimnia avena avena (Sowerby, 1832) P

Pedicularia Swainson, 1840

Pedicularia califomica Newcomb, 1864 C

Phenacovolva Iredale, 1930

Phenacovolva breinrostris (Schumacher. 1817) P & IP

Pseudocypraea Schilder, 1927

Pseudocypraea adamsonii (Sowerby, 1832) P & IP

Simnialena Cate, 1973

Simnialena inflexa (Sowerby, 1832) P

Simnialena rufa (Sowerby, 1832) P

Spiculala Cate, 1973

Spieulata barbarensis (Dall, 1892) C

Spiculata loebbeckeana (Weinkauff. 1881) C

Subsimnia Cate, 1973

Suhsimnia bellamaris (Berry, 1946) C

Xandarcnnda Cate, 1973

Xandarovula hammesi Bertsch & Bibbey sp. nov. P

LITERATURE CITED

Allan. Joyce. 1956. Cowry shells of world seas. Georgian

House, Melbourne, x + 170 pp.; 15 pis.

Azuma, Masao. 1972. Descriptions of four new gastropods

from South China Sea. Japanese Journal of Malacology,

Venus 31(2):55-61; 10 text figs. (30 July 1972)

1976. Systematic studies on the recent Japanese

family Ovulidae (Gastropoda)-IV. Genera Lacrima Cate,

1973, Globoimla Cate, 1973, Pseudosimnia Schilder,

1927 and Xandarovula Cate, 1973. Japanese Journal of

Malacology 35(3):106-117; 1 pi.; 7 text figs. (20 October

1976)

Bertsch, Hans. 1979. Tropical fauna! affinities of opistho-

branchs from the Panamic province (eastern Pacific). The

Nautilus 93(2-3):57-61. (23 April 1979)

Vol. 96(2)

April 21, 1982

THE NAUTILUS 45

Gate, Crawford Neil. 1969. Two new species of the genus

Volva Roding, 1798 (Ovulidae Fleming, 1828). The Veliger

ll(4):364-366; pi. 56. (1 April 1969)

1973. A systematic revision of the recent

Cypraeid family Ovulidae (Mollusca: Gastropoda). The

Veliger 15(Supplement):l-116; 51 pis. (31 January 1973)

1974a. The Ovulidae: a key to the genera, and

other pertinent notes (Mollusca: Gastropoda). The Veliger

16(3):307-313. (1 January 1974)

1974b. Five new species of Ovulidae from the

western Pacific (Mollusca: Gastropoda). The Veliger

16(4):381-384; 1 pi. (1 April 1974)

1975. New Cypraeacean species (Mollusca:

Gastropoda). The Veliger 17(3):255-261; 2 pis.; 1 text fig.

(1 January 1975)

1976a. Three new Cypraeacean species (Mollus-

ca: Gastropoda). The Veliger 18(4):383-384; 1 pi. (1 April

1976)

1976b. Five new species on Ovulidae (Mollusca:

Gastropoda). Th£ Veliger 19(2):159-162; 1 pi. (1 October

1976)

1978. Recently discovered new species of Ovuli-

dae, chiefly from off Wakayama Prefecture, Japan (Mol-

lusca: Gastropoda). Japanese Journal of Malacology.

Venm 37(4):191-204; 2 pis. (November 1978)

Emerson, William K. 1978. Mollusks with Indo-Pacific

faunal affinities in the eastern Pacific Ocean. The Nautilus

92(2):91-96. (27 April 1978)

Emerson. William K. and William E. Old, Jr. 1965. New

molluscan records for the Galapagos Islands. The Nautilus

78(4):116-120. (April 1965)

Iredale, Tom. 1930. Queensland molluscan notes. No. 2.

Memoirs Queensland Museum 10(l):73-88; 1 pi. (28

August 1930)

Kay. E. Alison. 1979. Hawaiian marine shells. Reef and

Shore Fauna of Hawaii. Section 4: Mollusca. Bernice P.

Bishop Museum Special Publication 64(4):xviii -i- 652 pp.;

195 text figs. (December 1979)

Kuroda, Tokubei. 1928. New Japanese Mollusca. Venus

1(1):10-15; 1 pi.

Pennant, Thomas. 1777. British Zoology. Vol. IV. Crusta-

cea, Mollusca, Testacea. Benjamin WTiite, London viii -i-

10 + 154 pp.; 93 pis.

Petuch. Edward J. 1979. New gastropods from the AbroUos

Archipelago and reef complex, Brazil. Proc. Biol. Soc.

Washington 92(3):510-526; 4 text figs. (18 October 1979)

Schilder, M. and F. A. Schilder. 1971. A catalogue of living

and fossil cowries. Taxonomy and bibliography of Trivia-

cea and Cypraeacea (Gastropoda Prosobranchia). Institut

Royal des Sciences Naturelles de Belgique Mewaries,

Deuxieme Ser. 85:246 pp. (31 July 1971)

Schumacher, C. F. 1817. Essai d'un nouveau systeme des

habitations des vers testaces. Copenhagen, pp. 1-287;

22 pis.

(Contribution number 2 from the Escuela de

Ciencias Marinas, U.A.B.C, Ensenada, B.C.,

Mexico.)

A NEW FAVORINUS (NUDIBRANCHIA: AEOLIDOIDEA)

FROM THE CANARY ISLANDS

Jesus Ortea

Department of Zoology, Oviedo University, Spain

ABSTRACT

Favorinus vitreus, a new nudibranch is described from the Canary Islands with

a discussion of other Atlantic species.

In July 1980, among material collected from

Tenerife, during a trip supported by the Junta

de Canarias and La Laguna University, we

found 40 species of Ascoglossa and Nudibran-

chia; of the latter, I collected two specimens of a

small Favorinm with two white swellings in the

rhinophores which is here described as a new

species.

Favorinus vitreus n. sp.

(Figs. 1-3)

Type locality: Los Cristianos beach (26°00'N;

16°30'W), Tenerife, Canary Islands, 23 July

1980, two specimens found on the brown algae

(Sargasum sp. and Cystoseira sp.) with small

polyzoans and spaws of an undetermined Poly-

ceridae.

46 THE NAUTILUS

April 21, 1982

Vol. 96(2)

I I white pigment

FIG. 1. Favorinus vitreus n. sp.: A, dorsal view of li ping

animal; B, rhinophores.

Holotype: I deposited in the Museum national

d'Histoire Naturelle, Paris.

Description: The two living animals are 3 mm

in length; oral tentacles, elongate, 1 mm; rhino-

phores 0.5 mm; foot 0.3 mm; and cerata up to

0.7 mm. Animals translucent white (glassy) with

the head, rhinophores, oral tentacles and cerata

white opaque. Body translucent, with an opaque

white marking in both specimens. This broad

band is discontinuous in one specimen (fig. 1,A)

and continuous in the other. Rhinophores have

two small and inconspicuous white bulbs (fig.

FIG. 2. Favorinus vitreus n. sp. Left: radular tooth: right:

jaw.

1,B). Cerata arranged in arches in the first and

second group; the third and fourth groups ar-

ranged in rows. The fifth group is a solitary

ceras. The second arch is nearly a row. The

number of cerata per a arch or row in the 3-mm-

long holotype are as follows: left side: 6, 5, 3, 2,

1; right side: 6, 4, 3, 2, 1.

We have not observed any cnidosacs in the

cerata. Liver branches in the cerata not visible

in the living animal, as the white surface of

the cerata is opaque. Liver ducts in body com-

pletely transparent. Foot transparent, with two

translucent anterior corners. Cardiac area not

prominent.

The animal did not autotomize its cerata when

it was poked with tweezers, nor when narco-

tized with magnesium chloride. There is a dorsal

indentation to the jaw (fig. 2) the masticatory

border is not complete but has several irregular

rows of pointed teeth (fig. 2,E). The radula has

17 teeth, 30-35 fjm in height. Each tooth has a

strong central cusp with four or five acutely

pointed denticles on each side (fig. 2). The penis

is unarmed.

Derivation of name: This species is called F.

vitreiis, because of the transparency of its body

(vitreo = glassy).

Discussion: The Atlantic species of Favorinus

can be artificially divided into two groups ac-

cording to the color of the rhinophores:

a) animals with brown rhinophores.

F. branchialis (Miiller) from Northern Europe

Vol. 96(2)

April 21. 1982

THE NAUTILUS 47

1 mm

FIG. 3. Atlantic species of Favorinus. E, F. branchialis.

animal from Asturias (northern of Spain): F, F. blianus,

animal from Galicia (northern Spain); G, F. auritulus,

drawing adapted from Marcus (19801: H, F. ghanensis,

dramng adapted from Edmunds (1968).

and the Mediterranean (Thompson & Brown,

1976), Morroco (Pruvot-Fol, 1953) and Cape

Verde Islands (Eliot, 1906 as F. cameus), found

also in the Canary Islands (personal observa-

tion); F. ghanensis Edmunds, from Ghana (Ed-

munds, 1968; 1974); and F. auritulus Marcus,

from the tropica! west Atlantic (Edmunds, 1964;

Marcus, 1955; Marcus & Marcus, 1963, 1970;

Marcus & Hughes, 1974 and Edmunds & Mar-

cus, 1977) possess brown rhinophores.

b) animals with white rhinophores.

F. blianus Lemche & Thompson (fig. 3,F),

from Northern Europe (British Isles and Scan-

dinavia) (Lemche & Thompson, 1974; Hunnam

& Brown, 1975), found also in the northern

Spain (Ortea & Urgorri, 1981); andF. vitreus n.

sp. from Tenerife possess white rhinophores. A

Favorinus sp. which possibly belongs to this sec-

ond group, has been reported by Eliot (1906, p.

159) from the Cape Verde Islands.

F. blianus differs from F. vitreus by having

voluminous bulbs on the rhinophores and by the

distribution of the white opaque pigment in its

body, with discontinuous patches on the cerata

and anterior corners of the foot. It also reaches

a larger size, and its radular teeth lack denticles

on the sides of the large central cusp.

Among the species with brown rhinophores,

F. ghanensis seems to be a clearly defined

species due to its penial stylet and because it

feeds upon bryozoans, an exceptional diet for an

eolid nudibranch (Edmunds, 1974). However, F.

branchialis also feeds upon Bryozoa when the

edible spawn of opisthobranchs is scarce, as we

have been able to observe in Asturias, northern

Spain, where it is frequently collected during

the winter on Bugula fastigiata and B. fulva.

When F. branchialis eats Bugula, the color of

the liver in its cerata is violet-brown, as in those

of F. ghanensis. Three rhinophoral bulbs may

also be rarely present in F. branchialis, so that,

the differences between these species is limited

to the existence of the penial stylet in F. ghanen-

sis and small details of coloration (few white

dots on the body of F. ghanensis, a dorsal band

in F. branchialis).

The difference between F. branchialis and F.

auritulus is mainly in the 3 bulbs usually present

in F. auritulus. The spawn, which may help to

separate species, is only known from F. bran-

chialis, and it consists in a regular and con-

48 THE NAUTILUS

April 21, 1982

Vol. 96(2)

centric spiral cord (Loven, 1841; Alder & Han-

cock, 1845-55; Meyer & Mobius, 1865 and

Vayssiere, 1888) with eggs of 65-70 ^^m in dia-

meter in northern Spain (personal observation),

while Haefelfinger (1962) observed eggs of

45-60 yim from a spawn from Villefranche-sur-

Mer, Vayssiere (1888) noted eggs of 50 ^^m from

the Mediterranean and Rasmussen (1951) of 70

Jim from Copenhagen (Denmark).

RESUMEN

Descripcion de una nueva especie, Favorinus

vitreus. recolectada en Tenerife, islas Canarias,

caracterizada por tener rinoforos blancos y con

dos bulbos, tentaculos orales largos (1/3 del

cuerpo) y ceratas, pigmentados uniformemente

de bianco en la superficie.

ACKNOWLEDGMENTS

I wish to thank Dr. Terrence Gosliner and the

editor, R. Tucker Abbott, for assistance with the

manuscript.

LITERATURE CITED

Alder, J. and Hancock, A. 1845-1855. A monograph of the

British Nudibranchiate Mollusca. London, Ray Society.

Edmunds, M. 1964. Eolid Mollusca from Jamaica, with

description of two new genera and three new species. Bull.

Mar. Set. Gulf. Caribbean 14:1-32.

1968. Opisthobranchiate Mollusca from Ghana.

Proc. Malac. Sor. London 38:8.3-100.

1974. An eolid nudibranch feeding on Bryozoa.

Veliger 17:269-270.

Edmunds, M. and Marcus, E. 1977. On Favorinus auritulus

Marcus and Favorinus hranchialis (Miiller). ./. moll. Stud.

43:200-201.

Eliot, C. N. E. 1906. Report upon a collection of Nudibran-

chiata from the Cape Verde Islands, with notes by C.

Crossland. Proc. Malac. Soc. London 7:131-159.

Haefelfinger, H. K. 1962. Quelques faits concernant la nutri-

tion chez Favorinua branchialis (Rathke, 1806) et Stiliger

vesicidosu.s (Deshayes, 1864). deux mollusques opistho-

branches. i?«'. Suisse Zool. 69(2):31 1-316.

Hunnam, P and Brown, G. 1975. Sublittoral nudibranch Mol-

lusca (sea slugs) in Pembrokeshire waterns. Field Studies

4:131-159.

Lemche, H. and Thompson, T. E. 1974. Three opisthobranch

gastropods new to the British fauna. Proc. Malac. Soc.

Lowrfon 41:185-193.

Loven, S. L. 1841 Bidrag till Kannedomen of Molluskernas

utveckling. K. Vet. Acad. Himdl. Stockholm for ar 1839:

227-241.

Marcus, E. 1955. Opisthobranchia from Brazil. Bol. Fac. Fit

Cien. Let., Univ. S. Paulo. Brazil, Zoologia. 20:89-261.

Marcus, E. and Marcus, E. 1963. Opisthobranchs from

the Lesser Antilles. Studies on the Fauna of Curasao

and other Caribbean islands. Stud. Fauna Curasao

19(79):l-76.

1970. Opisthobranchs from Cura(;ao and faunis-

tically related regions. Studies on the Fauna of Curasao

and other Caribbean islands. Stud. Fauna Curagcui

33:1-129.

Marcus, E. and Hughes, H. P. I. 1974. Opisthobranch Mol-

lusks from Barbados. Bull. Mar. Sci. 24:498-532.

Meyer, H. A. and Mobius, K. 1865. Fauna der Kieler Bucht.

Die Hinterkeimer oder Opisthobranchier. Leipzig. Engel-

mann l:l-xxx:l-88.

Ortea, J. and Urgorri, V. (1981). Opistobranquios nuevos

para el literal Iberico colectados en Galicia. Boletin Insti-

tuto Espanol de Oceanografia 6(288):49-60.

Pruvot-Fol, A. 1953. Etude de quelques opisthobranches de

la Cote Atlantique du Maroc et du Senegal. Trav. Ins. Sci..

Cherifien Zool. 5:1-105.

Rasmussen, E. 1951. Faunistic and biological notes on

marine invertebrates. 2. The eggs and larvae of some

Danish marine gastropods. Vidensk. Meddel. Naturh,

Foren. 113:201-249.

Thompson, T. E. and Brown, G. H. 1976. British Opistho-

branch Molluscs. Synopses of the British Fauna (New

Series) 8:1-201.

Vayssiere, A. 1888. Recherches zoologiques et anatomiques

sur les mollusques Opisthobranches du Golfe de Marseille.

Pt. 2, Nudibranches et Ascoglosses. Ann. Mus. Hist. Nat.

Marseille 3(.4yA-ieO.

PHILLIP W. CLOVER

COLLECTOR & DEALER IN

WORLD WIDE SPECIMEN

SEA SHELLS p. o. Box 83

GUn Ellen, CA 95442

FREE SHELL AND

BOOK LISTS

SPECIALISTS IN RARE

CYPRAEA, CONUS, VOLUTA

MARGINELLA, MITRA, MUREX

Specimen Shells

Offering microscopic and miniature (to '/t inch) shells

from the Florida Keys, urith accurate locality data. Also

unsorted grunge; write for list.

Margaret Teskey

P.O. Box 273

Big Pine Key. Ft. 330J,3

Vol. 96(2)

April 21, 1982

THE NAUTILUS 49

THE CHANGING OHIO RIVER NAIAD FAUNA: A COMPARISON

OF EARLY INDIAN MIDDENS WITH TODAY

Ralph W. Taylor and Beverly D. Spurlock

Department of Biological Sciences

Marshall University

Huntington, West Virginia 25701

ABSTRACT

In prehistoric times the upper Ohio River supported a population of at least 32

species of freshwater mussels. By the turn of the Twentieth Century at least seven

species had been extirpated from the river. At the present time there are only 13 of

the original 32 species still occasionally found in the river. The river habitat has

been modified by man and at least 15 species new to the river have moved in. While

the total number of individuals currently living in the river is only a small

percentage of what it was prehistorically. the total number of species has only

changed from 32 to 28.

A survey of the freshwater naiads of the upper

Ohio River was conducted by a team of biolo-

gists from Marshall University during the sum-

mer of 1979 (Taylor, 1980). This work was con-

ducted with support by the United States Army

Corps of Engineers, Huntington/Pittsburgh

Districts. As a result of the survey, it was found

that today there exists a fairly diverse naiad

faunal assemblage (composed of at least 27 resi-

dent species) in this part of the river. Upon com-

paring these data with those presented by Ort-

mann (1921) it is immediately obvious that radi-

cal changes in the faunal make-up have taken

place since the turn of the century.

Within the past 200 years man has severely

modified the river through industrial and human

pollution, damming and dredging. The Ohio

River is a much different river from the pristine

Ohio that freely flowed for eons of time prior to

the coming of European man. Native Ameri-

cans, in apparently large numbers, had lived in

harmony with the river utilizing its resources in

moderation and adding nothing which would

seriously harm or modify the river.

The habit by some American Indians of dis-

posing of their wastes in garbage pits has pro-

vided contemporary paleoecologists with a very

useful tool through which much can be learned

about the lifestyle of these primitive people. A

major component of most midden piles, along

this portion of the river, is a large amount of

well-preserved freshwater mussel shell mate-

rial. Early man used the mussel meat as a food

at least part of the year, and the shells were

ground and used as a tempering agent in the

manufacture of clay pottery. Relatively few of

the shells were incorporated into pottery; the

majority was discarded after the flesh had been

removed for human consumption. These shells

remain so well preserved that even today, using

conchological characters only, they can be read-

ily identified to species level.

Method

The initial phase of this study was a boat

survey of the freshwater mussels which present-

ly (1979) inhabit a 340-mile long portion of the

Ohio River between Pittsburgh, Pennsylvania

and Greenup, Kentucky. Collecting techniques

included brailing, handpicking in the shallows

and collecting on the banks where muskrats had

disposed of empty shells after eating the flesh.

The shells were then identified, cataloged and

accessioned into the Marshall Univeristy Mala-

cological collections.

Literature records from Parodiz (1953) and

Stansbery (1977) provided information on two

sites. We report the results, in this paper, of the

excavation of three additional sites. Figure 1

50 THE NAUTILUS

April 21, 1982

Vol. 96(2)

PENN.

BARILEn— BIRD

W. VA.

FIG. 1. Map of the upper Ohio River showing the approx-

imate location of the archeological sites bearing freshwater

naiads.

shows the respective archeological sites, while

Table 1 gives additional pertinent information.

In addition, the published works of Ortmann

(1921) and Rhoads (1899), in conjunction with

Carnegie Museum records, were used to deter-

mine the status of freshwater mussel popula-

tions in the upper Ohio River at the turn of this

century.

Results and Discussion

By combining the data derived from all five ar-

cheological studies we were able to produce a

composite list which shows that within the last

2,000-or-so years at least 32 species of mussels

have lived in this part of the Ohio. By the middle

1800's the radical modification of the Ohio River

(required for navigation) had begun, and in addi-

tion the large industrial complexes and cities

along the Ohio began using the river as a waste

disposal system. Because of these alterations

Ortmann in 1921 was unable to find seven of the

32 species used by the Indians. Those species

which were present in goodly numbers in earlier

times, but which were extinct in the river by

1900, are listed in Table 2.

The misuse and modification of the river has

continued until the present. Between the years

of 1900 and 1980 an additional 12 species have

been eliminated (Table 3). Of the original 32

species of mussels which inhabited the Ohio

River in large numbers as recently as the 1600's,

only 13 were found in the 1979 survey. Nineteen

species had been extirpated from the entire up-

per Ohio over a relatively short period of time.

The picture is not, however, as bleak as it may

seem. Although the river is still far from being

clean, steps have been taken by state and

federal agencies that will insure a continued im-

provement in water quality and thus improve

the habitat. The fact remains that while water

quality may someday improve to an acceptable

Table 1. Additional information on the archeological sites.

Site Name

Investigator

Exca\-at ion

Date

Habi tat 1 un

Date

Species Identified

7

24

32 species

Vol. 96(2)

April 21, 1982

THE NAUTILUS 51

TABLE 2. Mussel species which became extinct in the Ohio

River between 1600 and 1920.

Plethobasus cicatricosus (Say, 1829)

Pleurobema claim (Lam., 1819)

Pleurobem.a sintoxia (Raf., 1820)

Pleurobema rubrum (Raf., IS20) { = plenum)

Epioblasma Jlex-itosa (Raf., 1820)

Epioblasma torulosa (Raf., 1820)

Ptychobranchus fasciolaris (Raf., 1820)

TABLE 3. Species extirpated Jrnw the Ohio River between

1920 and 1980.

Plethobasus striatus (Raf., 1820)

Plethobasus cyphyus (Raf., 1820)

Quadrula cylindrica (Say, 1817)

Elliptio dilatata (Raf., 1820)

Cyprogenia stegaria (Raf., 1820)

Actinonaias I. carinata (Barnes, 1823)

Obovaria retusa (Lam., 1819)

Obovaria suh-otunda (Raf., 1820)

Ligumia recta (Lam., 1819)

Lampsilis abrupta (Say, 1831) i = orbiculata)

Lampsilis ovata (Say, 1817)

Obovaria olivaria (Raf., 1820)

TABLE 4. Mussel species which have established residence

in the upper Ohio River in historical times.

Anodonta imbecillus Say, 1829

Anodonta g. grandis Say, 1829

Anodonta g. corpulenta Cooper, 1834

Strophitus u. undulatus (Say, 1817)

Lasmigona costata (Raf., 1820)

Lasmigona complanata (Barnes, 1823)

Lasmigona compressa (Lea, 1829)

Qiiadrula quadrula (Raf., 1820)

Fusconaia flava (Raf., 1820)

Unionwrus tetralasmus (Say, 1830)

Leptodea fragilis (Raf., 1820)

Potamilus alatus (Say, 1817)

Potamilus ohiensis (Raf., 1820)

Toxolasma parvus (Barnes, 1823)

Villosa i. iris (Lea, 1829)

level the river is irreversibly altered as a result

of damming. The river exists today as a series of

impoundments rather than a free-flowing river.

As the river habitat became intolerable for

some naiad species, it became acceptable for

others. There is a totally new mussel fauna

presently found in the Ohio River. In addition to

the 13 remaining species of the original fauna,

there are 15 species which have established

residence within the last century (Table 4). None

of these species has been found in the archeolog-

ical material from the five test sites. While none

of these species is presently found in commer-

cially harvestable quantities, most are found in

goodly numbers throughout the entire upper

Ohio. One noteworthy example is Quadrula

quadrula. Rhoads (1899) stated that this species

did not extend above Cincinnati, but it is

presently found in many large beds throughout

the study area.

ACKNOWLEDGMENTS

We wish to express our appreciation to the

U.S. Army Corps of Engineers, Huntington/

Pittsburgh Districts, for support of parts of this

study. As always, many thanks are due Dr.

David H. Stansbery of the Ohio State University

Museum of Zoology for his help in identification

and/or confirmation of some of our specimens.

LITERATURE CITED

Ortmann, A. E. 1921. A monograph of the Naiads of Penn-

sylvania. Annals of the Carnegie Museum 8:1-384.

Parodiz, J. J. 195.5. Shell remains from the Globe Hill Site.

Appendix 2:29-30 in Myers-Oaks, W. J. 1955. Excavations

at the Globe Hill Shell Heap (46HK34-1), Hancock County,

West Virginia. West Virginia ArcheoL Soc. 3. 32 pp.

Rhoads, S. M. 1899. On a recent collection of Pennsylvania

Mollusks from the Ohio River system below Pittsburgh.

The Nautilus. 12(12):12:133-137'.

Stansben,', D. H. 1977. The Molluscan fauna from Neale's

Landing. Appendix C:l-4 in Hemmings, E. T. 1977.

Neale's Landing: An archeological study of a Fort Ancient

settlement on Blennerhassett Island. West Virginia. Un-

published manuscript on file at the Archeological Section,

W. Va. Geol. and Econ. Surv., Morgantown, West Vir-

ginia.

Taylor, R. W. 1980. A survey of the freshwater mussels of

the Ohio River from Greenup Locks and Dam to Pitts-

burgh, Pa. U.S. Army Corps of Engineers, Huntington,

West Virginia. District. 71 pp.

52 THE NAUTILUS

April 21, 1982

Vol. 96(2)

FRESHWATER MUSSELS (UNIONIDAE) OF BIG MOCCASIN CREEK,

SOUTHWESTERN VIRGINIA

Richard J. Neves

Virginia Cooperative Fishery

Research Unit'

Department of Fisheries

and Wildlife Sciences

Virginia Polytechnic Institute

and State University

Blacksburg, Virginia 24061

and

Alexander V. Zale

Florida Cooperative Fishery

and Wildlife Unit^

Newins-Ziegler Hall

University of Florida

Gainesville, FL 32611

ABSTRACT

A survey of the micssel fauna of Big Moccasin Creek, an 88-km tributary of the

North Fork of the Holston River, was conducted during 1979 and 1980. Eight

species were collected, in comparison with fourteen taken during an earlier

survey in 1915. The endangered fine-rayed pigtoe, Fusconaia cuneolus, has seem-

ingly been extirpated from this stream.

The Upper Tennessee River drainage histori-

cally contained one of the most diverse naiad

faunas in the world. Ortmann's (1918) summary

of early mussel surveys recorded an abundant

and diverse fauna in major headwater tributar-

ies, which included the Clinch, Powell, and

Holston Rivers, Virginia. Recent mussel

surveys in these rivers (Stansbery 1972; Hill et

al. 1974; Stansbery and Clench 1973, 1974;

Bates and Dennis 1978; Ahlstedt and Brown

1979; Neves et al. 1980) have demonstrated a

significant decline in species diversity and abun-

dance, due largely to habitat alteration and

water quality degradation.

The loss of mussel populations in Virginia has

been greatest in the North Fork of the Holston

River between Saltville and the Virginia-

Tennessee state line (120 km of river). The 38

mussel species that once occurred in this section

of river (Ortmann 1918) were eradicated by

chemical waste inputs from a now defunct chem-

'The Virginia Unit is jointly supported by the U.S. Fish and

Wildlife Service, the Virginia Commission of Game and In-

land Fisheries, and Virginia Polytechnic Institute and State

University.

'The Florida L'nit is jointly supported by the U.S. Fish and

Wildlife Service, the Florida Game and Freshwater Fish

Commission, and the University of Florida.

ical plant at Saltville. Big Moccasin Creek

(BMC), the largest tributary of the North Fork,

was not affected by these chemical wastes. Ort-

mann (1918) reported 14 mussel species from

two sites on BMC, including the now endan-

gered fine-rayed pigtoe, Fusconaia cuneolus.

Because the naiad fauna of this creek had not

been examined for more than 60 years, we sur-

veyed several sites on BMC during 1979 and

1980 to determine what changes in species com-

position had taken place and whether F.

cuneolus still occurred in the stream.

Study Area

Big Moccasin Creek, an 88-km tributary of the

North Fork Holston River, flows through Rus-

sell and Scott counties in southwestern Virginia

(Fig. 1). The creek has a mean discharge of 0.37

m^/s, mean gradient of 3 m/km and drains 247

km^ of Valley and Ridge Province, which is

characterized by sedimentary strata of lime-

stones, dolomites, shales, and sandstones.

Roughly 58% of the watershed is forested; most

of the remaining land is used for grazing of live-

stock and cultivation of tobacco. Upstream

water chemistry measurements during low flow

in October 1979 were as follows: temperature.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 53

FIG. 1. Stations (1-9) suri'eyed for mussels in Big Moccasin

Creek, Virginia.

14°C; pH, 8.2; dissolved oxygen, 9 mg/i; con-

ductivity, 250 fimhos; and hardness, 175 mg/i.

Methods

Seven sites on Big Moccasin Creek were sur-

veyed by at least three biologists with water-

scopes during low water flows in 1979 and 1980.

Two of these sites were at locations surveyed

earlier in this century by Ortmann (1918). Live

specimens or recently dead (lustered) shells

were recorded at each station, which included

about 0.4 km of stream bottom. Collections at

two additional sites (stations 3 and 7) sampled

during summer 1977 (Steve Ahlstedt, TVA, per-

sonal communication) are included with our sur-

vey results. Reference material is housed at

Virginia Polytechnic Institute and State Univer-

sity.

Collecting Stations

1. Above confluence with the North Fork of the Holston

River on state route 614, 2.2 km east of route 23 at Wil-

helm (river kilometer 0.0; lat. 36°36'32"N long.

82°32'40"W; Scott County). Mussels were uncommon.

2. Adjacent to a state picnic area on state route 23, Moc-

casin Gap (river kilometer 4.0; lat. 36°38'00"N, long.

82°33'10"W; Scott County). No mussels were collected.

3. Adjacent to state route 613 north of Snowflake (river

kilometer 28.8; lat. 36°41'17"N, long. 82"27'56"W; Scott

County). Mussels were uncommon.

4. McConnell Mill at intersection of state routes 613 and

687 (river kilometer 32.8; lat. 36''4r43"N, long.

82°26'56"W, Scott County). Mussels were uncommon.

5. Bridge on state route 613. 100 m east of state route 891

intersection (river kilometer 4.5.1; lat. 36°42'30"N. long.

82°23'55"W; Scott County). Mussels were uncommon.

6. Dean's Farm on state route 613, 1.9 km west of Russell-

Scott county line (river kilometer 52.6; lat. 36°43'00"N,

long. 82°21'1.5"W; Scott County). Mussels were abun-

dant.

7. Adjacent to state route 613 ju.st south of Collinwood

(river kilometer 58.4; lat. 36°44'00"N, long. 82°19'15"W;

Russell County), Mussels were uncommon.

8. Fugate's Farm on state route 613, 0,4 km south of Wil-

low Spring (river kilometer 75.6; lat. 36"46'30"N, long.

82°14'18"W; Russell County), Mussels were common.

9. Owen's Farm at the intersection of state routes 676 and

677 (river kilometer 82.6; lat. 36°47'30"N, long.

82"ir50"W; Russell County). Mussels were abundant at

this site, having an average density of 18.7 mussels/m'

(Zale and Neves 1982).

Results and Discussion

A total of eight species of freshwater mussels

were collected in Big Moccasin Creek (Table 1).

Ortmann (1918) reported 14 mussel species at

Moccasin Gap (our station 2) and two species at

Willow spring (our station 8). We collected two

species from Moccasin Gap and six species near

Willow Spring. Based on the earlier and current

collection records, the following species have

apparently been extirpated from the creek:

Ptychobranchus subtentum, Lampsilis ovata,

Dysnomia capsaejormis, Quadrula cylindrica,

Pegias fabula. Alasmidonta marginata, and

Fusconaia cuneolus. Lexingtonia dolabelloides

was not reported by Ortmann (1918), but we col-

lected this species at three stations. No recent

or relic shells of the endangered F. cuneolus

were found.

Results of this survey and water quality data

from the U.S. Geological Survey and the Vir-

ginia State Water Control Board (1976) indicate

that the lower 10 km of Big Moccasin Creek has

been subjected to water quality degradation

from urban development. At Moccasin Gap,

stream alterations resulting from road construc-

tion, housing development, and channelization

were readily apparent. The State Water Control

Board (1976) reported that treated sewage dis-

charge at Gate City and raw sewage discharge

from Weber City created severe pollution with

fecal coliform bacteria and high biological ox-

ygen demand in BMC, particularly during sum-

mer. This degradation in water quality surely

has adversely affected all downstream biota.

The diverse mussel fauna that once occurred

54 THE NAUTILUS April 21, 1982

Table 1. Checklist of mussel species collected (X) from Big Moccasin Creek, 1977-1980.

Vol. 96(2)

Mussel Species

1

Station No.

4 5 6

7

Anodontinae

Alasmidonta minor {Lea, 1845)

Lampsilinae

Lampsilis fasciola (Rafinesque, 1820)

Medionidus conradicus (Lea, 1834)

Villosa nebulosa (Conrad, 1834)

Villosa vanuxemi (Lea, 1838)

Unioninae

Fusconaia barnesiana (Lea, 1838)

Lexinqtonia dolabel loides (Lea, 1840)

Pleurobema ovi forme (Conrad, 1834)

X X X X X X X

X X X X X X X X

X XXXXXXX

XXXXX XXX

X XXXXXXX

X XX

X XXXXXXX

in lower Big Moccasin Creek has been elimi-

nated. In contrast the upper watershed has re-

mained rural, and the water quality and biota

have apparently remained essentially un-

changed over the past 70 years. Recent im-

provements in water quality below Moccasin

Gap may allow the gradual recolonization by

mussels either from upstream areas or from the

North Fork of the Holston River.

ACKNOWLEDGMENTS

We thank Lynn Russell Weaver and Jane Bain

for assisting with the mussel survey, Steve

Ahlstedt for providing the two complementary

collections, and Uavid H. Stansbery of the Ohio

State University for confirming several iden-

tifications. Funding for this work was provided

by the Virginia Commission of Game and Inland

Fisheries.

LITERATURE CITED

Ahlstedt. S. A. and S. R. Brown. 1979. The naiad fauna of

the Powell Kiver in Virpnia and Tennessee (Bivalvia;

Unionacea). Bull. Amer. Mai Union 1979:40-43.

Bates. J. M. and S. D. Dennis. 1978. The mussel fauna of the

CHnch River, Tennessee and Virginia. Sterkiana

69-70:3-23.

Hill. D. M., E. A. Taylor and C. F. Saylor. 1974. Status of

faunal recovery in the North Fork Holston River, Tennes-

see and Virg:inia. Proc. Southeust. Assoc. Game Fish

amim. 28:398-413.

Neves, R. J., G. B. Pardue, E. F. Benfield and S. D. Dennis.

1980. An evaluation of endangered mollusks in Virginia.

Va. Comm. Game Inland Fish. Proj. Rep. E-F-1. 140 p.

Ortmann, A. E. 1918. The nayades (freshwater mussels) of

the Upper Tennessee Drainage with notes on synonymy

and distribution, Proc. Amer. Phil. Soc. 57:521-626.

Stansbery, D. H. 1972. A preliminary report on the naiad

fauna of the Clinch River in the southern Appalachian

Mountains of Virginia and Tennessee (Mollusca: Bivalvia:

Unionoida). Bull. Amer. Mai. Union 1972:20-22.

Stansbery, D. H. and W. J. Clench. 1973. The Pleuroceridae

and Unionidae of the North Fork Holston River above

Saltville, Virginia. Bull. Amer. Mai. Union 1973:33-36.

Stansbery, D. H. and W. J. Clench. 1974. The Pleuroceridae

and Unionidae of the Middle Fork Holston River in Vir-

ginia. Bull. Amer. Mai. Union 1974:51-54.

Virginia State Water Control Board, 1976, Tennessee and

Big Sandy River Basins, Comprrhensiw Water Resour.

Plan Vol."V-A, Pt. 2 of 3. 298 p.

Zaie, A. V. and R. J. Neves. 1982. Reproductive biologj' of

four freshwater mussel species (Mollusca:Unionidae) in

VWirixim. Freshwater invert. Biol. 1:17-28.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 55

MOLLUSCA OF THE ROCKY SHORES OF EAST CENTRAL

VERACRUZ STATE, MEXICO

George N. Wiley, Ronald C. Circe and John W. Tunnell, Jr.

Division of Biology

Corpus Christi State University

Corpus Christi, Texas 78411

ABSTRACT

The molluscan fauna ofthePunta del Morro-Punta Delgada region, of east cen-

tral Mexico located approximately 75 km north of Veracruz, was studied during

June, 1973, August, 1976. and March and October, 1977. One hundred twenty-one

species (55 alive) consisting of 80 Gastropoda, 36 Pelecypoda, 3 Polyplacophora,

and 2 Cephalopoda were collected or observed from the volcanic rocky shores of

this region.

The lack of natural rocky shores in the north

and northwestern Gulf of Mexico excludes ex-

tensive hard-bottom shore communities. Lim-

ited fauna and flora, however, have become

established on the various man-made jetties

along the Gulf coast (Whitten, et al., 1950). The

most northern natural rocky shore areas in the

southwestern Gulf are located on the eastern

coast of Mexico. The first is a minor limestone

proturberance at Punta Jerez, Tamaulipas, and

the second is an intrusive Cenozoic volcanic out-

cropping in the Punta del Morro-Punta Delgada,

Veracruz, region (Galtsoff, 1954).

This paper is based upon the mollusks col-

lected from the Punta del Morro-Punta Delgada

region, Veracruz, Mexico (Atlantic). The pur-

pose of this investigation is to describe the rocky

shore molluscan populations of this unique Gulf

of Mexico shore community, especially noting

habitat distribution and ecological zonation.

The molluscan fauna of the southwestern Gulf

of Mexico was first reported by F. C. Baker

(1891). This expedition, under the direction of

Professor Angelo Heilprin, collected mollusks

from the beaches, islands, and shallow reefs

around Veracruz. Sixty-seven years later,

Moore (1958) collected 24 species of molluscs by

wading and snorkeling on Blanquilla Reef, lo-

cated approximately 90 km south-southeast of

Tampico. Chavez, et. al. (1970) listed 126 species

from the lagoon of Lobos Reef, about 10 km

south of Blanquilla Reef. A few species of mol-

lusks common to or characteristic of different

ecological zones at La Blanquilla Reef, located

off Veracruz, were reported by Villalobos

(1971). Tunnell (1974) conducted a comprehen-

sive study of the mollusks of Lobos Reef and

Enmedio Reef, 15 km southeast of Veracruz,

during May and June, 1973. He reported 220

species from Lobos and 219 from Enmedio (290

together). Included in his study were discussions

concerning the zoogeographical distribution and

ecological distribution of species within the dif-

ferent reef biotic zones.

Rice and Kornicker (1962 and 1965, adden-

dum) provided environmental and distributional

data for 149 species from Alacran Reef along

the outer edge of the Campeche Bank. Ekdale

(1974), who sampled the benthic invertebrate

fauna on the western side of the Yucatan Strait,

collected 316 species of molluscs. He grouped

the mollusks into five major habitat assem-

bleges, each with a set of characteristic species.

Molluscan investigations in the Caribbean

have been published from Puerto Rico and the

Virgin Islands by McLean (1951); Coomans

(1958); Usticke (1959); Warmke and Abbott

(1961); and Weber (1961). Rehder (1962) and

Work (1969) reported on Los Roques, Vene-

zuela, and Abbott (1958) on Grand Cayman.

Mollusks from the Yucatan Peninsula have

been listed by Weisbord (1926) and Jaume

(1946). Olsson and McGinty (1958) and Radwin

(1969) compiled a list of molluscs from the Carib-

56 THE NAUTILUS

April 21, 1982

Vol. 96(2)

bean coast of Panama, and Houbrick (1968) did

the same for Costa Rica.

Study Area

The Punta del Morro-Punta Delgada region

lies approximately 75 km north of Veracruz

(Fig. 1). These volcanic outcroppings are the

eastern most extension of the Volcanic Cor-

dillera which extends across Mexico from the

west coast to the east coast.

Seven prominent volcanic rocky points are

found in the Punta del Morro-Punta Delgada

region (Fig. 2). Much confusion exists concern-

ing the names of the various points. Ernesto

Chavez of the Escuela Nacional de Ciencias

Biologicas, Mexico, D.F. {pers. comm.) has pro-

vided a list of the more commonly used names of

these points. Names which are occasionally used

FIG. 1. Mapofth: ii-ealnyiGulfofMexiroshov'tngllu'Piintit

del Morro-PwnXa r>etg<ida region. Bathymetry contoured in

fathoms.

FIG. 2. Map of the Punta del Morro-Punta Delgada region

showing the seven prominent rocky points. (Adapted from:

Comision Intersecretarial Coordinadora Del Levantamiento

De La Carta Geografica De La Republica Mexicana Vera-

cruz UQ-VJ)

by either local residents or listed on other maps

are in parentheses following the names provided

by Chavez.

Extensive collecting was done exclusively at

two points: Punta del Morro (19°52'15"N,

96°27'30"W); and Punta Boquilla de Piedras

(19°51'15"N, 96°26'45"W). These points were

chosen for their accessibility and differing physi-

ographical features.

There is a large variation in available types of

habitats on these rocky shores, ranging from

steep, rough jagged boulders, which receive full

energy of the waves breaking on them (Figure 3)

to high and low tide pools (Figure 4) to shallow

calm-water areas which are commonly lined

with small, smooth rocks and pebbles located on

sandy beaches and behind larger boulders (Fig-

ure 4 and 5). The large grapsid crab Gra-psiis

Vol. 96(2)

April 21, 1982

THE NAUTILUS 57

FIG. 3. Sfeep rocky cliffs, approximately 60 m high, at Pun-

ta Los Munecos fPvnta Boca Andrea).

FIG. 4. Large, steep jagged boulders and high to low tide

pools at Punta del Morro (Punta Delgada).

grapus (Linnaeus) is commonly observed scurry-

ing over the outer rocks in the supratidal zone

and the small acorn barnacle Chthamalus

fragilis Darwin dominates the upper intertidal

to lower supratidal zones. The mid to lower in-

tertidal areas are dominated by sabellariid

polychates and luxurient algal growths, primari-

ly browns with some reds and greens. Subtidally

the rocks are covered to a great extent by the

zooanthid anemone Palythoa mamrnillosa (Ellis

and Solander), and the sea urchin Echinometra

lucunter (Linnaeus) is also very abundant.

The Punta del Morro-Punta Delgada region

has an annual temperature range of 22-26°C

'^.^

««i^ .'..'»,:a*j^

FIG. 5. Large, steep jagged, boulders, protected rocky tide

pool, a7id sandy beach at Punta del Morro in the foreground.

Low profile headland, similar to Punta Boquilla de Piedras.

in the background.

with an annual rainfall of 1200-1500 mm (Gar-

cia, 1970).

Methods

Specimens were collected and observed dur-

ing four trips to the area: June, 1973; August,

1976; and March and October, 1977.

Techniques utilized in the collection of

mollusks included: collection and examination of

sediment samples from tidepools, beaches, and

subtidal substrates for micromollusks; collecting

intertidal algae samples for associated mollusks;

gathering rock samples for boring and attached

species; snorkeling, when weather permitted,

for subtidal species; and general collecting in-

volving walking and wading, examining cracks,

crevices, tidepools, and overturned rocks.

All samples were put in Whirlpak plastic bags

and placed in buckets containing 10% formalin

for fixation. Specimens were returned to the

lab, washed with freshwater and placed in a 45%

solution of isopropanol for preservation. Sedi-

ment samples were rinsed with freshwater and

placed in an oven for 24 hours at 100°C. After

drying, micromollusks were picked from each

sample. Algae samples were rinsed over a U.S.

Standard Sieve #35 (500u), and any mollusks re-

tained were kept for identification. Algae

58 THE NAUTILUS

April 21, 1982

Vol. 96(2)

samples were also examined under a dissecting

microscope for attached species.

All mollusks reported in this paper are de-

posited in the Corpus Christi State University

museum collections.

Results

A total of 121 species of Mollusca (55 alive)

v^ere collected from the Punta del Morro-Punta

Delgada region. The 121 species represented 61

families, 87 genera, and consisted of 80 species

of gastropods, 36 species of bivalves, 3 chitons,

and 2 cephalopods. The most abundant families

of gastropods were Columbellidae (7 species),

Littorinidae (4 species), and Neritidae (4

species). Arcidae (8 species), Veneridae (4

species), Chamidae (3 species), and Isogno-

monidae (3 species) were found to be the most

abundant bivalve famiHes.

Table 1 contains a list of mollusks collected or

observed from the Punta del Morro-Punta Del-

gada region. Included in the table are notations

SYSTEMATIC LIST OF MOLLUSCS COLLECTED OR OBSERVED

FROM THE PUOTA DEL MORRO-PUNTA DELCADA REGION

Species Ecological

_^ Notation +

Class Cascropoda

Subclass Prosobranchla

Order Archaeogastropoda

Flssurellldae

*Dl_od_ora caycnensls (Lamarck, 1822) Int

*Fla8urella barbadensls (Cinclin. 1791) Int

ACRiaeldae

*Acnaea leucopleura (Cinelln, 1791) int

Trochldac

Calllostoma euglypCum (A. Adams, 1854) bd

C. Jububinum (Cinelln, 1791) bd

Tefjula fasclata (flcrn, 1778) bd

Turbinldae

Astraea tecta Llghtfoot, 1786 bd

Phaslanellidap

•Tricolia afflnis

(C.B, Adams, 1850)

Norltidae

•Serita fulgurane Cmclln. 1791

*N. tgsspUata Cmelin. 1791

■N. versicolor Cmelin. 1791

•Nerltlna virglnca (Linnaeus, 1758)

Order HesoK3"Cropoda

. Littorinidae

*Lictorlna llncolatn QrblRny, 1840

*L. nrleoRrla (Potlez and Hlchaud, 1838)

*L. nebulowa (Lamarck, 1922)

*L. ilciac (Coelln, 1791)

Vltrincllldae

Vitrtnclla floridana Pllsbry and McGlnty, 1946

Parviturbqidos IntirruptuB (C.B. Adans, 1850)

Atchitceionlcidac

Arch 1 tec tonicn noblllw Rodlng, 1709

Hejjacyj* hiBulcatuB (Orblgnv, 1842)

H. cyllndrlcus {Gmflln, 1791)

Caecldae

*'-i''£.'^ p"lchellun Stimpaon, 1851

'C. rvi'-iu'iltiim Folln, 1867

't- Ke^iJl"^ (OrblRny. IStl)

Vcrsfit Idrti^ ~

'Pttnloronchun var Una (Orbignv, 1841)

Csrlthidai!

CTlChlun IjitoBiin Mt-nke. 182R

sab

sab

Order Mytllolda

Vol. 96(2)

April 21, 1982

THE NAUTILUS 59

TABLE 1 (continued)

•Collected Alive

+Ecological notation: spt-supratidal zone, int-Intertidal zone, sbt-subtidal

zone, al-algae associated, sab-sabellariid colony, sed-sediment samples,

tp-tidepool, ob-observed, bd-beach drift (shell only)

designating whether the moUusk was collected

alive or dead, the specific collecting site or zone,

and, in some cases, a notation relating to some

ecological aspect which deserves mention. The

systematic list follows Keen (1971) except for

genera or families which are absent from her

coverage of tropical West America, in which

case Abbott (1974) was employed. The species

are arranged alphabetically within each genus.

Supratidal zone

Littorina-Nerita zonation, common in many

rocky shore areas throughout the Caribbean,

characterized the supratidal zone. Littorina

zizac and L. lineolata were commonly found on

supratidal rocks and boulders in the upper por-

tion of the supratidal zone. Littorina zizac was

usually found above L. lineolata whenever they

occupied the same substrate. Two individuals of

L. nebulosa were collected from Punta Boquilla

de Piedras but are considered uncommon. Lo-

cated between the Littorina zone and the

water's edge, Nerita fulgurans was observed in

large numbers. Nerita tessellata and A^. ver-

sicolor were also considered uncommon as only

two individuals of each species were collected

both from Punta Boquilla de Piedras in March,

1977. Planaxis nucleus was observed in close

proxhnity to A^. fulgurans in the lower portion of

the supratidal zone and upper portion of the in-

tertidal zone but was less abundant.

Intertidal zone

Fifteen species of mollusks (11 Gastropoda, 1

Pelecypoda, and 3 Polyplacophora), were col-

lected from the intertidal zone; only the subtidal

zone yielded more species (18). The intertidal

zone of the Punta del Morro-Punta Delgada

region can be divided into 3 sub-zones: upper,

middle and lower intertidal. Each zone can be

characterized by certain mollusks.

The upper intertidal zone is characterized by

Siphonaria pectinata, Nerita fulgurans, and

Thais haemastoma. These species were also

found in the supratidal zone but were more

abundant in the upper intertidal zone.

Two keyhole limpets, Diodora cayenensis, and

Fissurella barbadensis. and the muricid Pur-

pura patula were the characteristic species of

the middle intertidal zone. Fissurella was the

most abundant limpet. One specimen of Acmaea

leucopleura was collected but considered uncom-

mon. In calm-water tide pools under rocks right

at the water line Planaxis lineatus appeared

gregarious often in clusters of 15-20 in-

dividuals, but sometimes as many as 80-100.

The lower intertidal zone was more of an in-

terface with the subtidal zone. Petaloconchu^

varians formed extensive mats in this area.

Brachidontes exustus and Isognomon hicolor

|:|

60 THE NAUTILUS

April 21, 1982

Vol. 96(2)

were commonly found attached to Petaio-

conchus. The polyplacophorans collected were

found in this zone, but only one specimen of each

was collected and were considered uncommon.

Of two muricids mentioned above, Purpura

patula was common on rough water, outer rocks

and Thais haemastoma was more common on

rocks found in moderate to slight wave energy

areas.

Subtidal zone

Barbatia domingensis, found to be most abun-

dant in turbulent environments on the Bermuda

platform by Bretsky (1967), was the most abun-

dant ark collected from the subtidal zone. Area

imbricata, Arcopsis adamsi, and Barbatia

Candida, which are perhaps the most con-

spicuous arks in the shallow waters of Lobos

Reef (Tunnell, 1974), were only occasionally col-

lected or observed in the subtidal zone. Locally

abundant in the upper portion of this zone were

clusters of Brachidontes exustus and Isognomon

bicolor. Aplysia of. brasiliana, A. dactylomelo

and two unidentified nudibranchs were collected

in 1976 while snorkeling.

Algae

Numerous species of mollusks are found asso-

ciated with marine algae. Warmke and Almodo-

var (1963) presented data on mollusk-algae

associations of Puerto Rico. They collected 90

species of molluscs from 25 species of algae.

Four species of algae, Sargassum cf. filipen-

dula C. Agardh, Padina vickersiae Hoyt, Caul-

perpa racemosa (Forskal) J. Agardh, and Ulva

lactiica Linnaeiis were collected at Punta Bo-

quilla de Piedras and examined for mollusks.

These four species appeared to be the dominant

algae.

Nine species of gastropods were found asso-

ciated with the four algal species collected. Only

two species, Tricolia affinis and Caecum

pulchellum, were found exclusively on the algae.

Littorina meleagris was also found intertidally

on the rocks. Caecum vestitum and Fossarus or-

bignyi were found in the sabellariid polychaete

colonies, as well as on algae. Planaxis lineatus

was collected both from calm-water tidepools

and algae, while Nitidella laevigata was found

crawling subtidally on the rocks among the

algae.

Sabellariid polychaete colonies

Sebellariid polychaete colonies were com-

monly observed covering portions of intertidal

boulders. These colonies formed a layer approx-

imately 2-10 cm thick and were composed of ag-

glutinated tubes of sand grains. Larger holes

and channels probably formed by wave action

were inhabited by numerous crustaceans, poly-

chaetes, and mollusks, Fossarus orbignyi,

Brachidontes exustus, and the nestling bivalve

Sphenia antillensis were the most common

mollusks collected from the sabellariid colonies.

Discussion

Although the majority of the molluscan fauna

collected from the Punta del Morro-Punta Del-

gada region is considered to be Caribbean in

origin, many of the families and genera were

represented by only one or two species. Tunnell

(1974) stated that southwestern Gulf reefs ap-

pear to have several reduced or lacking tropical

elements which are generally common in areas

more centrally located within the Caribbean

Province. This situation appears to be the case

in the Punta del Morro-Punta Delgada region

also. Although not universally accepted, the in-

shore northern boundary of the Caribbean prov-

ince is located in the vicinity of Cabo Rojo

(Briggs, 1974). This places the study area near

the periphery of the Caribbean province where

physical and ecological conditions are favorable

for tropical species, but are far from being op-

timal. The molluscan fauna of the Punta del

Morro-Punta Delgada rocky shores would there-

fore, most accurately be described as "depau-

perate Caribbean."

ACKNOWLEDGMENTS

The authors would like to gratefully thank the

Direccion General de Pesqueras, Departamento

de Pesca for allowing us to collect and study in

Mexico.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 61

LITERATURE CITED

Abbott, R. T. 1958. The Marine Mollusks of Grand Cayman

Island, British West Indies. Monograph 11. Acad. Nat.

Sci. Phila.. 138 pp., 5 pis.

Abbott, R. T. 1974. American Seashelh. Van Nostrand

Reinhold Co., New York, 663 pp.

Baker, F. C. 1891. Notes on a collection of shells from South-

ern Mexico. Proc. Acad. Nat. Sci. Phil. pp. 45-55.

Bretsky, S. S. 1967. Environmental factors influencing, the

distribution of Barbatia domingensis (Mollusca: Bivalvia)

on the Bermuda platform. PostiUa. Peabody Museum of

Nat. Hisf., Yale Univ. 108:14pp.

Briggs, J. C. 1974. Marine zoogeography. Mc-Graw-Hill,

New York, 475 pp.

Chavez, E. A., E. Hidalgo, and M. L. Sevilla. 1970. Datos

acerca de las comunidades bentonicas del arricife de

Lobos, Veracruz. Revista de la Sociedad Mexicana de

Historia Natural 31:211-280.

Coomans, H. E. 1958. A survey of the littoral Gastropoda of

the Netherlands Antilles and other Caribbean islands.

Study. Fauna Curaeao 8:42-111.

Ekdale, A. A. 1974. Marine molluscs from shallow-water en-

vironments (0 to 60 meters) off the northeast Yucatan

coast, Mexico. Bull. Mar. Sci. 24(3):638-668.

Galtsoff, P. S. ed. 1954. Gulf of Mexico, its origin, waters,

and marine life. U.S. Fish Bull. 55(89):604 pp.

Garcia, E. 1970. Los climas del estado de Veracruz. Ann.

Inst. Biol. Univ. Anton Mexico 41(Ser. Bot l):3-42.

Houbrick, J. R. 1968. A survey of the littoral marine mol-

lusks of the Caribbean coast of Costa Rica. The Veliger

ll(l):4-23.

Jaume, M. L. 1946. Moluscos marinos litorales del Cabo

Catoche, Yucatan, Mexico. Revista Soc. Malac. Carlos de

la Tor-re 4(3):95-100.

Keen, A. M. 1971. Sea shells of tropical west America. Stan-

ford Univ. Press, Stanford, Calif., 1064 pp.

McLean, R. A. 1951. The pelecypod or bivalve mollusks of

Porto Rico, and the Virgin Islands. Sclent. Surv. Porto

Rico. N.Y. Acad. Sci. 17(1):1-183.

Moore, D. R. 1958. Notes on Blanquilla Reef, the most north-

eriy coral formation in the western Gulf of Mexico. Puhl.

Inst. Mar. Set. Texas 5:151-155.

Olsson. A. A. and T. L. McGinty. 1958. Recent marine mol-

lusks from the Caribbean coast of Panama with the de-

scription of some new genera and species. Bull. Amer

Paleo. 39(177):58 pp.

Radwin, G. E. 1969. A recent molluscan fauna from the

Caribbean coast of southeastern Panama Trans. San Diego

Soc. Nat. Hist. 15(4):229-236.

Rehder, H. A. 1962. Contribucion al conocimiento de los

moluscos marinos del archipielago de Los Roques y La

Orchila. Mems. Soc. Sctenc. nat. Xa Salle' 22:116-138.

Rice, W. H. and L. S. Kornicker. 1962. Molluscs of Alacran

Reef, Campeche Bank, Mexico. Puhl. Inst. Mar. Sci..

Texas 8:366-403.

1965. (Addendum to 1962 paper). Puhl. Inst.

Mar. Set.. Texas 10:172.

Tunnell, J. W., Jr. 1974. Ecological and geographical distri-

bution of Molluscs of Lobos and Enmedio coral reefs,

Southwestern Gulf of Mexico. Ph.D. Dissertation. Texas

A&M Univ., 158 pp.

Usticke, G. W. N. 1959. A checklist of the marine shells of St.

Croix. U.S. Virgin Islands, unth random annotations.

Lane Press, Burlington, Vermont, 90 pp.

Villalobos, A. 1971. Estudios ecologicos en un arrecife cora-

lino on Veracruz, Mexico. Pp. 532-545 in Symposium on

investigations and resources of the Caribbean Sea and ad-

jacent regions, preparatory to CICAR, organized jointly

by UNESCO and FAO.

Warmke, G. L. and R. T. Abbott. 1961. Caribbean seajshells.

Livingston Publishing Co., Narbeth, Penn., 348 pp.

Warmke, G. L. and L. R. Almodovar. 1963. Some associa-

tions of marine molluscs and algae in Puerto Rico. Mala-

cologia 1(2):163-177.

Weber, J. A. 1961. Marine shells of Water Island, Virgin

Islands, The Nautilus 75(2):55-60.

Weisbord, N. E. 1926. Notes on marine mollusks from the

Yucatan Peninsula, Mexico. The Nautilus 39(3):81-87.

Whitten, H. L., H. F. Rosene, and J. W. Hedgepeth. 1950.

The invertebrate fauna of Texas coast jetties; a pre-

liminary survey. Publ. Inst. Mar. Sci.. Texas 1:53-87.

Work, R. C. 1969. Systematics, ecology, and distribution of

mollusks of Los Roques, Venezuela. Bull. Mar. Sci.

19(3):614-711.

1^

SIX NEW SPECIES OF INDO-PACIFIC TEREBRIDAE (GASTROPODA)

Twila Bratcher

8121 Mulholland Terrace

Hollywood, CA 90046

and

Walter 0. Cernohorsky

Auckland Institute and Museum

New Zealand

While doing research for a forthcoming book,

we have come across a number of undescribed

terebrid species, some in museums, others from

private collectors. Some other species were

62 THE NAUTILUS

April 21, 1982

Vol. 96(2)

FK ;S 1 i):Tcrebra mactanensis Bratcher & Cemohorsky. new species. Holotype, LACM no. 1968. oi.i mm. 2 & 9.

Terebra ■o^.-^.v..- Bratcher & Cemohorsky. n^ species. Holotype. LACM no. 1969. 26.1 mm. 3& 8: Dupl.cana mozambiquen-

sis Bratcher & Cemohorskv. new species. Holotype NM no. H78iS. 22.3 mm. 4 & 12: Terebra caddeyi Bratcher & Cemohorsky.

new mcie.s. Holotype LACM no. 1967. 52.7 mm. 5 & 11: Duplicaria baileyi Bratchei- & Cemohorsky. new species. Holotype

LACM no. 1070. ii.f m.m.. 6 & 7: Terebra burchi Bratcher & Cemohorsky. new .species. Holotype MNHN. 17.9 mm.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 63

represented only by a single specimen, and we

will wait for more material before describing

them. Six are being described here.

Terebra burchi new species

(Figs. 6. 7)

Diagnosis: A pure-white shell with small

brown dots scattered at random just below the

suture and with a broadband of yellowish brown

on the base of the body whorl.

Description: Shell about 18 mm in length, with

12 whorls of teleoconch; color pure-white with

small yellowish brown dots scattered at random

immediately anterior to the suture and with the

same color on the base of the body whorl; outline

of whorls convex; protoconch with IV2 whorls

remaining (paratype with intact protoconch of

3V2 amber conical whorls); sculpture of slightly

arcuate axial ribs from suture to suture, 13 on

penultimate whorl; ribs well-developed, quite

sharp, and narrower than interspaces; inter-

spaces with 5 evenly-spaced grooves, the

posterior being deep; interstical grooves not

crossing ribs; no subsutural band; body whorl

with ribs fading at periphery; 8 grooves anterior

to periphery, 6 posterior; aperture quadrate,

with basal plication; columella recurved.

Dimensions: Holotype 17.9 x 4.3 mm. Para-

types from 18.8 x 4.3 to 22.4 x 5.4 mm.

Type locality: Northern Mozambique Channel

(12 45'S; 45 18;E), at 15 -20 m, Benthedi Ex-

pedition sta. 32.

Type Material: Holotype MNHN Paris. Para-

types LACM no. 1971 (1); MNHN (2); Bratcher

coll. (2): Cernohorsky coll. (2): Tursch coll. (2).

Distribution: From Mozambique and Papua

New Guinea; 2 to 20 m. Also Philippines.

Discussion: There are several other Indo-

Pacific terebrids with dark areas anterior to the

periphery of the body whorl and scattered spots

below the suture. None of them has a pure-white

background. Terebra amoena Deshayes, 1859,

T. conspersa Hinds, 1844, and T. pertv^a (Born,

1778), all have larger shells with beige to tan

backgrounds and have obvious subsutural

bands.

This species is named in honor of a late friend

and colleague, R. D. Burch.

Terebra caddeyi new species

(Figs. 4, 12)

Diagnosis: A long, slender, flat-sided terebrid

shell, shiny tan, and with 3 or 4 spiral grooves

per whorl.

Description: Shell long, slender, with 25

whorls; color shiny tan; outline of whorls

straight; protoconch missing; 3 spiral bands,

each defined by a spiral groove, occur anterior

to suture; posterior band narrow, without

nodes; center band of obsolete nodes; anterior

band flatter and scarcely noded; no axial sculp-

ture except axial striae; body whorl with deep

groove posterior to periphery, followed by 3

weak grooves; aperture small, quadrate; col-

umella curved, with moderately heavy parietal

callus; siphonal notch broad.

Dimensions: Holotype 52.7 x 6.9 mm. Para-

types from 44.4 x 5.4 to 53.4 x 6.6 mm.

Type Locality: Korere Village, Rabaul, Papua,

New Guinea, 3 m. in depth, on volcanic sand.

Type Material: Holotype LACM no. 1967.

Paratypes Bratcher coll. (1); Caddey coll. (2).

Distribution: Known only from type locality.

Discussion: Terebra lima Deshayes, 1857, also

has a flat-sided shell, but it lacks the sheen of

this species and has almost cancellate sculpture

with spiral cords and axial sculpture forming

small nodes at intersections. T. jenningsi Burch,

1965, has a concave outline, a cord at the periph-

ery of the body whorl, and more spiral grooves.

It also lacks sheen. This species is named for P.

T. Caddey, who collected the type lot.

Terebra mactanensis new species

(Figs. 1. 10)

Diagnosis: A long, extremely slender terebrid

shell with dark spots between axial ribs anterior

to the suture.

Description: Shell extremely slender, long;

color, shiny brown with black spots between ax-

ial ribs anterior to suture and a row of small

white dots at periphery of body whorl; outline of

whorls convex; protoconch of 3 white, conical

whorls; sculpture of axial ribs from suture to

suture, narrower than interspaces; ribs on body

whorl terminating at periphery in a white spot;

aperture elongate; columella recurved, white;

siphonal fasciole extremely heavy, long.

64 THE NAUTILUS

April 21, 1982

Vol. 96(2)

Di7nensions: Holotype 54.4 x 7.2 mm. Para-

types from 47.8 x 6.1 to 62.1 x 8.1 mm.

Type Locality: Punta Egano, Mactan Island,

Cebu, Philippine Islands at 200 m.

Type Material: Holotype LACM no. 1968.

Paratypes Bratcher collection. (1); Marrow col-

lection. (1).

Distribution: This species is known only trom

the type locality.

Discussion: One of the paratypes is light

orange-brown with dark brown spots between

the ribs anterior to the suture. The other is

white with a few brown areas anterior to the

suture and with extremely faint brown spots at

the periphery of the body whorl. All have similar

sculpture. There is no Indo-Pacific terebrid

species with which T. mactanensis could be con-

fused. The white paratype might be compared

with T. fortunae Deshayes, 1857, which has an

inflated shell with longer whorls, has more

spiral sculpture; and has no brown ornamenta-

tion. The type lot was collected by Max Marrow.

Terebra marrowae new species

(Figs. 2, 9)

Diagnosis: A cream-colored terebrid shell

with a few small orange-brown dots and with an

exceptionally large, blackish mamillate proto-

conch.

Description: Shell medium-sized for the

genus; color, cream with a few small scattered

orange brown dots, protoconch and first whorls

of teleoconch brown; outline of whorls concave

in early whorls, flat in later ones, with convex

subsutural band; protoconch of IV2 broad, ma-

millate blackish brown whorls; subsutural band

of large pearl-like nodes, occasionally spotted

between by orange-brown; suture shallow; sub-

sutural groove deep, narrow; subsutural band

followed by a narrow, almost smooth, inconspic-

uous band, becoming slightly nodulous on last 2

whorls; remainder of whorl finely concellate

with 2 spiral cords per whorl, forming shallow

pits between intersections; body whorl with

finely cancellate sculpture ending posterior to

periphery, spiral cords only continuing to

siphonal fasciole; aperture short, quadrate; col-

umella recurved.

Dimensions: Holotype 26.1 x 5 mm. Para-

types from 21.4 x 4.8 to 32.1 (Apex missing) to

6.4 mm.

Type Locality: Cleaverville, N.W. Australia;

intertidal.

Type Material: Holotype LACM no. 1969.

Paratypes Bratcher coll. (1); Caddey coll. (2);

Cernohorsky coll. (1); Cooper coll. (2).

Distribution: Northwest Australia, intertidal.

Discussion: The most outstanding feature of

this species is the extremely large blackish

brown or purplish brown mamillate protoconch,

with a lighter brown or purplish area extending

through two to four whorls of the teleoconch,

becoming progressively lighter. The sculpture of

some of the paratypes is finer than that of the

holotype, with smaller nodes on the subsutural

band. The color varies from white to yellowish

white with a white subsutural band. All speci-

mens except one have a few tiny scattered

brown dots, and that one was collected dead.

Two of the paratypes have brownish pin-point

dots at the periphery of the body whorl.

There are several other Indo-Pacific terebrid

species with cancellate sculpture, all of which

are easily separable from Terebra marrowae. T.

swobodai Bratcher, 1981, is more slender, has

convex whorls and very heavy, rough sculpture

with small nodes forming where axial and spiral

cords cross. T. elliscrossi Bratcher, 1979, and T.

amanda Hinds, 1844, both have slender, non-

mamillate protoconchs of more than IV2 whorls.

Although T. fenestrata Hinds, 1844, has a

mamillate protoconch, it is about Vs as large as

that of T. marrowae, and the sculpture of the

teleoconch is extremely coarse and heavy. This

species is named in honor of Lorna Marrow, who

collected the holotype.

Duplicaria baileyi new species

(PI. 5, 11)

1965 Duplicaria australis (E. A. Smith), J. Gate & R. D.

Burch, Veliger 6(3): 145; 1967 Cernohorsky, Mar. SMls

Pacific 1;208, pi. 52, fig. 392. [non Terebra australis E. A.

Smith, 1873).

1966 Duplicaria (Duplicaria) sp. Cernohorsky & Jennmgs,

Veliger 9(1):58, pi. 6, fig. 44.

1978 Terebra australis (E. A. Smith), Hinton, Guide Austra-

lian Shells, pi. 59, fig. 6. [non Terebra australis E. A.

Smith, 1873].

Vol. 96(2)

April 21, 1982

THE NAUTILUS 65

Diagnosis: A beige Duplicaria shell with a few

■yellowish brown splotches scattered irregularly

and with brown anterior to the periphery of the

body whorl.

Description: Shell medium sized, with 13

whorls of teleoconch; color, beige with a few

reddish brown splotches and dots scattered at

random, and with the same color anterior to the

aeriphery of the body whorl; outline of whorls

5lightlyconvex; protoconch of 4V2 pink, translu-

cent, conical whorls; axially ribbed subsutural

mnd defined by shallow groove, with deeper

junctations between ribs; remainder of whorl

vith straight axial ribs contiguous with ribs on

)and; interspaces about equal to ribs, smooth;

)ody whorl with ribs ending at periphery; peri-

)hery marked with light band between rib end-

ngs and light-brown area; aperture elongate;

:olumella recurved; siphonal fasiole striate, with

noderate keel.

Dimensions: Holotype 24.9 x 5.3 mm. Para-

ypes 21.5 x 4.5 to 40.0 x 7.9 mm.

Type Locality: Guadalcanal, Solomon Islands

9 25'S; 159 56'E); in sand at 20 m.

Type Material: Holotype LACM no. 1970.

^aratypes ANSP no. 352482 (1); AM no.

]132464 (1); CAS no. 60674 (1); MCZ no. 290426

1); MORG no. 21.275; NM no. H-766 (1); USNM

10. 782262 (1); Bratcher coll. (2); Cernohorsky

oil. (2); Morrow coll. (2).

Distribution: Mozambique to the Philippines

.nd the Solomon Islands; from intertidal to

;00m.

Discussion: The number of axial ribs varies

rom 20 to 29 on the penultimate whorl, and the

■asic color varies slightly from cream to beige,

'he brown splotches may vary in number and

.ize, but they are always inconspicuous. Dupli-

aria teramachii Burch, 1965, is somewhat

imilar in color pattern although it has a broad

and of reddish brown instead of the occasional

iterrupted blotches. It also has a narrower

pical angle, flatter subsutural band, and

horter whorls. This species is named in honor

f Brian Bailey who collected the holotype.

Duplicaria mozambiquensis neu^ species

(Figs. 3, 8)

Niiii Terehm gracilis Reeve, Conch. Icon. 12, pi. 24, fig. 1.31

[non Lea 1833; non Gray, 1834] Hab.: Africa; holotype

BM(NH). 1874.10.29.2; 22.0 mm.

Diagnosis: A Duplicaria with extremely in-

flated early whorls, a punctate subsutural

gi-oove, and a narrow, noded subsutural band.

Description: Shell color light-brown with ivory

ribs, nodes, and narrow peripheral stripe on

body whorl; outline of whorls convex; proto-

conch mamillate with IVz brown, moderately in-

flated embryonic whorls; sculpture of early

whorls of teleoconch consisting of a narrow sub-

sutural band with wide-spaced round nodes (8 or

9), and below each node an arcuate rib almost

noded in center, giving an angulate outline to

the early whorls; a punctate subsutural groove

developing after the 3rd whorl of teleoconch;

ribs on remainder of whorl remaining wide-

spaced, sharp, and arcuate, but losing the

angulate form on later whorls (12 on penulti-

mate whorl); nodes on subsutural band angulate

on later whorls; body whorl with sharp ribs con-

tinuing below periphery to keel of siphonal

fasciole; aperture elongate; columella almost

straight.

Dimensions: Holotype 22.3 x 5.6 mm. Para-

types from 17.0 x 4.4 to 33.0 x 6.8 mm.

Type Locality: Mozambique, trawled off

Chinde Island.

Type Material: Holotype Natal Museum no.

H7843/T2541. Paratypes in BM(NH) 198021;

Natal Museum nos. J2973/T2542 (1), 566/T2543

(1), B2137/T2544 (1), H765/T2545 (2); Bratcher

coll. (1).

Distribution: From Durban, South Africa, to

Mozambique.

Discussion: Four of the paratypes are the

same color as the holotype; one is peach color;

and two are ivory. Some individuals have small,

inconspicuous nodes at the posterior end of the

axial ribs. Some have more numerous, less wide-

ly spaced ribs.

Duplicaria spectabilis (Hinds, 1844) has

similar sculpture, but it has a distinctive color

pattern of rich brown and cream stripes, and a

protoconch of 2V2 whorls. Duplicaria evoluta

Deshayes 1859, has a broad subsutural space but

no groove. It has axial ribs on the subsutural

band instead of round nodes, and the early

66 THE NAUTILUS

April 21, 1982

Vol. 96(2)

whorls are quite flat in outline. This is the same

species as Terebra gracilis Reeve, 1844, a name

which was preoccupied.

Abbreviations have been used for a number of

institutional collections cited in this paper. They

are:

AM - Australian Museum

ANSP- Academy of Natural Science of Phila-

delphia.

CAS - California Academy of Sciences.

LACM-Los Angeles County Museum of Natur-

al History.

MCZ- Museum of Comparative Zoologj-, Har-

vard University.

MNHN - Museum National d'Histoire Naturelle,

Paris.

MORG-Museu Oceanografico de Rio Grande,

Brazil.

NM- Natal Museum, South Africa.

ACKNOWLEDGMENTS

We wish to thank the following for the loan of

tjpe material for this study: Brian Bailey of

Honiara, Solomon Islands; Dr. Philippe Bouchet

of MNHN Paris; P. T. Caddey of N.S.W., Aus-

tralia; Dr. George Davis of ANSP; Dr. R. Kil-

burn of NM; Max Marrow of Victoria, Australia;

Dr. Ben Tursch of Brussels, Belgium; Dr.

Joseph Rosewater of USNM.

LITERATURE CITED

Born, Ignatius. 1778. Index Rerum NaturcUium Caesaeri

Vindobonensis. Testacea.

Bratcher, Twila L. 1979. Taxonomic Changes in Eastern Pa-

cific Terebridae, with Description of a New Species. The

Veliger 22(l):61-64; figs. 1-12.

1981. Four Previously Undescribed Indo-Pacific

Terebrids. The Veliger 23(4):329-332; figs. 1-8.

Burch, Robert D. 1965. New Species from the Indo-Pacific

Ocean and from the Gulf of Mexico. The Veliger

7(4):241-253: pi. 31, figs. 1-10.

Deshayes, G. P. 1857. Description d'especes nouvelies du

genre Terebra, Jour, de Conchy I. 6:65-102, pis. 3-5.

1859A. General review of the genus Terebra,

and a description of new species. Proc. Zool. Soc. London.

p. 270-321.

Hinds, R. B. 1844. Synopsis of the known species of Terebra.

Proc. Zool. Soc. London for 1843, pt. 11:149-168.

THE FRESHWATER MUSSELS (NAIADS) OF BIG INDIAN

CREEK, A SMALL SOUTHERN INDIANA TRIBUTARY OF THE

OHIO RIVER (BIVALVIA: UNIONIDAE)

Ralph W. Taylor

Department of Biological Sciences

Marshall University

Huntington, West Virginia 25701

ABSTRACT

A 1979-81 survey of Big Indian Creek in southern Indiana prodiLced a total of

16 species of freshwater mussels (Naiads). In addition to these unionid species, the

exotic clam Corbicula fluminea was also found to be present throughout the

stream. The faunal make-up proved to be typically Ohioan in nature. The most

common species were Lampsilis r. luteola, Lampsilis ventricosa and Amblema p.

plicata. Lampsilis fasciola, Leptodea fragilis and Toxolasma parvus were each

represented by a single specimen.

In their Revision of the Mollusca of Indiana, together, in a condensed form, all the early

Goodrich and van der Schalie (1944) brought work done on the freshwater mussels in Indiana

Vo). 96(2)

April 21, 1982

THE NAUTILUS 67

streams. A more recent work by Krumholz et al

1970) provided valuable information on the

vVhite and Wabash Rivers. Most of the pub-

ished information available on the mussels of

Indiana is in fact concerned with the naiad

aunas of the larger rivers. There appears to be

I dearth of information available on the mussels

)f small streams, and in particular those

streams of southeastern Indiana. It is the pur-

)ose of this paper to report work on such a

.tream, Big Indian Creek, a tributary of the

)hio River located in southern Indiana.

Big Indian Creek originates in the low-rolling

ill section of southeastern Indiana, locally

nown as Floyd Knobs, and flows in a generally

outhwesterly direction to its confluence with

he Ohio River at the village of New Amsterdam

ome fifty river miles downstream of Louisville,

Kentucky. The stream presently exists in a fair-

-' undisturbed condition. Big Indian Creek flows

irough land that has for over a hundred years

een dedicated to agricultural use. A good per-

entage of the lower portion of the creek flows

irough Harrison/Crawford State Forest. The

nly towns of any size in this drainage are the

illages of Crandall and Corydon, with a com-

ined total population of less than one thousand

ihabitants. Water quality is apparently good,

s the only possible source of pollution would be

lat which results from limited agricultural ac-

vities within the floodplain. The substrate is

rimarily limestone bedrock with an occasional

and and gravel bar. There is very little

vidence of excessive silt deposition anywhere

long the stream's course.

Methods

During the summers of 1979 and 1981, Big In-

ian Creek was surveyed for freshwater mussels

t irregular time intervals. Seven collecting sta-

ons were established. Fresh dead shells were

andpicked from the water and shore line. Very

Id, severely weathered specimens were dis-

irded and only specimens of those species con-

dered to be part of the present naiad fauna are

icluded in this report. Specimens were cleaned

nd cataloged in the Marshall University Mala-

)logical Collections. Voucher specimens have

been placed in the Ohio State University Muse-

um of Zoology. Scientific names are those used

by Stansbery (1980).

Collecting Stations

Under bridge on Indiana St. Rt. 335, .4 mi S of St. Rt. 64,

1 mi N of village of Crandall; Harrison County, Indiana.

Take St. Rt. 335 SW of Crandall to Bethlehem" Cemetan,-

Rd., .5 mi N on Cemetary Rd. to Indian Creek; Harrison

County, Indiana.

Take St. Rt. 135 S of village of New Salisbury to un-

named, unimproved dirt road, .4 mi S of Southern Rail-

way tracks, .7 mi E on dirt road to bridge; Harrison

County, Indiana.

Take St. Rt. 135, 3.7 mi S of New Salisbury to Conrad

Cemetary Rd., .3 mi E to Creek; Harrison County, In-

diana.

Take St. Rt. 135 2.5 mi SW of Corydon to Heidleburg

Church Rd, then Mathis Rd. 1 mi to Creek; Harrison

County, Indiana.

Take St. Rt. 135 to Heidleburg Church Rd., then to Val-

ley View Rd. 1 mi to creek, Ca 3.5 mi SW of Corydon;

Harrison County, Indiana.

Indiana St. Rt. 335, .1 mi E of Indiana St. Rt. 135, Ca 2

mi S of New Salisbury; Harrison County, Indiana.

Table 1. Freshwater Naiads collected 1979-81 from Big Indian Creek.

Species listed by site.

Anodonta

grandis Say, 1829

Strophitus u. undiilatus (Say, 1817)

Alasmidonta vindis (Raf., 1820)

Lasmigona complanata (Barnes, 1823)

Lasmigona costata (Raf.. 1820)

Aroblcma £. plicata (Say, 1817)

Fusconaia flava (Raf.. 1820)

EUiptlo dilatata (Raf., 1820)

Leptudea fragiUs (Raf., 1820)

Potamilus alatus (Say. 1817)

Toxolasma parvus (Barnes, 1823)

VI 1 Ipsa I, ins (Lea, 1829)

Vlllosa lienosa (Conrad, 1834)

l.ampai 1 is radiata luteola (faoi., 1819)

LampBi 1 IS vcntncosa (Barnes, 1823)

Lampsilis fasciola Haf . . 1820

corbicula fluminea

68 THE NAUTILUS

April 21, 1982

Vol. 96(2)

Summary

Survey collecting at seven stations along Big

Indian Creek yielded a total of sixteen species of

freshwater Naiads plus the exotic Asiatic import

Corbiculajluminea (Table 1). Most species were

present in good numbers at several different lo-

calities. Five species (Anodonta grandis, Lepto-

dea fragilis, Elliptio dilatata, Toxolasma par-

vus and Lampsilis fasciola) were, however,

found at only one locality each.

None of the species found is currently con-

sidered to be rare or endangered. The faunal

make-up of this small stream is typically Ohioan

and is fairly similar to that found in Floyd's Fork

and Salt River (Taylor, 1980) which enter the

Ohio River from the south in the proximity of

the mouth of Big Indian Creek which enters

from the north.

LITERATURE CITED

Goodrich, Calvin and H. van der Schalie. 1944. A Revision of

the Mollusca of Indiana. The Amer. Mid. Nat. 32(2):

257-326.

Krumholz, Louis A., R. L. Bingham and E. R. Meyer. 1970.

A Survey of the Commercially Valuable Mussels of the

Wabash and White Rivers of Indiana. Proc. of the Indiana

Acad, of Set. 79:20.5-226.

Stansbery, David H. 1980. Naiad Mollusks of the Ohio River

Drainage System. The Ohio State University Museum of

Zoology, 1 p. mimeo.

Taylor, Ralph W. 1980. Mussels of Floyd's Fork, A Small

Northcentral Kentucky Stream. The Nautilus 94(1):

13-15.

MEETINGS

July 14-17. CO. A. (Conchologists of America).

Sanibel Island, Florida.

July 19-23. A.M.U. (American Malacological

Union. New Orleans, Louisiana.

SHELL SHOWS

July 30 - Sept. 1. Jacksonville Shell Show.

Jacksonville Beach, Florida.

June 20-23. 15th annual meeting of the West-

ern Society of Malacologists at the University

of Redland, CA.

RECENT DEATHS

Wataru W. Sutow, M.D., well-known cancer

expert, author of medical textbooks, and active

member of the Houston (Texas) Shell Club, died

Dec. 20, 1981, at the age of 69. He was born

Aug. 3, 1912, in Guadalupe, California. See

American Malacologists (1973), p. 457 and Texas

Conchologist (1982), vol. 18, no. 2, p. 25.

^<^^MMA^AM^^^W^^^^^«MMMVMM^W^V^VVN^^^

Carlos G(uillermo) Aguayo, leading Cuban

malacologist, exiled in Puerto Rico, died Feb.

12, 1982, at the age of 83, in Puerto Rico. He

was born in Havana, Cuba on Dec. 19, 1899, and

became a professor at the University of Puerto

Rico in Mayaguez in 1958. He published over

130 papers on mollusks, especially those of

Cuba, and was a Guggenheim Foundation Fel-

low at Harvard in 1931-33. (see American

Malacologists (1973), p. 188).

LICENSED APPRAISALS

AND

IDENTIFICATION SERVICE

Collections of shells and libraries of shell

books expertly appraised for estate, gift and

tax purposes. Moderate fees, plus travel ex-

penses. Services confidential. Inquire below.

Professional identifications of marine mol-

lusks for biological surveys, environmental

studies and private shell collectors. Fees bas-

ed on hourly basis. Minimum $20.00.

R. Tucker Abbott, Ph.D.

P. 0. Box 2255

Melbourne, Florida 32901

U.S.A.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 69

MASSIVE DESTRUCTION OF SCLERACTINIAN CORALS BY THE

MURICID GASTROPOD, DRUPELLA, IN JAPAN

AND THE PHILIPPINES'

Jack T. Moyer

Tatsuo Tanaka Memorial Biological Station

Ako, Miyake-jima, Izu Islands, Japan 100-12

William K. Emerson

American Museum of Natural History

New York, New York 10024, U.S. A

and

Michael Ross

Bureau of Fisheries and Aquatic Resources

Region 7 District Office

Cebu City, Philippines

ABSTRACT

Aggregates of the muricid gastropods Drupella fragum, at Miyake-jima, Japan,

and Drupella rugosa, at Mactan Island, Cebu. Philippines, were noted to result in

widespread destruction of scleractinian corals. Destruction was limited to fast-

growing foliate forms such as Montipora, Acropora, Pocillopora, etc., but other

genera, e.g. Porites, Goniopora, Favia, Fungia, etc.. were avoided. Muricid pre-

dation at Mactan appeared to represent a biological control on coral reef struc-

ture, "weeding out" fast growing forms and providing space and settlement sites

for other coral species, thus increasing diversity over time. At Miyake-jima, in the

warm, temperate waters of southern Japan, Acropora spp. are dominant. Coral

destructio7i by Drupella fragum has been massive on Acropora reefs at Miyake-

jima, amounting to as much as 35% of our study reef (Toga Reef No. 1) in two

years. The sudden population explosion o/D. fragum in 1976 at one site on the

island and in 1980 at another location correlates with massive siltation from

heavy bulldozing on the volcanic island, followed by heavy rainfall. A hypothesis

to explain this correlation is presented.

Although many marine animals are known to

feed on hermatypic corals (Robertson, 1970;

Reese, 1977; Taylor, et ai, 1980), only the

asteroid Acanthaster planci has previously been

reported as representing a serious threat to

coral reefs (Randall, 1972; Branham, 1973; En-

dean, 1973; and others). Robertson (1970) re-

ported that certain species of prosobranch gas-

tropods of the muricid genus Drupella are

known as predators on scleractinians of various

genera and that some have been found living in

fairly large colonies among the branches of and

under stony corals. However, to our knowledge.

'Contribution Nc

Station.

36. Tatsuo Tanaka Memorial Biological

population explosions of Drupella with

widespread coral destruction have yet to be

reported.

Since 1976, population explosions of varying

sizes involving Drupella fragum (Blainville,

1832) have been observed annually at the island

of Miyake-jima (34°05'N, 139°30'E), one of the

Izu Islands of southern Japan. The combination

of heavy predation of D. fragum and Acan-

thaster planci (see Moyer, 1978) have destroyed

much of the island's coral fauna. Since 1979, a

similar phenomenon, but with less destructive

results, has been under observation at Mactan

Island, Cebu, Philippines (10°18'N, 123°54'E),

involving Drupella rugosa (Born, 1778) and

various species of branching corals. In this

70 THE NAUTILUS

April 21, 1982

Vol. 96(2)

paper we document the role of Drupellafragum

and D. rugosa in the destruction of corals and

identify them as potentially harmful to coral

reefs.

Methods and Materials

Early observations of Drupella predation at

Miyake-jima were made incidental to behavioral

studies on reef fishes from 1976 to 1979. The ex-

tent of the damage caused by the gastropods

was not realized until 1980. Accurate measure-

ments and observations of Drupella destruction

were obtained for two months in 1980 and one

month in 1981. Data were collected both diur-

nally and nocturnally using scuba. Specimens

were collected as they fed on coral polyps 2 to 4

hours after sunset and as they were concealed

under coral branches by day. At Mactan Island,

three transects, each measuring 100 m in

length, were placed parallel to shore, 4 m apart

and at depths ranging from 1-2 m, at two study

sites (Tambuli Beach and Malingin Beach) for

the purpose of coral community studies. At

every .25 m point along these transects, a point

sample was made using a small weight (5 mm in

diameter). This provided 1200 points of informa-

tion for each reef site. Data were collected from

1979-1981. The percent of dead and grazed cor-

als was recorded at each site according to speci-

fic genera of corals and observations of Drupel-

la rugosa swarms were recorded, both diurnally

and nocturnally. Specimens were collected as

they aggregated under coral branches by day.

All specimens from both Japan and the Philip-

pines are deposited in the mollusk collection of

the American Museum of Natural History

(AMNH). Underwater photographs were taken

by Moyer at Miyake-jima and by Ross at Mactan

Island.

The four species of neogastropods observed

feeding on the corals at Miyake-jima and Mactan

are referable to the superfamily Muricacea.

Three are species of Drupella, a genus assigned

to the Thaidinae, a subfamily of the Muricidae

(Emerson and Cernohorsky, 1973; Ponder,

1973). The fourth species belongs to the nomi-

nate genus Coralliophila. of the Coral lio-

philidae, which are anatomically similar to the

Muricidae, but lack a radula. The radular denti-

tion of Drupella is atypical of the Muricidae, in

which three teeth are aligned in each transverse

row, with the central tooth commonly armed

with three cusps and the triangular lateral teeth

distally cuspid. In Drupella, the much larger and

more slenderly elongated lateral teeth are com-

monly basally denticulate on the inner margin

and distally terminate in a hooked tip (Cerno-

horsky, 1969, pp. 303-307, figs. 8-11). These

radular modifications probably reflect an adap-

tation to a specialized mode of feeding (Cooke,

1895; Arakawa, 1958; Wu, 1965; Robertson,

1970; and others). The reed-like lateral teeth

apparently sweep the polyps of coral onto the

basal denticles of the lateral teeth and the cusps

of the central teeth where the food particles are

abraded (cf. Cernohorsky, 1969, p. 304;

Fankboner, 1970, p. 15; Taylor, 1976, p. 185).

Results

Species identification

Japan: The two Miyake-jima forms of Dru-

pella clearly represent distinct species. The

specimens that were found in massive clusters

on coral are identified as Drupella fragum

(Blainville, 1832). These specimens (Figs. lA,

IB), which attain a maximum height of 23-)-

mm, have weakly developed axial ribs (12 per

whorl) and an essentially white aperture. The

exterior is pale orange-ochre, giving a yellowish-

white appearance to the shell. Many specimens

have a spirally arranged, brownish maculation

between the axial ribs, especially on the body

whorl. In some specimens, as many as six faint,

orange spots occur on the outer lip and in the in-

terior of the siphonal canal. The identification of

these specimens must remain tentative until

comparison can be made with the typological

specimens of Purpura fragum in Blainville's col-

lection, which is deposited in the Museum Na-

tional d'Histoire Naturelle, Paris. In the original

description, Blainville (1832, p. 203, pi. 9, fig. 4)

noted that the yellowish-white shell possessed

dark rose nodules and a white aperture. In the

present specimens, the nodules in some individ-

uals are dark to pinkish brown.

The second Miyake-jima species, Drupella

e^ata (Blainville, 1832, p. 207, pi. 11, fig. l)[ = D.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 71

FIGS. 1-3. lA iind IB, Drupella fraguni {Blainville). Miyake-jima. Jajjan; A, all white form. AMNH 203816a: B, maculated

form. AMNH ■203816h: x2. 2A and 2B, Drupella elata (Blainville), Miyake-jima. Japan, AMNH 202766; x2. 3A and 3B,

Drupella rugosa (Blainville). Marian Island. Philippines; A, all white form, AMNH 20266I,a; B,/orm mith colored nodules and

tannish orange aperture, AMNH 2027J,6b; x2.

'.pectrum (Reeve, 1846)], numerous individual

specimens of which were observed on the corals,

las a more massive shell with fewer (9 per

vhorl) axial ribs that are more obtuse. These

specimens, the largest of which examined by the

iecond author measures 28 + mm in height, are

vhite within and without, but the edge of the

)uter lip is tinged with greenish-yellow. The

;pecimens are largely overgrown with calcare-

)us algae and bryozoans (Figs. 2A, 2B). It

■hould be noted that this species was previously

)laced in the synonmy of Drupella cornus

Roding, 1798), see Cernohorsky (1969, pp. 304,

;05, pi. 48, figs. 12, 12a, b, and figs. 14, 14a [fide

Cernohorsky, 1978, p. 70]). H. A. Rehder (in

delville, 1980), however, notes that "D. comus

is restricted to the Indian Ocean . . . [and] D.

elata ... is found in the Pacific from Indonesia

eastwards." On morphological grounds, largely

the color of the aperture (orange in D. comus,

white in D. elata), Rehder does not believe the

two forms are conspecific, although he does not

rule out the possibility that D. elata may prove

to be a geographic subspecies of D. comns. We

follow Dr. Rehder in referring the western

Pacific specimens of this complex to D. elata

(Blainville).

The third muricacean gastropod observed

preying on corals at Miyake-jima is the wide-

ranging Indo-Pacific species, Coralliophila

violacea (Kiener, 1836) [=Murex neritoideus

Gmelin, 1791, not Linne, 1767; =C. diversi-

72 THE NAUTILUS

April 21, 1982

Vol. 96(2)

formis (Kiener, 1836); = C. squamulosa (Reeve,

1846)]. This distinctive suctorial-feeding species

is characterized by the dark violet aperture and

whitish squamulosely striate exterior (Cerno-

horsky, 1972, p. 131, pi. 37, fig. 5).

Philippines: The muricid snails found swarm-

ing on corals at Mactan Island are referable to

Dnipella rugosa (Born, 1778, p. 303) [=D. con-

catmata (Lamarck, 1822)]. These specimens

(Figs. 3A, 3B), the largest of which measures

24.6 mm in height (mean height approximately

18 mm in a sample of 360 specimens), has a con-

spicuously nodular appearance. The body whorl

is sculptured with 3 to 4 rows of prominent

nodules, which are reduced to a single row on

the penultimate whorl. The color varies consid-

erably, ranging from creamish-white specimens

to ones with dark brown or tannish nodules. The

aperture and siphonal canal may be entirely

white or a bright tannish-orange. Two speci-

mens of Drupella elata (Blainville, 1832) were

among the 348 specimens of D. rugosa collected

in one of the sites sampled on Mactan Island.

The larger specimen (AMNH catalogue no.

202766) measures 29.6 mm in height.

Field observations

Although situated at more than 34° North

latitude, Miyake-jima supports a surprisingly

diverse tropical fauna (Ida and Moyer, 1974;

Shepard and Moyer, 1980; Tribble and Randall,

MS). Tribble and Randall (MS) identified 92

species of scleractinian corals representing 45

genera. These are widely scattered in small out-

croppings, rather than as true coral reefs such

as those typically found at tropical latitudes.

Some relatively large Acropora patches are

present in scattered locations around the island,

the largest of which is located at Toga Bay on

the southwest side of the island. This patch

covers an area of approximately 1200 m\ and

will hereafter be referred to as Toga Reef No. 1.

"Swarming" of Drupella fragum was first

noted on a large platform of Acropora sp. (1.5 m

in diameter) in Igaya Bay, on the northwest side

of Miyake-jima, in August, 1976. Recent evi-

dence of gastropod predation was evident in the

form of fresh, white coral skeletons and small

swarms of D. fragum, but when the entire plat-

form was found dead and overgrown with algae

in the Spring of 1977, it was not known whether

it was killed by D. fragum, Acanthaster planci

(which appeared on the island's corals in the

winter of 1976-77; Moyer, 1978), or from silta-

tion from construction near the bay followed by

heavy rains.

A similar Acropora platform at Igaya Bay was

used as a cleaning station by a juvenile labrid

fish, Thalassoma lutescens, in 1978, and was

photographed and repeatedly observed at that

time (Fig. 4A). During September and October,

1978, a swarm of D. fragum appeared on this

coral, which was subsequently totally destroyed,

apparently by Drupella predation (Fig. 4B).

Accurate observations and measurements of

D. fragmn destruction were not possible until

the autumn (September and October) of 1980,

when one of us (JTM) was engaged in daily

Acanthaster monitoring as part of a removal

program. At this time, sizeable areas of freshly

killed white Acropora spp. were discovered,

with the area of destruction widening on a daily

basis. Careful searches in and under the corals

disclosed only swarms of Drupella fragum. Dur-

ing an evening dive, 17 September 1980, an in-

festation of several thousand Drupella fragum

was found massed around the border between

living and dead corals, devouring the polyps.

The snails were bunched around and on top of

each other in such numbers that it was impossi-

ble to estimate how many individuals were pre-

sent. Between 20:25 h and 21:00 h, a total of

about 1,500 were collected, 662 of which were

kept and later deposited at AMNH (catalogue

no. 203816). The remaining specimens were dis-

carded. Hundreds of others fell through the cor-

al branches and could not be recovered. In-

terestingly, all 12 AMNH specimens selected at

random for radular study were found to be

females. This swarm of D. fragum seemed to

disperse as waters cooled in winter, however, an

area of 35 m^ of reef was destroyed by their

predation during the two month period of obser-

vations (Fig. 5). No egg cases were observed on

or under the corals, suggesting that the

gastropods were swarming to feed. There was

no noticeable evidence that the swarms were

Vol. 96(2)

April 21, 1982

THE NAUTILUS 73

"IG. 4. A. Healthy Acropora coral platfimn at Miyake-jima. Japan. !!)?:). with the juvenile fish. Thaiassoma lutescens

I.'ifindae). cleaning Chromis flavoniaculata (Pomacentridae): B. Same Acropora platform in 1981. totally dead and overgrown

filli algae, after invasion o/" Drupella fragum. Note faviid coral lo7i far side in both photographs) remains unharmed by

II 1st ropcid invasio n.

74 THE NAUTILUS

April 21, 1982

Vol. 96(2)

FIG. 5. A. /";/"(

Japan, Sept. I'JSO.

, , \. ;. i/-,.ir. jniin. Japan, 1979, prior to Drupella fragum irM^aston; B, Toga Reef No. 1. Miyake-jimi

• il imik i/f Drupeila explosion: 5C, (on opposite page) Dead reef at same location in October. 1981.

Vol. 96(2)

April 21, 1982

THE NAUTILUS 75

FIG. 6. Toga Reef No. 1. Miynkejniia. Jiijiini, during Drupella fnigiini rxphision, Norcinhfr, i:lsi. A'nlc evidence of

movement from left to right in photo, with fresh algal growth over earlier coral kill at the left of photo.

76 THE NAUTILUS

April 21, 1982

Vol. 96(2)

directly related to reproductive purposes, but an

indirect relationship (intensive feeding either

directly before or directly after the reproductive

season) cannot be discounted.

A similar explosion of D. fragum was moni-

tered at Toga Reef No. 1 throughout the month

of November, 1981 (Fig. 6). During this period,

a total of 17 m^ of coral was destroyed. The com-

bined damage observed during two months in

1980 and one month in 1981 amounted to more

than 4% of the 1200 m^ reef being destroyed by

what was unquestionably predation by D.

fragum. Between 1979 and 1981, approximately

35% of Toga Reef No. 1 was destroyed. Records

from a carefully monitored ^ca?ii/iaster removal

program carried out in Toga Bay in 1980 and

1981 strongly suggest that Drupella fragum.

predation was responsible for most of the coral

destruction at Toga Reef No. 1. Acanthaster

were rarely seen at that reef, but were present

in small numbers at Toga Reef No. 2 and, more

commonly, at isolated patches deeper in the bay.

Surveys in November, 1981, at Igaya Bay, site

of the first detection of Drupella fragum out-

breaks in 1976, disclosed numerous patches of

freshly killed Acropora spp., similar to those at-

tributed to probable Acanthaster predation in

1978-1980. Close observation in and under the

branches of such corals invariably disclosed

clusters of D. fragum and numerous individuals

of the much larger D. elata (Blainville, 1832).

Relatively low numbers of Acanthaster were

removed from Igaya Bay, in spite of intensive

searches during the Acanthaster removal pro-

grams in 1979-1980. This fact, plus incidental

observations oi D. fragum outbreaks from 1976

to 1980 and the widespread occurrence of iso-

lated patches of corals under Drupella attack in

1981 (when the first intensive searches for the

muricids were attempted), suggest in retrospect

that a large amount of the massive Acropora

destruction at that site (perhaps as much as 50%

or more) is attributable to population explosions

of Drupella fragum.

Failure to discover the important role of muri-

cid predation at the onset of coral destruction at

Miyake-jima resulted from (1) presence of mod-

erate numbers oi Acanthaster planci at various

locations around the island and the perhaps

understandable assumption that fresh coral kill

resulted from asteroid predation, and (2) the

secretive behavior of D. fragum, which ag-

gregate far back under coral branches by day,

coming out at night to feed on coral polyps (Fig.

7). Observations in 1981 suggest t\\aL.tD. fragum

are more active during the dark phases of the

moon than around the full moon (more were visi-

ble on both night and day dives around the new

moon than at the full moon, and more fresh cor-

al kill was noted, although accurate measure-

ments on a daily basis could not be obtained, due

to the position of the study site on the windward

side of the island). D. fragum, also appeared to be

negatively phototactic. Many dropped between

the coral branches when caught in the flashlight

beam or as the strobe flashed.

At Mactan Island, data on swarming by Dru-

pella rugosa were collected from three different

sites, two at Tambuli Beach (AMNH catalogue

nos. 202763, 202764) and one at Malingin Beach

(AMNH catalogue no. 202765), see Table 1. On

Philippine reefs studied by the third author,

these snails are usually found widely dispersed,

in small groups or as individuals, and they ap-

parently are restricted to shallow waters. Since

April, 1980, it has been noted that at the Mactan

study sites, D. rmgosa has occurred on numerous

occasions in higher densities, including swarms

of up to 1500 individuals per .5 m\ comparable

to swarms of D. fragum on Acropora corals at

Miyake-jima (Fig. 8). Significant destruction of

specific corals was noted in localized areas; how-

ever, massive reef destruction, such as reported

above on the reefs at Miyake-jima, has not been

observed. Grazing by swarms oi D. rugosa, like

D. fragum, is nocturnal, with observed preda-

tion to date limited to the Astrocoeniina, specifi-

cally, Acroporidae and Pocilloporidae. These

corals, including the genera Montipora, Acro-

pora, Seriatopora and Pocillopora, are all small

polyped and branching or foliate in growth form

(Table 2). For these commonly grazed corals,

unlike the Japanese situation, predation appears

random and incomplete, resulting in scattered

patches of surviving corals in the grazed areas.

Corals with larger polyps (greater than 1 mm)

and massive growth forms are avoided and not

subject to predation by these molluscs, either in

/ol. 96(2)

April 21, 1982

THE NAUTILUS 77

FIG. 7. A, Diurnal aggregation (//' Drupella fragum under Acropora sp. at Toga Reef No. 1, Miyake-jima.

Japan, November. 1981; B, Nocturnal feeding y/'Drupella fraguni on Acropora sp. at Toga Reef No. 1. Miyake-

jima. Japan. November. 1981.

78 THE NAUTILUS

April 21, 1982

Vol. 96(2)

Vol. 96(2)

April 21, 1982

THE NAUTILUS 79

Table 1. Composition of dominant corals on

Mactan Island study sites

'able 2. Grazing preferences of Drupel la rugosa

at Mactan Island

^omnonly Grazed Corals

tontipora fruatiaosa

I. gvasi lis

1. alaiaormia

1. ramosa

1. foliosa

1. prolifera

^eriatopora hystrix

loderately Grazed Corals

'ocillopora vermoosa

^eriatopora oaliendmm

^onmonly Avoided Corals

tontipora aactus

1. verpuaosa

'OT^ tes

''avia

'avi tes

'latygyra

'■oniastrea

■oniopora

'yphastrea

■ardenoseris

'oeloseris

H.ploc2strea

Aaropora formosa

A. tenuis

A. florida

A. palifera

A. robusta

A. humilus

Aaropora hyaointhus

A. millipora

Galaxea

Pavona

Merulina

Euphyllia

Hydnophora

Peotinia

Lobophyllia

Symphy I Ha

Fimgia

Polyphyllia

Cyalosevis

Eahinopora

warms or as individuals (Table 2). Contrary to

he report by Demond (1957), wliich found Dru-

lella elata "most frequently on Porites sp.", and

hat of Taylor (1980) of Z). rugosa predation on

Writes colonies, Porites spp. appear to be en-

irely avoided by Drupella rugosa in the Philip-

lines. In addition, several highly tuberculate

(course surface projections) species of the most

commonly grazed genus, Montipora, are also

entirely avoided, suggesting a degree of host

selection.

Discussion

Robertson (1970) reported that species of

Drupella are known to be associated with scler-

actinian corals, and Fankboner (in Robertson,

1970) described feeding of Drupella comus

(Roding, 1798) on polyps of stony corals. Taylor

(1976, pp. 178, 185; 1978, pp. 95, 98) records

that Drupella comus and D. nigosa (Born, 1778)

feed on coral polyps. The literature, however,

suggests that not all the species of Drupella, at

least at the presently recognized generic alloca-

tions, may be obligate coral associates (Cerno-

horsky, 1969, p. 304; Robertson, 1970; Taylor,

1978, p. 99). This is further suggested by the

fact that four individuals of D. elata (Blainville,

1832) appeared on a species of Goniopora (Pori-

tidae) within a week after 6 of 8 parasitic muri-

cacean gastropods, Coralliophila uiolacea

(Kiener, 1836), were removed from the coral at

Miyake-jima. The coralliophilids had been ob-

served regularly over a period of six months in

the same Goniopora sp. with no appearance of

muricid predators during that period (M. Zaiser,

pers. cornm.). It is tempting to speculate that the

parasitic relationship of C. violacea and its

host may include some elements of mutualism.

Taylor (1978, p. 99), however, found on Can

Island, Addu Atoll, Maldives, that Drupella cor-

nus and Coralliophila violacea occur mostly on

different species of stony corals, and, as noted

above, Drupella rugosa avoids Poritidae in the

Philippines, but such avoidance is not necessar-

ily the case elsewhere (Taylor, 1980). The possi-

bility of competitive interactions between C.

violacea and Drupella spp. is currently under in-

vestigation by the senior author.

Our information on the composition of living

and grazed corals at the Mactan Island reef sites

suggests a degree of host selection by Drupella

rugosa, with specific preferences for Montipora

and Acropora and an obvious avoidance of

Porites, which is more common at Mactan than

Acropora (Tables 1 and 2). The effect of such

80 THE NAUTILUS

April 21, 1982

Vol. 96(2)

prey-preference is similar to that reported for

Acanthaster planci by Goreau, et al. (1972), and

Glynn (1976). These studies revealed a parallel

selection by Acanthaster for numerically com-

mon, fast growing species of Acropora, Monti-

pora. Pocillopora. Striatopora, and Stylophora.

as well as some faviids and fungiids. Other cor-

als appeared to be resistant to Acanthaster, e.g.

Millepora, Heliopora, Goniopora, and certain

species of Porites and Pocillopora (Goreau, et

al., 1972). Under "normal" conditions on tropical

reefs, such as those at Mactan, the results of se-

lective predation both by Acanthaster and Dru-

pella appear to be the same, i.e., a periodic

"weeding out" of the fast-growing, foliate

species (particularly the very abundant Monti-

pora at Mactan), providing space and settlement

sites for other corals and thus increasing diver-

sity over time. As discussed by Goreau, et al.

(1972) for Acanthaster predation, there will be

understandably different immediate and long

term effects as far as diversity and coral cover

are concerned, depending upon the intensity of

predation. At Mactan, Drupella rugosa appears

to represent an additional biological control

(with Acanthaster) on the coral community

structure. This seems to be characteristic on

tropical reefs elsewhere. R. Randall {pers.

comm.) reports that at Guam Drupella sp. feeds

exclusively on Pocillopora. Entire Pocillopora

colonies are destroyed, but destruction of large

segments of the reef, such as that occasionally

caused by Acanthaster population explosions,

does not occur.

The situation in the warm temperate waters

at Miyake-jima is quite different. As noted, true

coral reefs do not occur there, although coral

diversity is remarkable for such a high latitude.

Acropora spp. are dominant and form large

patches at certain localities, e.g. Toga Reefs

Nos. 1 and 2. However, other corals are rela-

tively scattered in isolated, small colonies on

the lava substrate (Tribble and Randall, MS).

Patches of branching Acropora are preferred

prey by Drupella fragum, and we have never

seen evidence of this snail feeding on other coral

genera.

Why Dnipella J'nuvun population explosions

began to occur in about 1976 at one site on

Miyake-jima and in 1980 at another site is a

puzzling question. The senior author has worked

on both sites since 1957, finding no evidence of

widespread coral destruction until 1976. It may

be more than a coincidence that heavy silting

from construction programs near the study sites

preceded each outbreak by from 2 to 4 years in

both cases. Beginning in 1972 and continuing

until 1974, large amounts of sand were pumped

with seawater through a large sandpipe from

the sandy interior of Igaya Bay to a filtering

center on shore, where the sand was collected.

The murky water, containing fine sediments,

flowed back into the bay, settling over coral

patches. Occasional breaks in the sandpipe

added to this siltation. During the same period,

a large pasture for dairy cattle was constructed

on the volcanic mountain above the bay. Lum-

bering, followed by bulldozing to level the land,

preceded the planting of cattle feed. Five dry

river beds run into Igaya Bay, all of which pass

through this recently constructed pasture.

Heavy rains during the period of bulldozing

flooded the rivers with muddy water and cut

visibility in the bay (normally 15-40m) to less

than 2 m on many occasions during this period.

Drupella destruction was noted from 1976, four

years after the beginning of both sand mining

and pasture construction.

Similarly, in 1976 a road was constructed to

Toga Bay for the purpose of development of a

marine park featuring Toga Bay's two large

coral patches (Toga Reef No. 1 and Toga Reef

No. 2). Silting from this road into the bay began

immediately, but increased greatly in 1978,

when the road was paved and drainage ditches

constructed to a point about 300 m above the

bay. Erosion along the remaining 300 m of road

since 1978 has been massive. Drupella destruc-

tion in Toga Bay first appeared in 1980, four

years after the construction of the road and two

years after severe siltation.

Although the relationship between massive

siltation and population explosions of Drupella

fragum at Miyake-jima must remain purely

speculative at this time, it is of interest to con-

sider the gastropod population explosions in the

light of Birkeland's (in press) hypothesis relating

Vol. 96(2)

April 21, 1982

THE NAUTILUS 81

cycles of Acanthaster planci destruction with

long periods of drought followed by heavy rains.

Birkeland presents evidence to show that heavy

rainfall following an extended dry season or

drought results in great increases in nutrients,

especially phosphorus, from terrestrial runoff

into the shallow waters near shore, thus stimu-

ating unusually lush blooms of phytoplankton.

This increased food supply permits Acanthaster

arvae to survive in much higher densities than

inder normal conditions, resulting in population

explosions and extensive coral damage three

/ears later, when the postlarvae reach maturity.

A similar phenomenon may account for the

Drupella fragum. explosions at Miyake-jima;

leavy siltation from human activities accom-

)anies tyjihoon or rainy season downpours, thus

ntroducing nutrients into coastal waters. Many

ihallow water marine invertebrates experience

)utbreaks at irregular intervals (Coe, 1956), and

;uch outbreaks have been related to terrestrial

lutrient runoff (see Sutcliffe, 1972, 1973). J. B.

Taylor (1975) identified 14 species of thaidid

'eligers (all four-lobed) in the plankton of

Caneohe Bay, Hawaii, including Drupella elata

as D. comtis). It is possible that Drurpella larvae

■at phytoplankton and that the heavily silted

orals may be unable to consume these larvae in

lumbers sufficient to control their population.

)f course, we recognize that this is pure conjec-

ure, but we present it for the purpose of stimu-

ating research on this intriguing problem.

ACKNOWLEDGMENTS

We thank the Fishermen's Cooperative of

Jiyake-jima and the Fisheries Divisions of the

I >\ernments of Tokyo Prefecture and Miyake-

ima for financial and logistic support during

his study. The efforts of J. A. Dalton, J. L.

)icksved, G. Hodgson, T. Takada, K. Tateyama,

1. Umeda and M. J. Zaiser in field observations

nd data collection are greatly appreciated. R.

uiiidall is thanked for sharing observations of

h-iipeUa predation at Guam. Dr. C. Birkeland

itidly offered expert advice and supplied impor-

int literature. Drs. R. T. Abbott, H. A. Rehder,

Is. B. W. Myers, and Mr. A. D'Attilio gener-

usly provided information on the taxonomy of

the Thaidinae and C. J. Ferraris, Jr. on Philip-

pine corals. Dr. J. Rosewater kindly supplied a

needed reference. W. E. Old, Jr. and S. S.

Horenstein and M. Werner, respectively, pro-

vided bibliographic assistance, photographed

the specimens (Figs. 1-3), and tj-ped the

manuscript. We are indebted to Dr. R. Robert-

son for critically reading the manuscript. We

also thank P. Kott and M. Cope.

LITERATURE CITED

.\rakawa, K. Y. 1958 ("1957"). On the remarkable sexual

dimorphism of the radula of Drupella. Venus (Japanese

Jour. Malacol.) 19(3-4):206-214.

Birkeland, C. (In press). Terrestrial runoff as a cause of out-

breaks of Acanthaster planci (Echinodermata: Asteroi-

dea). Mar. EcoL, Progress Series.

Blainville, H. M. D. de. 1832. Disposition methodique des

especes recentes et fossiles des genres Pourpre, Ricinule,

Licorne et Concholepas de M. de Lamarck. Nouv. Ann.

Mus. Nat. d'Hist.. Paris 1:189-263.

B(jrn, I. 1778. Index Rerum Naturalium Musei Caesarei

Vindobonetisis. Pt. 1: Testacea. Vienna, xiii + 458 p.

Branham, J. M. 1973. The crown of thorns on coral reefs.

BioScwnce. 23{4):219-226.

Cernohorsky, W. 0. 1969. The Muricidae of Fiji, Pt. 2, Sub-

family Thaidinae. The Veliger 11(4):293-315.

1972. Marine Shells of the Pacific. Pacific Pub-

lications, Sydney, 411 p.

1978. Tropical Pacific Marine Shells. Pacific

Publications, Sydney, 352 p.

Coe, W. R. 1956. Fluctuations in populations of littoral

marine invertebrates. Jour. Mar. Res. 15:212-232.

Cooke. A. H. 1895. The Cambridge Natural History. Mol-

luscs, vol. 3. pp. i-ix, 1-459. Macmilian & Co., New York.

Demond, J. 1957. Micronesian reef-associated gastropods.

Pacific Sci. 11(3):275-341.

Emerson. W. K. and W. 0. Cernohorsky. 1973. The genus

Drupa in the Indo-Pacific. Indo-Pacific Mollu.tca

3(13):801-864.

Endean, R. 1973. Population explosions of Acanthaster

planci and associated destruction of hermatypic corals

in the Indo-West Pacific region. In: Biology and Geology

of Coral Reefs, vol. 2, Biology 1, Jones, 0. A. and R.

Endean (eds.). Academic Press. New York, p. 390-438.

Fanklioner, P. V. 1970. Notes on feeding and the functional

morpholog\' of the gut in the reef dwelling muricid Morula

elata (Blainville). Western Soc. Malacologists. Abst. &

Proc. for 1969, pp. 15, 16.

Glynn, P. W. 1976. Some physical and biological determi-

nants of coral community structure in the eastern Pacific.

Ecological Monographs 46:431-456.

Goreau. T. F., J. C. Larg, E. A. Graham and P. D. Goreau.

1972. Structure and ecologj- of the Saipan reefs in relation

to predation by Acanthaster planci (Linnaeus). Bull.

Mar. Sci. 22(1):113-152.

Ida, H. and J. T. Moyer. 1974. Apogonid fishes of Miyake-

82 THE NAUTILUS

April 21, 1982

Vol. 96(2)

jima and Ishigaki-jima, Japan, with description of a new

species. Japan. Jour. Ichthyol. 21(3):113-128.

Melville, R. V. [Sec] 1980. International Commission on

Zoological Nomenclature. Opinion 1154, Dnipella Thiele,

1925 (Mollusca, Gastropoda): Designation of a type species

by use of the plenary powers. Zooi Norrmi. Bull.

37(2):85-88.

Moyer, J. T. 1978. Crown of thorns starfish invades Miyake-

jima.' Marine Park Jour. (Japan) 44(1): 17 (in Japanese).

Ponder, W. F. 1973. The origin and evolution of the Neo-

gastropoda. Malacologia 12(2):295-338.

Porter, W. P. 1972. Predation by Acanthaster and its effect

on coral species diversity. Amer. Natur. 106(950):

487-492.

Randall. J. E. 1972. Chemical pollution in the sea and the

crown-of-thorns starfish (Acanthaster planci). Biotropica

4(3): 132- 144.

Reese, E. S. 1977. Coevolution of corals and coral feeding

fishes of the family Chaetodontidae, p. 267-274. Third

International Coral Reef Symposium Proc. vol. 1, Univ.

Miami. Miami, Florida.

Robertson, R. 1970. Review of the predators and parasites

of stony corals, with special reference to symbiotic proso-

branch gastropods. Pacific Sn. 24(l):43-54.

Shepard, J. W. and J. T. Moyer. 1980. Annotated checklist

of the fishes of Miyake-jima, Japan. Pt. 1, Pomacentridae,

Chaetodontidae, and Pomacanthidae. Publ. Seto Mar.

Biol. Sta. 15(l-4):227-241.

Sutcliffe, W. H., Jr. 1972. Some relations of land drainage.

nutrients, particulate material, and fish catch in two east-

ern Canadian bays. Jour. Fish. Res. Bd. Canada 29(2):

357-362.

1973. Correlations between seasonal river dis-

charge and local landings of American lobster (Homarus

americanus) and Atlantic halibut (Hippoglossus hippo-

glossus) in the Gulf of St. Lawrence. Jour. Fish. Res. Bd.

Canada 30(6):856-859.

Taylor, J. B. 1975. Planktonic prosobranch veligers of

Kaneohe Bay. Ph.D. Diss. Univ. Microfilms Inst., Ann

Arbor, Michigan, 331 p.

Taylor, J. D. 1976. Habitats, abundance and diets of muri-

cacean gastropods at Aldabra Atoll. Linnean Soc. Zool.

(London) Jour. 59(2):155-193.

1980. Diets and habitats of shallow water preda-

tory gastropods around Tolo Channel, Hong Kong. In: The

malacofauna of Hong Kong and southern China, Morton,

B. (ed.) Univ. Hong Kong Press, p. 163-180.

1978. Habitats and diet of predatory gastropods

at Addu Atoll, Maldives, Exp. Mar. Biol. Jour. 3(1):

83-103.

Taylor, J. D., N. J. Morris and C. N. Taylor. 1980. Food

specialization and the evolution of predatory prosobranch

gastropods. Paleont. (London) 23(2):375-409.

Tribble, G. W. and R. Randall. [MS]. Description of coral

environment and community structure of the shallow

water corals of Miyake-jima, Japan.

Wu, S.-K. 1965. Studies of the radulae of Taiwan muricid

gastropods. Inst. Zool., Acad. Sinica Bull. 4:95-106.

SUCCINEA AVARA SAY FROM THE SOUTHERN GREAT PLAINS

OF THE UNITED STATES

Dorothea S. Franzen

Illinois Wesleyan University

Bloomington, Illinois 61701

ABSTRACT

Succinea avara Sa?/, 182Jt, was described from shells collected from the North-

west Territory. This study is o/S. avara collected at 21 stations in four states of

the Southern Great Plains Region of the United States. Shells and anatomical

structures are described and illustrated; habitats of a selected representative

series from the 21 stations are described.

Succinea avara Say, 1824

Sucnnea avara Say, 1824, p. 250; pi. 15, Fig. 6.

Succitiea avara Say, Pilsbry, 1948, pp. 837-840;

Fig. 442. B, b, C, c, D.

Shells of Succineidae present only a few dis-

tinctive characteristics; specific determinations

depend largely upon anatomical characteristics

of the soft parts. It is questionable whether the

anatomy of the soft parts as now identified with

a species actually corresponds to the anatomy of

the soft parts of the holotypes and paratypes

which are known only from their shells. It is im-

V^ol. 96(2)

April 21, 1982

THE NAUTILUS 83

possible to determine the exact location of the

type locality when geographically it is desig-

nated only as being in a general area, as iorSue-

Hnea avara Say, "Inhabits the Northwest Terri-

:ory" (Say, 1824, p. 260). The succineas taken

Tom a locality today may not be representative

)f the fauna existing fifty to a hundred or more

/ears ago. Such are the problems concerning

;he identification of S. avara Say.

The shells of the holotype and paratj^aes of

'^uccinea avara, Academy of Natural Science

Philadelphia No. 59542, are small and probably

mmature.

^ype description:

"S. avara. Shell suboval, pale reddish-yellow,

subdisphanous, fragile, covered with an earthy

crust; whorls three, minutely wrinkled; body

whorl very large; spire small; aperture large,

subovate, about two-thirds of the whole length

of the shell. Length three-twentieths of an

inch. Inhabits the North-west Territory. This

small species of Succinea occurs in humid

places very frequently under stones and near

the water. The shell is always completely in-

crusted with a coating of earth. It may at once

be distinguished from either the ovalis or of

campestris by its very small size. PI. 15,

fig. 6." (Say, 1824, p. 260.)

Pilsbry (1948, pp. 837-840, Fig. 455 and Fig.

42, A, a, B, b, C, c, D) described what he con-

idered to be S. avara. Fig. 442, B, b, are draw-

igs of the reproductive system of a specimen

rom Warm Spring Canyon, Arizona, collected

y Ferriss in 1909; C, c, are of a specimen from

lurricane Fault, Arizona.

The purpose of this paper is to describe shell

nd anatomical features, as well as habitats, of

uccinea avara Say of the Southern Great

'lains of the United States. Collections were

lade by Artie L. Metcalf and by me at 21 sta-

ons located in the western half of Texas, east-

rn and southern New Mexico, southern and

orthwestern Oklahoma and southwestern Kan-

as (Fig. 1).

Succineidae are generally regarded as being

mphibious- living on wet ground, in woods,

long shores, or appearing on the surface

allowing a rainfall heavy enough to soak the

round. However, we have sometimes found

Fid. 1 . Four-state map shomng locations of the 21 collecting

sidlionti.

them in habitats far removed from wet areas.

At the stations where S. avara was obtained,

the annual precipitation ranges from about 9 to

44 inches. All stations are subjected to periods

of from 3 to 5 months in which the precipitation

is less than an inch per month (Yearbook of

Agriculture, 1941). S. avara included in this

study were taken in early spring, summer, and

early winter months during which time the

snails were either active or aestivating. They

were found in damp places under rocks, logs,

boards or dead stems of Yucca sp. The following

are the locations of the 21 collecting sites and

descriptions of habitats of a selected representa-

tive series often of those sites. The field number

designations and descriptions of habitats are by

Artie L. Metcalf. My field numbers, enclosed in

parentheses, serve as a reference to my records.

Sites of my collections are indicated by my field

numbers and the descriptions of such habitats

are mine.

Stations

1. Field No. ALM 968 (DSF 402); 30 June, 1969, along the

Cimarron River, along New Mexico Rd. 325; 7.5 mi W of its

junction with New Mexico Rd. 370, Union County. New Mex-

ico. Altitude: 5,250 ft. above mean tide (AMT). Annual

precipitation about 17". Habitat: under logs on moist

floodplain of the river and also under rocks on adjacent

hillside.

2. Field No. ALM 1058 (DSF 412); 30 ,Iuly, 1970; 2.5 mi N

of Mt. Dora, Union County, New Mexico.

84 THE NAUTILUS

April 21, 1982

Vol. 96(2)

3. Field No. ALM 1060 (DSF 41 1); 31 July, 1970; valley of

North Canadian River, 0.1 mi N of bridge over river on New

Mexico Rd. 370; about 22 mi N NW of Clayton (S 33, T 29 N,

R 52 W), Union County, New Mexico.

4. Field No. ALM 673, DSF .342; 8 May, 1966; .5 mi SE of

Rincon; HW^ 1-25 at junction with Engle Road, Dona Ana

County, New Mexico. Altitude: 4370 ft. AMT. Annual pre-

cipitation about 9.5". The locality is in Jornada del Muerto-

a basin between two mountain ranges. Habitat: Succinea

avara were aestivating on dead stems of Yucca data

Engelm. growing in a "tobosa grass swale" where the

predominant plant is tobosa grass, Hilaria mutica (Buckl.)

Benth. The snails were found on the side of the dead Yucca

stems lying on the ground where there was a trace of

moisture.

5. Field No. ALM 948 (DSF 398); 19 November, 1968:

valley of Tularosa River and falls, 0.5 mi SW of bridge over

Tularosa River, U.S. Hwy 70; 8.5 mi SW of Tularosa (NW '/i,

SW 'A, S 32, T 13 S, R 11 E), Otero County. New Mexico.

6. Field No. ALM 946 (DSF 397); 19 November, 1968;

Mescalero Indian Reservation, Valley of the Tularosa River,

1.05 mi S of Tularosa Spring (SW 'A.SW 'A. S 7, T 14 S, R 13

E), Otero County, New Mexico.

7. Field No. ALM 998 (DSF 407); 11 October, 1969, and

19 September, 1970; junction of Pehasco and Cox Canyons

(SW 'A, S 3, T 17 S, R 13 E), Otero County, New Mexico.

.•Mtitude: 7200' AMT. Annual precipitation about 22".

Habitat: grassy, riparian meadow where the Penasco River

is about 5 ft. wide. The canyon wall is wooded with pre-

dominantly Ponderosa pine.

8. Field No. (DSF 492); collected by Artie Metcalf 23

September, 1978; east-central part of Nash Draw Basin

(center of S 15, T 22 S, R 30 E), 20 mi E of Carlsbad, Eddy

County, New Mexico.

9. Field No. ALM 670; 8 February, 1966, and DSF 339, 7

May, 1966; Franklin Mountains (S 27, T 28 S, R 4 E), about 5

mi NE Canutille, El Paso County, Texas. Altitude: 4900 ft.

AMT. Annual precipitation about 9". Habitat: under dead

Dasylirion sp. and under moist limestone rocks. A light rain-

fall which occurred the first of May resulted in certain areas

being relatively moist.

10. Field No. ALM 1396 (DSF 426); 9 August, 1972; 0.6

mi SW of Dalhart on southeast side of U.S. Hwy 54 and

bordering Rita Blanca Creek, Hartley County, Texas.

11. Field No. ALM 1283 (DSF 418); 19 June, 1971; 2.3 mi

W Channing, south side of Texas Farm Rd. 767, Hartley

County, Texas.

12. (DSF 389); collected by Artie Metcalf 23 August,

1968; flood plain of Wolf Creek, 1 mi E of dam on Lake

Fryer, Ochiltree County, Texas.

13. (DSF :',90); collected by Artie Metcalf 23 August,

1968; bluffs along north shore of Lake Fryer, Ochiltree

County. Texas. Altitude: 2650 ft. AMT. Annual precipitation

about 19.31". Habitat: under loose limestone below scarp in

Ogallala Formation.

14. (DSF 392); collected by Artie Metcalf 13 September,

1968; bluffs on southeast .-side of Buffalo Lake, 3 mi S of Um-

barger, Randall County. Texas.

15. Field No. ALM 1801 (DSF 476); 20 March. 1977; 9 mi

ENE or 6.6 mi W of junction of Ranch Rd. 2084 (to

Christoval) with Texas Hwy 29. S side of Texas Hwy 29.

Schleicher County, Texas.

16. Field No. ALM 1451 (DSF 430); 16 April, 1973; east

side of conical hill immediately E of old Fort Lancaster; N of

U.S. Hwy 290, Pecos County. Texas. Altitude: 2075 ft.

AMT. Annual precipitation about 19". Habitat: under lime-

stone rocks on E side of conical hill. The vegetation consists

of sparse grass and low xeric shrubs.

17. Field No. ALM 1458 (DSF 428); 18 April, 1973; under

limestone rocks opposite Judge Roy Bean Visitor Center,

Langtry, Val Verde County, Texas.

18. Field No. DSF 422; 14 July, 1972; Buffalo Bayou,

Texas Hwy 6, 1 mi S of junction with HWY I-IO, near Ad-

dicks, Harris County, Texas. Altitude: 50 ft. AMT. Annual

precipitation about 44". Habitat: Succinea avara found

aestivating on a trunk of a pecan tree up to 7' above ground

level and on a fence rail 5' above ground level. The ground

vegetation comprised nettles and grasses which may have

been too dense for the succineas; therefore, they sought

more open areas.

19. Field No. DSF 387, 28 July, 1968, and 17 July, 1972;

west bank of a bay of Lake Texoma, University of Oklahoma

Biology Station, 2 mi E of Willis and Oklahoma State HWT

99, Marshall County, Oklahoma. Altitude: about 500 ft.

AMT. Annual precipitation about 38". Habitat: the snails

were aestivating under boards and logs lying on the un- '

shaded bank of the lake; temperature at time of collection,

95°F.

20. Field No. ALM 1066 (DSF 414); 1 August, 1970; 6 mi

S Turpin; IV2 mi N of Beaver Creek, Beaver County, Okla-

homa. Altitude: 2600 ft. above AMT. Annual precipitation I

about 18". Habitat: under stones of bluff of Ogallala Forma-

tion. The area is a short grass prairie with Yucca glauca Nut

tall.

21. Field No. ALM 1559 (DSF 442); 1974; Point of Rocks,

an escarpment of Ogallala Formation, 3 mi W of Kansas

Hwy 27 on N side of Cimarron River, Morton County, Kan-

sas. Annual precipitation about 16.2". Habitat: most of the

Succinea avara were under a single large stone lying at base

of scarp. Yucca glauca NuJt. and Rhus trilobata Nutt. were

common plants. Grama grass and annual ragweed were

abundant.

The Shell: Shell is conically-ovate, imperfo-

rate, amber, somewhat glossy, translucent, usu-

ally encrusted with grains of sand or soil. Com-

posed of 2V2 to 3% whorls, the shell attains a

height of 12.6 mm and a width of 6.6 mm (Table

1). Whorls inflated, sharply incised, increasing

rapidly in size from the knoblike nuclear whorl

to the large, inflated body whorl. Aperture

ovate, ranging from 47.6% to 77.7% of the

height of the largest shells obtained at the 21

stations, bounded by a sharp peristome which

Vol. 96(2)

April 21, 1982

THE NAUTILUS 85

TABLE 1. DtmermoTis of shells o/Succinea avara Say of selected ten oftli£ tumity-one collecting sites. The nwtisurements are

of the three largest shells of each locality. Ratios of measurements are listed in the last four columns.

■'.ontinues over the body whorl as a thin callus.

Columella is gently curved (Fig. 2). Surface of

the nuclear whorl appears to be, at high magnifi-

cation, malleated. Surface of the remaining

86 THE NAUTILUS

April 21, 1982

Vol. 96(2)

FIG. 2. Shells q/'Succinea avara Say of four- collecting stations: A, Station 19. Marshall County, Oklahoma

(Height 8.5 mm): B, Station 11, Hartley County, Texas (Height 11.1 mm): C, Station 13, Ochiltree County, Texas

(Height 11.2 mm): D, Station 1. Union County. New Mexico (Height 10.5 mm).

whorls are marked with striae, fine on the pe-

nultimate whorl, becoming heavier and coarser

producing a rough appearance on the ultimate

whorl, especially near the aperture (Fig. 2).

Body and Mantle Surfaces: Surface of translu-

cent, gray-white body is smooth; low tubercles

may be observed on head region. On either side

of the lower lateral body wall a suprapedal

groove, paralleled by a shallow pedal groove, ex-

tends from the labial palp to the posterior tip of

the body. Lateral body wall incised by shallow

vertical grooves which extend to the pedal

groove, resulting in a scalloped edge. Genital

aperture elongate-crescent, about 0.7 mm in

length, surrounded by a white, tumid lip, and

situated on the anterior right-hand side of the

body.

The bodies oi Surcinea avara of the geograph-

ical region included in this study are generally

so lightly pigmented that it is difficult to

recognize any distinct pattern. When intense

enough to be discerned, pigmentation is seen to

form a pair of median bands on the anterior-

most portion of the head. Just anterior to the

superior (posterior) tentacles these bands fuse

and expand to form a triangle. From this, at

level of the tentacles, a single band extends to

where the mantle attaches to the body. On the

anterior portion of the head each of the two me-

dian bands is flanked laterally by a not very

distinct band which extends to the base of the

tentacles. Blotches of pigment on the tentacles

form, at the base of each, a band which parallels

the median band to the edge of the mantle. Also

scattered flecks of pigment are to be noted. On

either lateral body wall a faint band of flecks of

pigment may run above and parallel to the

suprapedal groove. The sole of the foot exhibits

faint longitudinal bands of pigment.

Bands of pigment extend from the mantle col-

lar across the mantle surface, coalesce as they

approach the nephridium. Pigmentation in the

form of a fine line outlines the anterior margin

of the nephridium, diffuses over its surface, and,

Vol. 96(2)

April 21, 1982

THE NAUTILUS 87

as a heavier band, outlines its posterior margin.

The nephridium is white.

Reproductive System: The stout penis and

epiphallus fill the thin, translucent, non-pig-

mented or lightly flecked, penial sheath. Junc-

ture of epiphallus and penis is marked by a con-

■;triction (Fig. 3A). Epiphallus and penis may be

contained entirely within the sheath (Fig. 3B) or

:he epiphallus and the distal portion of the penis

•nay extend through the distally open-ended

sheath (Fig. 3B). Because the thin sheath tends

;o cling to the penis, it is difficult to note that it

s open-ended. Such variations occur within any

)f the populations included in this study. The vas

deferens enters the penial sheath near its distal

end, emerges from the sheath as it enters the

epiphallus terminally (Fig. 3A, B). The broad,

stout penial retractor muscle attaches to the

distal end of the epiphallus (Fig. 3A, B).

Prostate gland, globular, composed of small

acini, enclosed within a very thin, finely pep-

pered sheath, is approximately one-third the

size of the albumin gland. Albumin gland, gener-

ally triangular, composed of small acini, is

enclosed within a very thin, finely peppered

sheath. Hermaphroditic duct may be slightly or

moderately pigmented. Lobes of the bilobed

seminal vesicles, unequal in length, are gener-

ally narrow, elongate, and lightly to moderately

pigmented. Juncture of uterus and oviduct is

marked by a slight constriction. Spermathecal

duct is broad at its base where it enters the

uterus (Fig. 3A, B).

Radula and Jaw: Structure of the radula of

Succvnea avara exhibits characteristics of the

genus. Number of rows of teeth range from

about 70 to 90. Formulae of representative rows

of teeth are to be noted in Table 2. Ratio of

laterals to marginals approaches 1:1 as found to

be true of S. ovalis Say (Franzen, 1959, Table H)

and of S. vaginacontorta Lee (Franzen, 1971,

Table H).

Structures of representative teeth are illus-

trated in Fig. 4A. The central tooth has a long,

pointed mesocone which equals or exceeds

TABLE 2. Formulae of representalioe rows of teeth of

radulae o/Suceinea avara Say.

No. of Rows

of Teeth

Row

M

STATION

No. 402

14 8-11-1-10-10

32 10 - 9-1-10-10

60 - 1 - 15 - 6

61 - 1 - 15 - 6

62 - 1 - 13 - 8

8 - 18 - 1 ■ 18 -

'IG. 3. Reproductive organs of Succinea avara Say. PS.

eniai sheath; EP. epiphallus: PRM, penial retrattor mus-

le; VD, vas deferens: VA, vagina: SPD, spermathecal duct:

>D, oviduct: SP, spermatheca: HD, hermaphroditic duct:

iG, albumin gland: GA, genital atrium: PG. prostate gland:

'S. fertilization sac: SV, seyninal vesicle.

No. 428 (Slide 2)

88 THE NAUTILUS

April 21, 1982

Vol. 96(2)

The amber-colored jaw is illustrated in Fig.

4B. Anteriorly the collar has a pointed median

fold and posteriorly a rounded indentation. The

posterior margin of the basal plate has a pointed

median fold flanked on either side by a rounded

boss.

FIG. 4. A, Representative radula teeth of Succinea avara

Say. C, central tooth; 1-L-L, first left lateral; 1-L-M, first left

marginal; 2-L-M, second left marginal; 7-L-M, seventh left

marginal; 8-L-M, eighth left marginal. B, A jaw o/Succinea

avara Say.

length of the basal plate; mesocone is flanked on

either side by a small ectocone. Laterals have a

long, sharply pointed mesocone which, gener-

ally, exceeds length of the basal plate; there is a

single smaller ectocone; endocone is generally

wanting. Differentiation between laterals and

marginals is not always sharply defined; fre-

quently an endocone is present on the outermost

one to three laterals. There may be on either

side of a lateral with a split ectocone a tooth

with an undivided ectocone. Marginals are char-

acterized by a short endocone, a long, sharply

pointed mesocone, and an ectocone which is

divided into two, three or four cusps. In 5. ovalis

and S. vaginacontorta the three small cusps of

the ectocone are short and almost equal in size.

In S. avara the two medial-most cusps of the ec-

tocone are long; the outermost cusps are minute

(Fig. 4A). The basal plates of the marginals are

as in S. ovalis Say (Franzen, 1959, Fig. 3) and as

inS. vaginacontorta Lee {Franzen, 1971, Fig. 3).

They are not long and tapering as in the genus

Oxyloma (Franzen, 1963, Fig. 1) nor as short

and broad as in the genus Catinella (Quick,

1933, Fig. 4); (Franzen, 1979, Fig. 3).

ACKNOWLEDGMENTS

National Science Foundation Grants-in-Aid

No's. NSF G18000 and NSF GB2715 provided

laboratory equipment and supported, in part,

field studies. The Academy of Natural Sciences

of Philadelphia graciously lent shells of the

holotype and paratypes of Succinea avara Say.

Dr. A. Byron Leonard read the manuscript and

offered helpful suggestions.

LITERATURE CITED

Franzen, Dorothea S. 1959. Anatomy of Succinea ovaltK

Say. Proc. Mai. Soc. London 33(5):193-199, Tables MI,

Figs. 1-7.

1963. Variations in the Anatomy of the Succi

neid Gastropod Oxyloma retusa (Lea). The Nautilus

76(3):82-95, Tables MI, Figs. 1-4.

1971. Anatomy and Geographic Distribution of

the Succineid Gastropod Succinea vaginacontorta Lee.

The Nautilus 84(4):131-142, Tables MI, Figs. 1-3.

1979. Catinella parallela. a New Succineidae

(Puimonata) From Midwestern United States. The Nauti-

lus 93(2-3):63-69, Tables 1-2, Figs. 1-3.

Pilsbry, Henry A. 1948. Land Mollusca of North America

(North of Mexico). Acad. Nat. Sci. Philadelphia Mon. No.

3. Vol. II, Ft. 2: xlvii + 521-1113, 585 figs.

Quick, H. E. 1933. The Anatomy of British Succinea. Proc.

Mai Soc. London 20(6):295-318, Tables I-V, PI. 23-25,

Figs. 1-18.

Say, Thomas. 1824. In Appendix to Keating's Narrative

Expedition . . . source of St. Peter's River, etc. (North-

west Territory) 2:260, PI. 15, fig. 6.

Yearbook of Agriculture. 1941. Climate and Man. United

States Department of Agriculture, U.S. Government

Printing Office, Washington, D.C.: v-xii and 1-1248.

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THE

NAUTILUS

ISSN 0028-1344

Vol. 96

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Dr. David H. Stansbery

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THE

NAUTILUS

Volume 96, number 3 - July 26, 1982

ISSN 0028-1344

CONTENTS

Eileen H. Jokinen

Cipangopaludina chinensis (Gastropoda: Viviparidae) in North America,

Review and Update 89

Hans Bertsch and Barbara Myers

Comments on the Eastern Pacific Trivia ritteri (Gastropoda: Triviidae) 96

Bretton W. Kent

An Overlooked Busycon Whelk (Melongenidae) from the Eastern United States 99

M. G. Harasewych and Richard E. Petit

Notes on the Morphology ofCancellaria reticulata (Gastropoda: Cancellariidae) 104

William Miller, III and Sheree Kooser

The Distribution of Donax variabilis (Bivalvia) at Ship Island, Mississippi:

I Some Paleomalacological Aspects 113

J. Gibson-Smith and W. Gibson-Smith

Divarilima (Bivalvia: Limidae) and a New Subspecies from the Caribbean 115

J. Gibson-Smith and W. Gibson-Smith

The Subfamily Melampinae (Pulmonata: Basommatophora) in Venezuela,

with Descriptions of Two New Species 116

J. Gibson-Smith and W. Gibson-Smith

An Ancestral Stephopoma (Mollusca: Gastropoda) from the Tertiary

„ of Venezuela 120

K. E. Hoagland and Wesley R. Coe

Larval Development in Orepidula maculosa (Prosobranchia: Crepidulidae),

from Florida 122

Errata 96

COMPENDIUM OF SEASHELLS

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Two internationally known experts on mollusks have combined their knowledge and writing

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Vol. 96(3)

July 16, 1982

THE NAUTILUS 89

CIPANGOPALUDINA CHINENSIS (GASTROPODA: VIVIPARIDAE)

IN NORTH AMERICA, REVIEW AND UPDATE

Eileen H. Jokinen

Ecology Section, U-42

The Univeristy of Connecticut

Storrs, CT 06268

ABSTRACT

The Asian freshwater viviparid snail. Cipangopaludina chinensis (Gray), was

introdiwed into North America in the 1890's. The species has spread across the

United States and southern Canada and is well-established in the northeastern

U.S. Connecticut populations are limited to lakes and ponds of medium-hard to

hard waters of calcium levels in excess of 5 ppm. The shell growth ofC. chinensis is

allometric with the young snails having a lower shell height to shell width ratio

than the adults. The growth patterns and radular cusp number of the typical

chinensis and the japonicus morph differ from, each other. A literature review

notes the five-year life span, diatom diet, the importance of quality food sources in

regulating population parameters, and the role ofC. chinensis as a possible host

for Asian helminths.-

The Asian freshwater prosobranch snail

Cipangopaludina chinensis (Gray, 1834)

(Synonym; Viviparus chinensis malleatus

(Reeve, 1863)) was first introduced into North

America in the 1890's (Abbott, 1950). Since

then, malacologists have been documenting the

spread of populations of C. chinensis. This

species has become well-established in parts of

North America and it is time to review the

known North American biogeography, ecology

and natural history of C. chinensis. This paper

will also add information on the snail's en-

vironmental preferences, radular structure, and

shell growth pattern.

Dundee (1974) and Clarke (1978) have syno-

nymized C. japonicws (von Martens, 1861), with

C. chinensis. Clench & Fuller (1965) believed

that the two were different species with in-

dividuals of C. japonicus having more elongate

shells (higher height/width ratios) with smaller

apical angles than C. chinensis. Morphological

intergrades exist (Clench & Fuller, 1965), indi-

cating either hybrid populations of two species,

or different morphological types within one

species. Apparently Clarke (1978) has decided

upon the latter explanation but extensive com-

parative morphological and immunological

studies have not been published. Therefore, this

paper will differentiate, when possible, between

topical C. chinensis and C. japonicus by in-

dicating "japonicus" forms as "japonicus"

morphs, thereby cautiously following a middle

ground that may easily be assimilated into a

final resolution of the problem.

C. chinetisis was first brought to North

America in the early 1890's by sailors returning

from Yokohama, Japan. The sailors sold the

snails to merchandize dealers in San Francisco's

Chinatown (Wood, 1892a, 1892b; Stearns, 1901).

Although Wood called these specimens "japoni-

ca", the snails were actually C. chinensis, the

first introduced japonicus not being recorded

until 1911 (Hannibal, 1911; see discussion in

Clench & Fuller, 1965). A second west coast in-

troduction was reported by Rev. G. W. Taylor

who found "japonica" being sold in a Chinese

market in Victoria, British Columbia (Anony-

mous, 1894). It is not known whether these

snails were of the japo7iic2is or the chinensis

tj'pe (Clench & Fuller, 1965), and there has been

no report of established Asian viviparid popula-

tions in British Columbia. Additional west coast

I

90 THE NAUTILUS

July 16, 1982

Vol. 96(3)

colonies have been reported from California and

Washington (Clench & Fuller, 1965; Hanna,

1966; Branson, 1977).

Individuals of C. chinensis were first found on

the east coast by W. Clench who collected speci-

mens in Massachusetts from the Muddy River, a

stream dividing Boston from Brookline. The

snails may have been introduced when goldfish

were added to the stream for mosquito larvae

control (Johnson, 1915). Additional east coast

colonies have been reported from Quebec,

Maine, Vermont, New Hampshire, Massachu-

setts, Rhode Island, New York, New Jersey,

Pennsylvania, Delaware, Maryland, North

Carolina, and Florida (Johnson, 1916, 1918;

Richards & Adams, 1929; Schmeck, 1942;

Jacobson & Emerson, 1961; Clench & Fuller,

1965; Stanczykowska et aL, 1971; Perron &

Probert, 1973; Bucci, 1974; Dundee, 1974;

Clarke, 1978). Additional populations have

recently been found in Connecticut (see below

for details).

C. chinensis populations have also spread mid-

continentally in Ohio, Michigan, Wisconsin,

Indiana, Minnesota, Arizona, Colorado, Texas,

Iowa, Oklahoma, and Utah (Teskey, 1954; Bran-

son, 1959; Clench & Fuller, 1965; Huehner &

Etges, 1971, 1972; Barnhart, 1978). Figure 1 is

a map of all the recorded references to North

American populations of C. chinensis north of

Mexico (excluding Hawaii). This species is very

well established on the northeast coast where it

should now be considered as a permanent part

of the freshwater molluscan fauna. The species

is also becoming well entrenched in the St.

Lawrence-Great Lakes drainage. Is is also com-

mon on the west coast between San Francisco

and Seattle. Additional colonies are scattered

across the continent. The paucity of mid-

continental colonies may reflect a lack of infor-

mation but it should be noted that some reports

of recent continental malacological surveys

mention no colonies of C. chinensis (Harman &

Berg, 1971; Clarke, 1973).

Connecticut Records

Over the past several summers I surveyed 215

aquatic sites in southern New England, especial-

ly Connecticut, for gastropods. Populations of

typical C. chinensis have been found in seven

FIG. 1. Thu \iirth American 1982 distributiov q/'Cipangopaludina chinensis (Gray). See text for references.

Vol. 96(3)

July 16, 1982

THE NAUTILUS 91

lakes and ponds in Connecticut and one lake in

\'ermont (Lake Fairlee, Thetford Center, pre-

viously reported in Clench & Fuller, 1965). A

population of the japonicus-morph lives in one

lake in Connecticut (Black Pond). Table 1 sum-

marizes the physical/chemical data on the Con-

necticut sites. Methods of water chemical analy-

sis have been described elsewhere (Jokinen,

1978).

Populations of C. chinensis were generally

found in larger lakes or ponds with sandy to

muddy sand substrates. The snails were either

crawling on the substrate or on dock pilings.

Even though several investigators have noted

C. chinensis living in slower waters of streams

(Schmeck, 1942; Stanczykowska et ai. 1971) all

Connecticut sites were lentic.

Due to the fact that most of the bedrock in

Connecticut is of the highly insoluble schist-

gneiss type, the majority of the surface waters

(75% of those sampled) are very soft (calcium

values below 5ppm). Yet, five of the eight C.

chinensis populations were found in harder

waters (Ca*-' > lOppm) and none lived in waters

with Ca < 7ppm. If one statistically compares

expected to actual percent occurrence of C.

chinensis in soft (Ca < 5ppm) waters (56% ex-

pected occurrence, 0% actual) to medium-hard

to hard (Ca > 5ppm) waters (44% expected,

100% actual), using an arcsin transformation

(Sokal & Rohlf, 1969), there is a statistically

significant difference at the t = .02 level. This

indicates that C. chinensis is more likely to be

found in waters of higher calcium values than

Table 1. Physical/chemical data for the habitats of Clpangopaludina chinensis in Connecticut.

Lake or pond

Area CO. Cond. Ca Mg

Type ha S pH mgC/1 ymhos Color ppm ppm

Black Pond, Middlefield,

Middlesex Co.

Bunnell's Pond, Bridgeport,

Fairfield Co.

East Twin Lake, Salisbury,

Litchfield Co.

Goodwin Park Pond, Wethers- NR

field, Hartford Co.

Holbrook Pond, Hebron,

Tolland Co.

North Farms Reservoir,

Wallingford,

New Haven Co.

Tyler Pond, Goshen,

Litchfield Co.

Lake Wononskopomus ,

Salisbury ,

Litchfield Co.

NR 30.6 11 7.3 7.6 194 .024 11.4 4.5

A 16.6 10 7.1 3.5 207 .025 9.9 3.3

N 227.6 15 7.4 33.3 286 .005 35.0 11.6

1 7.0 14.2 280 .222 19.6 4.2

NR

NR

1.8

29.3 8 6.4 4.5

74 .059 7.1 2.0

25.3

73.7

7.5 8.6 158 .041 14.0 3.5

6.9 7.2 112 .030 7.8 4.1

NR 142.7 10 7.5 22.8 217 .010 26.6 8.3

Lake Type: N = natural, NR - natural, water level raised, A = artificial impoundment

S = total number of snail species in lake

-I- -I-

Na K

ppm ppm

8.2 0.5

22.0 2.9

3.9 2.1

11.9 2.0

4.8 0.3

5.5 1.8

3.1 0.5

4.5 1.6

I

92 THE NAUTILUS

July 16, 1982

Vol. 96(3)

lower, and that it may have a physiological need

for dissolved calcium levels above 5ppm.

Boycott (1936) termed mollusks which could not

exist in soft water as "calciphilic". Greater sam-

ple sizes and physiological tolerance testing are

needed for greater resolution of required envi-

ronmental calcium values.

Natural history and Ecology

Data on the life history pattern of C. chinensis

has been published by Stanczykowska et al.

(1971, 1972). They estimated the maximum life

span for female snails as five years, for males,

usually three years, sometimes four. All individ-

uals grew throughout life, with the female's

reaching larger sizes because of their longer life

span. The females may begin to produce em-

bryos by the end of their first year but the four

and five year classes produced the most off-

spring. Each female produced a minimum of 169

embryos during a life time (an estimation based

on embryo counts published by Stanczykowska

et aL, 1971). Crabb (1929) found up to 102 em-

bryos in one female. The embryos develop in the

uterus, and during spring and summer 100% of

the females are carrying embryos. The young

begin to appear in the population by June and

continue to be born through October. The snails

begin a migration into deeper water in October,

the females appearing to migrate first (Stanc-

zykowska et al., 1971).

Quality of food affects the population para-

meters of C chinertsis. Stanczykowska et al.

(1972) and Plinski et al, (1978) compared snail

populations from three different habitats near

Montreal, the Ottawa River, a canal, and a lake.

The snails fed on the bottom material which was

composed of inorganics, organics and algae. The

percent composition of substrate materials was

identical to the percent composition of the gut

contents, indicating that the snails were not

selectively ingesting any specific bottom mate-

rial. Most of the algae ingested were epiphytic

or benthic, indicating that feeding was primarily

radular and not by filtration. The ingested algae

were primarily diatoms, especially species of

Fragilaria, followed by greens, blue-greens,

and flagellates. The percent composition of algal

forms differed between the three habitats. The

Ottawa River snails were larger, heavier, had

higher carbon and nitrogen values, and a

greater population density than snails from the

other two habitats. Sediment and gut analysis

revealed that the Ottawa River sediments had

not only a higher percentage of algae but a

higher diatom content. The lake snails made

some attempt to compensate for poorer quality

and quantity by consuming greater quanitities

of substrate. Increase in ingestion rate with a

decreased food quality has also been noted for

pulmonate snails (Calow, 1975). No studies have

been done to date on why diatoms appear to be

more nutritious to C. chirieyisis than other algae.

The entire area of freshwater gastropod feeding

and nutrition is poorly explored.

Allometric Shell Growth

Shell growth in C. chinensis is allometric, that

is, the proportions of the shell width to the shell

height change throughout development. Conse-

quently, the shell looks different in juvenile and

adult individuals. The subject of allometric

growth has been explored by Gould (1966 &

1971), White & Gould (196.5), and Jokinen (in

review). Allometric growth may be analyzed by

using linear regression and the resultant curves

graphed. Figure 2 illustrates the allometric

growth pattern of "typical" C. chinensis and

compares it to the growth pattern of the japoni-

CTis-morph. Shell measurements were made to

the nearest 0.1 mm with vernier calipers and in-

cluded embryonic shells. The raw data are avail-

able from the author. The data were analyzed

using the computer program SAS (Statistical

Analysis System) (Barr et al., 1979) and both

linear and curvilinear regressions (SAS-General

Linear Models and Non-linear Models) were run

on shell height vs. width and log shell height vs.

log width to determine which model best fit the

data. Analysis proved that shell growth in both

groups is best described by a linear log shell

height vs. log width curve. Log shell height was

used as the independent variable.

The ratio of the shell width to the shell height

(W/H) decreases as the shell increases in size,

the shell becoming relatively more elongate as it

Vol. 96(3)

July 16, 1982

THE NAUTILUS 93

18-

1 6-

X

I-

Q 14

12-

_l

—I

1^10

X

o

6-

C. CHINENSIS

» JAPONICUS-

MORPH

EMBRYO

.4 .6 .8 10 12 14

SHELL HEIGHT

16

OQi

0

FIG. 2. Allometric shell growth o/C. chinensis and the ]a.pomcus-morph. Sketches

illustrate the shape of the shells at various points on the curves. The scale bars

represent 5 mm. The regression equation C. chinensis is logW = 0.81, (±0.00U S.

E.jlogH + 0.17. The regression equation for the ]a.'pomais,-morph is logW = 0.78

(±0.01 S. E.jlogH + 0.23. The standard error values do not overlap, indicating

that the two growth patterns are significantly different. The regression coeffi-

cients (slopes) are less than one. meaning that the shell width increases in size at

a slower rate than the shell height. The shells, therefore, elongate as they grow. If

the slopes equaled unity, both height and width would grow at an equal rate and

the shell shape would not change with increase in size.

grows. The regression equation for the relation-

ship between height and width for "typical" C.

chiyiettms is logW = 0.84 logH + 0.17, r-square

= .998. A value of less than unity for the regres-

sion coefficient (slope), in this case 0.84, indi-

cates negative allometry, the shell width not in-

creasing as fast as shell height. The regression

equation for the japonicus-morph, the popula-

tion from Black Pond, is logW = 0.78 logH -i-

0.23, r-square = .997. The lower slope value,

0.78, indicates that the shell width of the japo-

nicM^-morph increases even less rapidly than C.

chinensis shell width. This is reflected in the

relatively more elongate shells of the japonicws-

morph.

Young individuals of C. chinensis not only

have a different shaped shell from the older

snails but demonstrate marked periostracal

spiral hirsuteness. When embryonic shells are

placed in water or alcohol and examined under a

dissecting microscope, they show a pattern of

three spiral rows of stiff periostracal hairs. The

hairs are distinctly hooked and the longest hairs

are 0.35 mm in length. The hooks face outward

from the shell. Some of the larger shells show

rows of pits in the same position as the em-

bryonic hairs (see Fig. 2). Since the hairs

become obliterated as the snails grow post-

partum, their function, if one exists, is most

likely intra-uterine. The hairs might serve to

hook the embryonic shells together to stabilize

their positions in the uterus, or they might func-

tion during birth.

94 THE NAUTILUS

July 16, 1982

Vol. 96(3)

Radular Structure

Radulae of C. chinensis individuals were ex-

amined for basic structure (Fig. 3). The teeth

were separated from each other and mounted in

Turtox's CMC-9AF mounting medium which

contains acid fuchsin for staining chitinous

material. The stain picked-up certain structural

details of the central tooth where the radular

material is thickened.

C. chinensis has the typical taenioglossid

tooth number of 7 per row. The central, first and

second laterals all have a large central cusp

bounded on either side by four smaller cusps.

The marginal teeth do not have an enlarged cen-

tral cusp but have seven small cusps (this

number appears to be somewhat variable).

Radular structure is similar to that of Viviparus

georgianus Lea (as illustrated in Clench, 1962)

except for the lack of an enlarged central cusp

on the marginals of C. chinensis. An examina-

tion of the radular structure of one individual of

the Black Pond population {japonicus-mov\>\\)

revealed five small cusps on either side of the

central cusp on the first lateral tooth and nine

cusps on the marginal tooth. It is not known if

these differences reflect individual variation (if

the two morphs are of the same species) or

reflect interspecific variation. A much more ex-

tensive examination of interpopulation variation

is needed before real conclusions can be drawn.

Parasites

Information is sparse concerning the role of C

chinensis as the final or intermediate host of hel-

FIG. 3. Thf mthilar teeth o/C. chinensis. C = central tooth.

L-I = first laivnl. LIl = I'eeond lateral. M = marginal

(greatly erdargi'li.

minth parasites. Penner (1942) found Massachu-

setts populations to be negative for schistosome

parasites. The snail may serve as a final host to

the normally clam-inhabiting trematode A spido-

gaster conchicola von Baer. The snails are in-

fected by eating embryonated eggs passed in the

feces of other snails (Michelson, 1970; Huehner

& Etges, 1971, 1972, 1977). The snail also serves

as intermediate host to a number of echino-

stome trematodes (U.S. Dept. Agriculture,

1946), including Echinostoma cinetorchis Ando

& Ozaki, a species which has been reported from

humans in Japan, Taiwan and Java (Schmidt &

Roberts, 1977). Angiostrotigylus cantonensis

(Chen), a nematode which infects vertebrate

nervous systems, may use C. chinensis as an in-

termediate host (Chang et al., 1968). This

parasite has been reported from Hawaii and

Costa Rica (Schmidt & Roberts, 1977).

Summary

The introduced Asian snail, C. chinensis. does

well in cool-temperate to warm-temperate cli-

mates in permanent ponds, lakes and the slow

parts of rivers with mud, silt or sand substrates.

Available data indicate that the snail requires a

minimum of 5ppm calcium in its habitat and is

excluded from very soft waters. This species

should be considered a well-established part of

the North American fauna, especially in the

northeastern U.S.

Some basic studies on the natural history and

ecology of this species have been undertaken in

Canada but more studies should be done to de-

termine what effect populations of C. chinensis

are having on the native fauna.

More studies need to be undertaken on the

parasites carried by C chinensis in North

America to determine its potential as a second-

ary host to both native and introduced Asian

parasites.

Shell growth is allometric, the snails tending

to become relatively more elongate with age.

The embryonic periostracal hairs are stiff and

hooked and may serve to stabilize the young

snails within the uterus. The shell growth pat-

tern as well as details of the radula structure dif-

fer between the "typical" C. chinensis and the

japonicus-morph. Whether or not these should

Vol. 96(3)

July 16, 1982

THE NAUTILUS 95

be considered to be intraspecific individual or in-

terspecific differences awaits more extensive

analysis.

ACKNOWLEDGMENTS

Data collection was supported by a research

contract from the Department of Environmen-

tal Protection, State of Connecticut. My thanks

to L. Penner for loan of shells from the Muddy

River.

LITERATURE CITED

Abbott, R. T. 19.50. Snail invaders. Ndtiirnl History

59:80-8.5.

Barnhart, M. C. 1978. Three intmiluced gastropods in Iowa.

The Nautilus 92:106-107.

Barr, A. J., J. H. Goodnight, J. P. Sail, W. H. Blair, C. M.

Chilko. 1979. SAS Users Guide. 1979 ed., SAS Institute,

Inc., Raleigh, N.C., 495 p.

Branson, B. A. 1977. The Chinese apple snail, Cipatigopnlu-

dina chinensis. on Orcas Island. Washington. The Nauti-

lus 91:76-77.

Bucci, D. A. 1974. Vivipnrus malleatus in Montreal,

Canada. The Nautilus 88:55.

Calow, P. 1975. The feeding strategies of two freshwater

gastropods, Ancylus fluviatilis Mull, and Planorhis con-

tortus Linn. (Pulmonata), in terms of ingestion rates and

absorption efficiencies. Oecologia (Berl.) 20:33-49.

Chang. P. -K.. J. H. Cross, Jr. & S.S.S. Chen. 1968. Aquatic

snails as intermediate hosts tor Angiostrongylus cantonen-

sis on Taiwan. Jour. Parasitol. 54:182-183.

Clarke, A. H. 1973. The freshwater molluscs of the Canadian

Interior Basin. Malacologia 13:1-509.

Clarke, A. H. 1978. The Asian apple snail, Cipangopaludina

chinensis (Viviparidae) in (.)neida Lake, New York. The

Nautilus 92:134.

Clench, W. J. 1962. A catalogue of the Viviparidae of North

America with notes on the distribution of Vivipariis

georgianus Lea. Occas. Papers on Mollu.tks. Mus. Comp.

Zool., Harvard Univ. 2(27):261-287.

Clench, W. J. & S. L. H. Fuller. 1965. The genus Viviparus

(Viviparidae) in North America. Occas. Papers on Mol-

lusks. Mus. Comp. Zool. Harvard Univ. 2(32):385-412.

Crabb, E. D. 1929. Egg laying and birth of young in three

species of Viviparidae. The Nautilus 42:125-129.

Dundee, D. S. 1974. Catalog of introduced mollusks of East-

ern North America (North of Me.xico). Sterkiana 55:1-36.

Gould, S. J. 1966. Allometry and size in ontogeny and phy-

logeny. Bio. Rev. 41:587-640.

Gould. S. J. 1971. Geometric similarity in allometric growth:

a contribution to the problem of scaling in the evolution of

size. American Naturalist 105:113-136.

Hanna, G. D. 1966. Introduced mollusks of western North

America. Occ. Papers Calif. Acad. Sci. 48:1-108.

Harman, W. N. & C. 0. Berg. 1971. The freshwater snails of

central New York. Neiv York State Agric. Exp. Sta.

Search: Entomology 1(4): 1-68.

Huehner, M. K. & F. S. Etges. 1971. A new gastropod host

(or Aspidogaxter conchicola. Jour. Parasitol. 57:1255.

Huehner, M. K. & F. S. Etges. 1972. Experimental trans-

mission of A.yjidogaster conrhicola von Baer, 1827. Jour.

Parasitol. 58:109.

Huehner, M. K. & F. S. Etges. 1977. The life cycle and devel-

opment ofAspidngaster conchicola in the snails, Viviparus

malleatus and Goniobasis livescens. Jour. Parasitol.

63:669-674.

.lohnson, C. W, 1915. Viviparus malleatus Reeve in Massa-

chusetts. The Nautilus 19:35.

Johnson, C. W. 1916. Viviparus malleatus Reeve. The

Nautilus 30:48.

.lohnson, C. W. 1918. Viviparus malleatus and contectoides

in Massachusetts. The Nautilus 31:107-108.

Jokinen, E. 1978. Habitats of two freshwater limpets {Fer-

ri.<:sia: Ancylidae) from New England. The Nautilus

92:156-160.

Jokinen, E. H. (In review). Allometric growth of planorbid

snails.

Michelson, E. H. 1970. Aspidogaster conchicola from fresh-

water gastropods in the United States. Jour. Parasitol.

56:709-712.

Penner. L. R. 1942. Studies on dermatitis-producing shisto-

somes in eastern Massachusetts, with emphasis on the

status oi Scihistosojnatium pathloc.opticum Tanabe, 1923.

Jour. Parasitol. 28:103-116.

Perron, F. and T. Probert. 1973. Viviparus malleatus in

New Hampshire. The Nautilus 87:90.

Plinski, M.. W. Lawacz, A. Stanczykowska & E. Magnin.

1978. Etude quantitative et qualitative de la nourriture

des Viviparus ?NaWea/Ms (Reeve) (Gastropoda, Prosobran-

chia) dans deux lacs de la region de Montreal. Canadian

Jour Zool. 56:272-279.

Schmeck, E. H. 1942. Viviparus malleatus in Niagara River.

The Nautilus 55:102-103.

Schmidt, G. D. and L. S. Roberts. 1977. Foundations of

Parasitology. C. V. Mosby Co., St. Louis., 604 pp.

Sokal, R. R. and F. J. Rohlf. 1969. Biometry. W. H. Freeman

& Co.. San Francisco, 776 pp.

Stanczykowska, A., E. Magnin and A. Dumouchel. 1971.

Etude de trois populations de Viviparus malleatus (Reeve)

(Gastropoda. Prosobranchia) de la region de Montreal. I.

Croissance, fecondite, biomasse et production annuelle.

Canadian Jour. Zool. 49:1431-1441.

Stanczykowska, A. M., M. Plinski and E. Magnin. 1972.

Etude de trois populations de Viviparus malleatus (Reeve)

(Gastropoda, Prosobranchia) de la region de Montreal. II.

Etude qualitative et quantitative de la nourriture. Cana-

dian Jour Zool. 50:1617-1624.

Stearns, R. E. C. 1901. Japanese Vivipara in California. Thv

Nautilus 15:91.

Teskey, M. C. 1954. The mollusks of Brown County. Wiscon-

sin. The Nautilus 68:24-28.

U.S. Department of Agriculture. 1946. Index-Catalogue of

Medical and Veterinary Zoology. Part 7. Hosts.

White. J. F. and S. J. Gould. 1965. Interpretation of the coef-

ficient in the allometric equation. American Naturalist

99:5-18.

96 THE NAUTILUS

July 16, 1982

Vol. 96(3)

FOR SALE

A matched pair of blond wood shell cabinets, each

42" wide, 25" deep, and 40" high, with a 3'/z" deep

display case on top, covered by 'A" plate glass.

Each cabinet contains 24 drawers .3" deep. Price

for pair $9.SO.OO F.O.B. Mobile.

One shell dredge, 12" wide, 7" high (with sloped lip

at mouth), 18" long; complete with break-aloose

steel rod bridle and 1" galvanized steel lead chain,

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hot-dipped galvanized after fabrication, covered

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20 lbs. Price $125.00 F.O.B. Mobile. Contact H. I.

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3(5fi05. I'hone (205) 471-1219.

ERRATA

Nautilus, vol. 94 (1) J. K. Buttner and R. C.

Heidinger, "Seasonal Variations - Corbi-

cula - Illinois Fish Pond." Page 9, Table 2,

bottom line, for 10"" read 10-\ Page 10, left

column near bottom, for P>0.01, read

P<0.01. The authors express their regrets.

COMMENTS ON THE EASTERN PACIFIC TRIVIA RITTERI

(GASTROPODA: TRIVIIDAE)

Hans Bertsch

Instituto de Investigaciones Oceanologicas

Universidad Autonoma de Baja California

Ensenada, B.C., Mexico

Barbara Myers

and Department of Marine Invertebrates,

Natural History Museum

P.O. Box 1390

San Diego, California 92112

Although Cate (1979) recently surveyed all

known species of Triviidae the information he

presented on Trivia ritteri Raymond, 1903, was

only a condensation of the original description.

Not only were no new data presented, but one of

the original collecting sites of T. ritteri (Cortez

Bank, west of San Diego) was omitted. The find-

ing of two specimens of T. ritteri from deep

water off San Diego, California, occasioned a

search for additional information.

References, Synonymy and Published Records:

Triiw ritteri Raymond, 1903: 85-86; Catalina Island

(60 fms); Monterey; Cortez Bank (54 fms); Catalina Island

near Avalon (40 fms); off San Pedro (50 fms); Lowe, 1904:

18-20; C:i(alina Island; Dall, 1921: 140; Oldroyd, 1927:

238-239; Grant & Gale, 1931; 44, 754; Upper Pliocene,

Los AiiKfl's; Keen, 1937: 47; Smith & Emer.son, 1955:

99, 101; Ficny, 1956: 151, 154; Anacapa Passage, ,34°00'

38-20" N; ll;»'30' 35-1.3" W (26-27 fms); Anacapa Pas-

sage, 34°00' IC" N; W.i'^-zr :j5-05" W (29-31.5 fms); 2 1/2

miles N. of W. end of Anacapa Island, 34°03' 05-45" N;

119°26' 02" to 25' 28" W (46-58 fms); Abbott, 1974: 149.

fig. 1637

Puiiula ritteri (Raymond). Burch, 1945: 27-29; 42; off

Redondo Beach (25 fms), White's Landing, Catalina

Island (40 fms); Pleistocene of Timm's Point, San Pedro.

Decoriatrivin ritteri (Raymond). Cate, 1979: 11, 98; figs.

94 and 94a.

Geographic and. Bathyynetric Distributions:

Trivia ritteri has been recorded from the

upper Pliocene and the Pleistocene of Los

Angeles, and in the Recent from Monterey to

San Pedro, Catalina Island, Anacapa Passage

and Island, and Cortez Bank. Our knowledge of

the distribution of Trivia ritteri has been based

on just a few specimens.

In the collections of California Academy of

Sciences (CAS), Los Angeles County Natural

History Museum (LACM), and San Diego Nat-

ural History Museum (SDNHM) are numerous

Vol. 96(3)

July 16, 1982

THE NAUTILUS 97

lots of Trivia ritteri. An analysis of the accom-

panying data of those specimens gives a better

understanding of the distribution of T. ritteri

(Table 1). Specimens in the LACM collection ex-

tend the range of T. ritteri southward more

than 400 miles (over 650 km) to midway down

the Baja California peninsula (27° N). Trivia

ritteri is endemic to the California Province

(sensu Valentine, 1966), with a few northern

records to Monterey (36° N).

Trivia ritteri is usually found deeper than 50

m (Figure 1). More than two-thirds of the 113

specimens analyzed for bathymetric occurrence

were found between 51-90 meters deep. Trivia

ritteri has a predominantly insular distribution:

the majority of specimens have been collected

from the offshore islands of southern California

and northern Baja California. Mainland collect-

ing records are either from several miles off-

shore or from deep trenches (such as La Jolla)

which rapidly drop off close to the shoreline.

TABLE 1. Recent records o/ Trivia ritteri with their depth

ranges.

Location Specimens Depth Range

Central California

Monterey Bay area

Southern California, mainland

Los Angeles area, offshore «12 mi.)

San Diego, La Jolla trench

Southern California , Channel Islands

San Miguel, Santa Rosa, Santa Cruz

and Ana cap a Islands

San Nicholas, Santa Barbara and

Santa Catalina Islands

San Clement e Island

Baja California. Mexico

West of Isla Cedros

South of Isla Cedros

Off Thurloe Head (27° 36" 50" N;

lU* 50' 50" W)

Off San Pablo Pt . (27° 12' 45" N;

114° 31' 45" W)

FIG. 1. Bathymetric distribution o/Trivia ritteri, bitsed on

113 spechnens. Vertical numbers are alternating 10 m

depths. The mid-point oftlie collection information was used

for specimen, lots with a range of depth.

TABLE 2. Shell size and teeth counts o/Trivia ritteri. Shells

in collection of the San Diego Museum of Natural History.

Number of Apertural Denticles

Inner Lip Outer Lip

98 THE NAUTILUS

July 16, 1982

Vol. 96(3)

Shell Maiyhology:

Raymond (1903) described the shell characters

of Trivia ritteri fairly comprehensively; the

first published illustration of the species appears

to be Abbott, 1974: fig. 1637. Among the mate-

rial we examined (Table 2) were several imma-

ture specimens, a bulloid juvenile form com-

pletely lacking apertural denticles and dorsal

ribbing (Figs. 2 and 3) and a subadult (Figs. 4

and 5) with teeth, the outer lip still thin but hav-

ing begun folding over the aperture, and the

dorsal surface nearly smooth. The animal begins

to develop the adult shell morphology (Figs. 6

and 7) when it reaches 6-8 mm. The British

Trivia arctica reaches the adult stage in 6-8

months (Lebour, 1933).

Radula:

The radula of Trivia ritteri is illustrated here

for the first time (Fig. 8). The radula we exa-

mined has 22 rows of fully formed teeth, with 2

scythe-shaped marginal teeth and a single large

admedian tooth on each side of the median tooth

(cf. Schilder, 1936, for terminology). The adme-

dian tooth has a single denticle on the inner side

of the erect shaft, and 3 or 4 small denticles on

its outer side. The median tooth has a major

cusp with 6-8 small accessory denticles on each

side.

ACKNOWLEDGMENTS

We are grateful to those who graciously

loaned us specimens or gave us data from collec-

tions in their care: Mr. Loyal J. Bibbey; Dr.

Barry Roth, California Academy of Sciences;

and especially Mr. Gale Sphon and Dr. James H.

McLean, Los Angeles County Natural History

Museum. We thank Anthony D'Attilio, San

Diego Natiiral History Museum, for the draw-

ings.

LITERATURE CITED

Abbott, R. T. 1974, American SciishMa. Second Edition.

Van Nostrand Reinhold Company, New York. 663 pp.;

24 pis.; numerous text figs.

FIGS. 2-7, Sh:lt <;/'Tri\na ritteri, dm-xat mid venlrai vinrs.

Photo(/rnj,hs hy B. Myers. 2-3, Juvenile hulla-form; . 4-5,

SuhrtdnlU<: 6-7. Ad^ill.-i.

FIG. 8, Radula of Trivia ritteri. Drawimj hy Anthony

D'Attilio.

Vol. 96(3)

July 16, 1982

THE NAUTILUS 99

Berry. S. S. 19,56. Mollusca dredged by the Orea off the

Santa Barbara Islands, California, in 1951. Jirur. Wash-

uigtoH Acml. Sri. 46(5):150-15T; 9 text figs. (5 June 1956).

Burch, J. Q. 1945. Distributional list of the west American

marine mollusks from San Diego to the Polar Sea. Pror.

Conchohyical Club So. California 53:27-29; 42 pis.

(October 1945).

Gate, C. N. 1979. A review of the Triviidae (Mollusca:

Gastropoda). San Diego Soc. Nal. Hi.-it. Mem. 10:1-126;

41 pis. (20 June 1979).

Dall, W. H. 1921. Summary of the marine shellbearing

mollusks of the northwest coast of America, from San

Diego, California, to the Polar Sea, mostly contained in

the collection of the United States National Museum,

with illustrations of hitherto unfigured species. Bull. U.S.

Nat. Mus. 112:1-217; 22 pis. (24 February 1921).

Grant, S., IV and H. R. Gale. 1931. Catalogue of the marine

Pliocene and Pleistocene Mollusca of California and adja-

cent regions. San Diego Soc. Nat. Hist. Mem. 1:1-1036;

32 pis. (3 November 1931).

Keen, A. M. 1937. An abridged check list and bibliography

of West North American Mollusca. Stanford Univ. Press.

Stanford, California. 87 pp.

Lebour, M. V. 1933. The British species of Triina: T. artica

and T. monacha. Jour. Marine Biol. Assoc. U.K. 18(2):

477-484; 1 pi. (January 1933).

Lowe, H. N. 1904. k dredging trip to Santa Catalina I.sland.

The Nautilus 18(2):18-20. (June 1904).

t)ldroyd, I. S. 1927. The marine shells of the west coos* of

North America. Stanford Univ. Press, Stanford, Cali-

fornia, Vol. II (2);l-304; pis. 30-72.

Riiymond, W. J. 1903. A new Californian Trivia. The

Nautilus 17(8):85-86. (December 1903).

Schilder. F. A. 1936. Anatomical characters of the Cypraea-

cea which confirm the conchological classification. Pruc.

Malar. Soc. London 22(2):75-112; pis. 11 & 12. (15 July

1936).

Smith, A. G. and W. K. Emerson. 1955. William James

Raymond, 1865-1947. The Nautilus 68(3):98-102.

(January 1955).

Valentine, J. W. 1966. Numerical analysis of marine mollus-

can ranges on the e.xtratropical northeastern Pacific

shelf. Limnology and Oceanography 11(2):198-211; 7 text

figs. (April 1966).

AN OVERLOOKED BUSYCON WHELK (MELONGENIDAE)

FROM THE EASTERN UNITED STATES

Bretton W. Kent

Department of Zoology

University of Maryland

College Park, MD 20742

ABSTRACT

Busycon laeostomum new species, a large living sinistral Busycon u>helk is

described from the eastern United States. This new species is compared with B.

carica (Gmelin, 1 791) and B. contrarium (Conrad, 18U0) and its subgeneric affini-

ties are discussed.

A large, sinistral Busycon is occasionally col-

lected offshore along the Atlantic coast of the

United States between southern New Jersey

and northern Virginia. This whelk frequently

has a pure white shell and has been referred to

as either a sinistral form of B. carica (Gmelin,

1791) or as B. contrarium (Conrad, 1840)

(Abbott, 1974, p. 222). Here I show that this

whelk is distinct from sympatric B. carica and

B. contrarium, and describe this new species.

100 THE NAUTILUS

July 16, 1982

Vol. 96(3)

FIGS. 1-.5. Busycon laeostomum tl sp. 1, 258 mm (Pope coUeclion); 2, 213 mm (USNM no. 679720), 3, 222 mm (with soft tissues

and operculum; Harasewych collection), 4, 221, mm (USNM no. 806850). and 5, 221 mm (siphonal canal broken; USNM wo.

806850).

Vol. 96(3)

July 16, 1982

THE NAUTILUS 101

Busycon laeostomum, new species

Figs. 1-5, 7, 10

Shell Description - Shell sinistral, solid, sub-

pyriform, inflated; shoulder rounded, tubercu-

late; spire low, subconical; spiral cords on spire

weak or absent; aperture broad, elliptical,

longer than siphonal canal; anal ridge on

parietal wall narrow, weak; siphonal canal open,

somewhat flaring; columella broad, strongly

bowed; spiral cords on body whorl restricted to

dorsum of siphonal canal, weak; shell exterior

usually chalky, white or pale pinkish-orange, oc-

casionally with indistinct slightly darker suffu-

sions; periostracum minutely ciliated; aperture

white or light pinkish-orange; apertural lirae ab-

sent or rarely weak.

Description of Animal - Head and foot black;

operculum corneus, elliptical, lingulate, nucleus

apical; radula with 3 teeth per row, 5-6 cusps on

central teeth.

Material Examined - Holotype - Length =

245 mm (U.S. National Museum no. 806849);

trapped in 10-15 m of water off Stone Harbor,

New Jersey, January 1979. Paratypes - Length

= 184 mm (USNM no. 678850); dredged 8-10 km

off Avalon, New Jersey, July 1967. Length =

213, 176 and 138 mm (USNM no. 679720);

dredged off Ocean City, Maryland, 1969.

Length = 224 and 221 mm (USNM no. 806850)

and 222 and 220 mm (M. G. Harasewych collec-

tion); trapped off Stone Harbor, New Jersey in

10-15 m of water, January 1979. Length = 220

mm (USNM no. 806851); off Avalon, New

Jersey, 1968. Length = 258 mm (Theresa R.

Pope collection) and 83 mm (author's collection);

collected as beach shells at Tom's Cove,

Assateague Island, Virginia, April 1977. Length

= 163 mm (Geerat J. Vermeij collection); off

Ocean City, Maryland, 1977.

Type Locality - Stone Harbor, New Jersey;

in 10-15 m of water.

Range - Southern New Jersey to northern

Virginia; offshore.

Etymology - laios (Gk.) = left, stoma (Gk.) =

mouth.

Remarks - B. laeostomum is sinistral and has

a ciliated periostracum hke B. contrarium. but

otherwise more closelv resembles B. carica in

general shell shape (Figs. 6-11, Table 1). B. con-

trarium is more slender and has a more angular

shoulder than B. laeostomum, although tuber-

cles along the shoulder make the latter differ-

ence somewhat difficult to observe in photo-

graphs (e.g. - Figs. 4 and 10). B. contrarium and

B. laeostomum also differ with respect to the

development of these tubercles, the nature of

the external spiral cords and apertural lirae, the

shape of the columella, and the color of the shell.

■WliileS. laeostomum has only rounded tubercles

along the shoulder, in B. contrarium these

tubercles are frequently elongated into promi-

nent spines. B. contrarium also has well-

developed spiral cords and apertural lirae which

become obsolete in later whorls. In B. laeosto-

mum these cords and lirae are very weakly

developed or more commonly completely ab-

sent, even in the early whorls (e.g. -compare

Figs. 9 and 10). The columella of 5. contrarium.

is slender and relatively straight compared to

broad, strongly bowed columella of B. laeo-

stomum. Finally, the dark brown axial streaks

that usually occur on the early whorls of B. con-

trarium (Fig. 9) do not occur in B. laeostomum.

In B. laeostomum 62% of the types (8 of 13) had

a pure white shell; the remaining shells were a

TABLE 1. Comparison of northern Busycon contrarium

and B. carica with B. laeostomum, with respect to spire

angle, the ratio of siphonal canal length to aperture length,

aperture shape ( = aperture uidth divided by aperture length)

and canal shape (= canal uridth halfway along canal divided

by canal length). The values listed are means with ranges in-

dicated in parentheses. Shells with badly broken lips or

siphonal canals were not measured. Differences were tested

for signif>.cance with a one-tailed Mann-Whitney U test and

the level of significance is indicated by asterisks (i.e.-' in-

dicates p <0.05. ** indicates p <0.01. and **' indicates p

<ri.ooii.

contrarium laeostomum carica

."^pire

Angle

Canal: Aperture

Ratio

Aperture

Shape

Canal

Shape

N

.Size Range

94°** 105° 81°"*

(85-121°) (92-126°) (75-90°)

1.10*** 0.77 0.82

(0.87-1.48) (0.66-0.93) (0.72-0.93)

0.60 0.60 0.58

(0.57-0.64) (0.57-0.64) (0.51-0.63)

0.19*" 0.33 0.29

(0.14-0.26) (0.24-0.41) (0.19-0.35)

15 10 12

75-224 mm 83-258 mm 93-219 mm

102 THE NAUTILUS

July 16, 1982

Vol. 96(3)

I

7

\

«<

I

I

0

FIGS. 6-11. Comparison of typical adults (upper row) and juveniles (lower row) o/Busycon contrarium (Fig. 6 and 91, B.

laeostomum (Fig. 7 and 10) and B. carica (Fig. 8 and 11). 6, 22J, mm, 7, 2^5 mm (holotype; USNM no. SOtiHi'J). 8, 191 mm, 9, 88

mm, 10, S3 mm (author's collection), and 11, 93 mm.

pale pinkish-orange with slightly darker suffu-

sions. Pure white shells (i.e. -no axial streaks

even on the early whorls) are known to occur in

B. contrarium, but are very rare (HoUister

1958). I have examined four white B. con-

trarium (USNM nos. 36298. 414722. 609947.

Vol. 96(3)

July 16, 1982

THE NAUTILUS 103

and 631770). Except for the white shells these

B. contrarium are indistingiiishable from other

B. contrarium. This suggests that B. laeosto-

mum is not merely a white-shelled morph of B.

contrarium. Further, B. laeostom,um is probably

not just a northern form of B. contrarium. The

two species are sympatric throughout the range

of B. laeostomum. rather than having con-

tiguous or disjunct ranges as would be expected

of geographic morphs.

Although B. laeostomum and B. carica are

similar in shape (Figs. 7, 8, 10 and 11, Table 1),

several lines of evidence suggest that they are

not merely sinistral and dextral morphs of the

same species. B. laeostomum and B. carica dif-

fer, not only with respect to handedness and the

nature of the periostracum, but also in spire

angle (compare spires in figs. 7, 8, 10 and 11;

Table 1), the shape of the anal ridge, and shell

color. The anal ridge in B. carica is a swollen

prominance (Hollister 1958) that is very dif-

ferent from the weak ridge in B. laeostomum. B.

carica also frequently has a bright reddish-

orange aperture (Hollister 1958), but this has

not been observed in B. laeostomum. The white

shells which commonly occur in B. laeostomum

have not been reported in B. carica.

If B. laeostomum, and B. carica were sinistral

and dextral morphs of the same species they

should have similar geographic and bathymetric

ranges. The geographic ranges of the two

species are very different; B. carica occurs from

Massachusetts to northern Florida (Abbott

1974), while B. laeostomum has a restricted

range between southern New Jersey and north-

ern Virginia. The bathymetric ranges are also

different. B. carica occurs both intertidally and

subtidally in areas where it is sympatric with B.

laeostomum (pers. observ., M. G. Harasewych,

pers. comm.), while all of the live-collected B.

laeostomum were subtidal.

Finally, differences in handedness may pre-

clude copulation between B. laeostomum and B.

carica. There is no direct evidence that sinistral

and dextral morphs are unable to copulate, but

as Pulley (1959) has argued, even slight diffi-

culty in copulation would reduce gene flow be-

tween the two forms. If copulation between the

two morphs were possible, egg capsules should

occasionally be found containing both sinistral

and dextral individuals. Gill (1867) examined

over 500 B. carica egg capsules, but found only

dextral individuals.

The presence of sinistral coiling and a ciliated

periostracum strongly suggests that B. laeo-

stomum is much more closely allied to B. con-

trarium than to B. carica. Consequently, B.

laeostoynum should be placed in the subgenus

Sinistrofulgur Hollister 1958 with B. con-

trarium and B. perversum Linne 1758.