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ANUARY 25. 1982 



1^«N ()()28-i:i44 

Vol. 96 

No. 1 

A quarterly 

devoted to 

malacology and 

the interests of 


Founded 1889 by Henn' A. Pilsbry. Continued by H. Burrington Baker. 
Editor-in-Chief; R. Tucker Abbott 



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 


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 

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 



Volume 96, number 1 - January 25, 1982 

ISSN 0028-1344 


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 


January 25, 1982 

Vol. 96(1) 


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,, 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. 

.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 
.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 


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, 

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 

I'ln' I'irsI a>vrhfdogiral wtirk liij mi .■\mmcaii 
jiuhHuhed in A mtn'icit. 


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, 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 



Joseph Rosewater 

Department of Invertebrate Zoology 

National Museum of Natural History 

Washington, D.C. 20560 


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- 


January 25, 1982 

Vol. 96(1) 

Vol. 96(1) 

January 25, 1982 


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 

No. Specimens 





Range of 






Range of 






Range of 






Range of 


Left Valve 












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 


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). 


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. 


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. 



Francesco B. Trama 

Department of Biological Sciences 

Rutgers LTniversity 

New Brunswick, New Jersey 


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 


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. 


January 25, 1982 

Vol. 96(1) 

specimens or empty shells were found in those 

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 


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 

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. 


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 

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. 


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 



Philippe Bouchet and Francoise Danrigal 

Museum National d'Histoire Naturelle 

55, Rue de Buffon 

75 Paris (5°), France 


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 

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 

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. 


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 

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 


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 

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. 


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 

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 

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 


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 


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. 

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. 


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 


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. 


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. 


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. 


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. 


isseli Nevill & Nevill, 1875 (Marginella) 
Nevill & Nevill 1875: 95 

Fig. 57 


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 

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. 


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. 


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 


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 


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 


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 


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 /, 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 



January 25, 1982 

Vol. 96(1) 

Vol. 96(1) 

January 25, 1982 



January 25, 1982 

Vol. 96(1) 

Vol. 96(1) 

January 25, 1982 



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 {\ 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. 






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. 


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 



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. 

' )-^**&::: 



,1 ■■ ' ap*"; 

102, Bursatella savignyana Audouin. 130 mm. 

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


103, Octopus horridus Orhigny. mantle length 30 mm. 


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. 


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- 

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: 

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., 

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. 

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. 

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 



Clement L. Counts, III and Robert S. Prezant 

College of Marine Studies 

University of Delaware 
Lewes, DE 19958 U.S.A. 


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 

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. 


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 


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 

Mounted specimens were coated with a thin 
layer of gold in a Denton Vacuum Evaporator 
and examined in a Philips PSEM 501 SEM. 


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- 

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 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 


(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 


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 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 


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). 


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 

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 


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. 


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. 


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 

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. 


Paul W. Parmalee and Walter E. Klippel 

Department of Anthropology 
University of Tennessee 
Knoxville, TN 37916 ' 


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 


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. 


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. 


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. 


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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Specimen Shells 

Offering microscopic and miniature (to '/t inch) shells 
from the Florida Keys, vnth accurate locality data Also 
unsorted grunge: write for list. 

Margaret Teskey 

P.O. Box 273 

Big Pine Key. Fl. 3301,3 


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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 

Recent Deaths 68 Meetings and Shell Shows 68 


April 21, 1982 

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Vol. 96(2) 

April 21, 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. 

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 

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 


Vol. 96(2) 

Terrence J. Frest and R. Sanders Rhodes II 

Department of Geology 

University of Iowa 

Iowa City, lA 52242 


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 

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 


'■■:'■• 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 


(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) 
Ste notrema 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 icodiscu s parallelus (Say, 1821) 
Vertigo tndentata Wolf. 18 70 
Vallonia parvula Sterki, 189J 
Zcnitoides arboreus (Say, 18161 
Ret inei la indentata (Say, 1823) 


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 


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











District of Columbia 

















New Jersey 



New York 











South Carolina 







West Virginia 


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. 


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): 

1946. Land Mollusca of North America (North 

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

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. 


April 21, 1982 

Vol. 96(2) 


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 



Js" '-■/ .''^' .* ■(! 

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

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

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



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- 


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 


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. 



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- 





37 mm 

19 mm (Figs. 1 and 2) 


31 mm 

14 mm (Figs. 3 and 4) 

(Hammes' collection) 

33 mm 

17mm (Figs. .5 and 6) 

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 


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. 


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 

Locality: This specimen was live-collected 
near Los Zurrones, west Panama, by Royce E. 
Hubert, in 1979. Mr. William E. Old, Jr., first 


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 

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 

Detailed comparison of internal and external 
(shell) anatomy will be necessary to determine 
the relationship between Phenacovola 
brevirostris and P. lenoreae Cardin & Walls, 

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 Ber tsch & Bibbey sp. nov. P 


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 

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 


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 

1976b. Five new species on Ovulidae (Mollusca: 

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


Jesus Ortea 

Department of Zoology, Oviedo University, Spain 


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 

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- 


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: 

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 

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 


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- 


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). 


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. 


I wish to thank Dr. Terrence Gosliner and the 
editor, R. Tucker Abbott, for assistance with the 


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. 

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 

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: 

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 

1970. Opisthobranchs from Cura(;ao and faunis- 

tically related regions. Studies on the Fauna of Curasao 
and other Caribbean islands. Stud. Fauna Curagcui 

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. 


SEA SHELLS p. o. Box 83 

GUn Ellen, CA 95442 



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 



Ralph W. Taylor and Beverly D. Spurlock 

Department of Biological Sciences 

Marshall University 
Huntington, West Virginia 25701 


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. 


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 


April 21, 1982 

Vol. 96(2) 



W. VA. 

FIG. 1. Map of the upper Ohio River showing the approx- 
imate location of the archeological sites bearing freshwater 

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 

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 


Exca\-at ion 

Habi tat 1 un 

Species Identified 

Globe Hill 





Neale's Landing 





Lewis-Old Town 

Taylor and 


197 9-80 



Rolf Lee 

Taylor and 


197 9-80 








Composite .Number 



32 species 

Vol. 96(2) 

April 21, 1982 


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. 


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. 


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- 

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. 


April 21, 1982 

Vol. 96(2) 



Richard J. Neves 

Virginia Cooperative Fishery 

Research Unit' 

Department of Fisheries 

and Wildlife Sciences 

Virginia Polytechnic Institute 

and State University 

Blacksburg, Virginia 24061 


Alexander V. Zale 

Florida Cooperative Fishery 

and Wildlife Unit^ 

Newins-Ziegler Hall 

University of Florida 

Gainesville, FL 32611 


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 

'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 


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. 


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- 

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- 

7. Adjacent to state route 613 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 


Station No. 

4 5 6 


Alasmidonta minor {Lea, 1845) 

Lampsilis fasciola (Rafinesque, 1820) 
Medionidus conradicus (Lea, 1834) 
Villosa nebulosa (Conrad, 1834) 
Villosa vanuxemi (Lea, 1838) 


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 






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. 


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 


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 

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 



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

Division of Biology 

Corpus Christi State University 

Corpus Christi, Texas 78411 


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- 


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 


1— Punta d 

bI Morro 
(Punta Delgada) 

A — Punta 

Boquilla de Piedras 


V- Pl 

nta Boca de Lome ~ 

^^x;;;:>=i— Barra Palma Sola 


- Punta Los Munecos 


iPunla Boca Andreal 


-—Punta Delgada 

(Punta Las Literas) 


- Punta Limon 


1 — Punta Villa Rica 





FIG. 1. Mapofth: ii-ealnyiGulfofMexiroshov'tngllu'Piintit 
del Morro-PwnXa r>etg<ida region. Bathymetry contoured in 

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 


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). 


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 


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. 


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 


Species Ecological 
_^ Notation + 

Class Cascropoda 

Subclass Prosobranchla 

Order Archaeogastropoda 

*Dl_ od _ ora caycnensls (Lamarck, 1822) Int 

* Fla8urella barbadensls (Cinclin. 1791) Int 


*Acnaea leucopleura (Cinelln, 1791) int 


Calllostoma euglypCum (A. Adams, 1854) bd 
C. J ububinum (Cinelln, 1791) bd 

Tefjula fasclata (flcrn, 1778) bd 


Astra ea tecta Llghtfoot, 1786 bd 


•Tricolia afflnis 

(C.B, Adams, 1850) 


• 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) 

Vitrtnclla floridana Pllsbry and McGlnty, 1946 
Parviturbq id os IntirruptuB (C.B. Adans, 1850) 

Arch 1 tec tonic n noblllw Rodlng, 1709 
Hejjacyj* hiBulcatuB (Orblgnv, 1842) 
H. cyl lndrlcus {Gmflln, 1791) 

*'-i''£.'^ p" lchellun Stimpaon, 1851 
'C. rvi'-iu'ilt iim Folln, 1867 
't- Ke^iJl"^ (OrblRny. IStl) 
Vcrsfit Idrti^ ~ 

' Pttnloronc hun var Una (Orbignv, 1841) 

CTlChlun IjitoBiin Mt-nke. 182R 


Cerithlopsis emersoni (C.B. Adams. 1838) 


Alaba incerta (Orbigny, 1842) 



•Planaxis lineatus (da Costa. 1778) 

al. tp 

*P. nucleus ^Brugulire. 1789) 



Strombus puf^ills Linnaeus, 1756 


S. ranlnus Gmelin. 1791 



•Epitonlum lamellosum (Lamarck. 1822) 



Janthlna Janthina (Linnaeus. 1758) 


J. globosa (Swainson, 1822) 



Helanella conoidea Kurtz and Stlmpson, 1851 


M. intermedia (Cantraine. 1835) 



*Fossarus orbignyl Fischer. 1864 

al, sab 


*CrepIdula aculeata (Gmelin. 1791) 

C. fornlcata (Linnaeus, 1758) 


*C. maculosa Conrad, 1846 



Pollnlces hepatlcus (Rddlng, 1798) 


P. lacteus CCulldlng, 1834) 


Stigmaulax sulcatus (Born, 1778) 



Cvpraea spurca acicularls Cmelin. 1791 



Cyphoma glbbosum (Linnaeus, 1758) 



Fhallum granulatum Born, 1778 



Cymaclum nicobaricum (Rodlng, 1798) 


C. plleare (Linnaeus, 1758) 



Bursa Eranularls cubanlana (Orbigny. 1842) 


Order Neogastropoda 


*Horula nodulosa (C.B. Adams. 1845) 


•Purpura patula (Linnaeus, 1758) 

Int. tp 

*Thals haemastoma (Conrad. 1837) 



Coralliophlla aberrans (C.B. Adams, 1850) 



Bailya Intricata (Dall. 1884) 


•Plsania tlncta (Conrad, 1846) 



Columbella mercatorla (Linnaeus. 1758) 


*Anachi9 floridana Rehder , 1939 


*A. semlpllcaca Stearns, 1873 


Zafrona Idallna (Ducloe, 1840) 


*Nltldella laevigata (Linnaeus, 1758) 

sbt. al 

N. nltlda (Lamarck, 1822) 


Mltrella ocellata (Gmelin, 1791) 



Busjrcon splratum plaposum (Conrad. 1863) 


Nassarius acutus (Say, 1822) 


N. albus (Say, 1826) 



*Leucozonla nassa (Gmelin, 1791) 


Utlrus infundlbulum (Cmelin. 1791) 



Oliva sayana Ravenel, 1834 


Ollvella cf. mlnuta (Link. 1807) 



Mltra nodulosa (Gmelin. 1791) 


Vexillum sp. 


•Conus mus Hwass , 1792 


Subclass Opisthobranchia 

Order Cephalaspldea 


*BulU striata flrugui^re, 1792 


Order Anaspldea 


•Aplysla cf. braslUana Rang, 1828 

*A. dactvlomela Rang, 1828 

Order Nudibranchla 

*Nudibranch sp. A 

*Nudibranch sp. B 

Subclass Pulmonata 

Order Basommatophora 


*Pedipes mirablis (Muhlfeld. 181'') 

spt, sab 


*Siphonaria pectinata (Linnaeus, 1758) 

Class Bivalvia 

Subclass Pteriomorpha 

Order Arcoida 


*Arca imbricata (Brugulere, 1789) 


A. zebra (Swainson, 1833) 

•Barbatia Candida (Helbling. 1779) 

*B. domingensis (Lamarck, 1819) 

Anadara braslUana (Lamarck. 1819) 

A. transversa (Say, 1822) 


A. sp. 


•Arcopsls adamsi (Dall, 1886) 



Glvcvmeris decussata (Linnaeus. 1758) 

Order Mytllolda 

Vol. 96(2) 

April 21, 1982 


TABLE 1 (continued) 



Notation + 


*LlthophaEa arlstata (Dlllwyn, 1817) 


*BrachldonCes exustus Linnaeus, 1758 

int. sab 

Order Pteriolda 


♦Plnccada ImbrlcaCa Rodlng. 1798 



*IsoRnonion alatus (Ciaelin, 1791) 


•1. blcolor (C.B. Adams. 1845) 


*I. radlatus (Anton, 1839) 



*Cras8ostrea vlrginlca (Gmelln. 1791) 

sbt. tp 


ArRopeccen sp. 


Chlamys ornaca (Lamarck, 1819) 



Spondvlus cf. Icterlcus Reeve. 1856 


Subclass KeCerodonCa 

Order Venerolda 


Polymesoda carollntana (Conrad. 1839) 



Luclna pectlnata (Cmelln. 1791) 



Arctnella cornuta Conrad, 1866 


Chama macerophylla (Cmelln. 1791) 


*Pseudochama radians (Lamarck, 1819) 



Dinocardluro robustutn (LiRhtfoot, 1786) 


Trachvcardium Isocardla (Linnaeus. 1758) 



Chione cancellata (Linnaeus. 1767) 


C. clenchi Pulley, 1952 


Hacrocalllsta maculaEa (Linnaeus, 1758) 


Doslnla cf. elegans Conrad. 1846 



RanRla flexuosa (Conrad. 1839) 



Ma coma sp. 


ScriRllla mlrabllls (Phllippi, 1841) 



Donax sp. 



Semele proflcua (Pulteney. 1799) 


Order Myoida 


*Sphenla antlllensis Dall and Simpson. 1901 


Class Polyplacophora 

Order ChiConlda 


*Tonicia schramml (ShutCleworth, 1856) 


Order Unknown 

Chiton sp. A 


Chiton sp. B 


Class Cephalopoda 

Subclass Coleoidea 

Order Seploldea 


Splrula splrula (Linnaeus, 1758) 


Order Octopoda 


♦Octopus sp. 


•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 



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 

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. 


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 

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 

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. 


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." 


The authors would like to gratefully thank the 
Direccion General de Pesqueras, Departamento 
de Pesca for allowing us to collect and study in 

Vol. 96(2) 

April 21, 1982 



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Twila Bratcher 

8121 Mulholland Terrace 
Hollywood, CA 90046 


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 


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 


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 

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. 


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- 


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; 


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 


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 

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 

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 

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 


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 


AM - Australian Museum 

ANSP- Academy of Natural Science of Phila- 

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, 

MORG-Museu Oceanografico de Rio Grande, 

NM- Natal Museum, South Africa. 

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. 


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. 




Ralph W. Taylor 

Department of Biological Sciences 

Marshall University 
Huntington, West Virginia 25701 


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 


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. 


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- 

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. 


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 


April 21, 1982 

Vol. 96(2) 


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. 


Goodrich, Calvin and H. van der Schalie. 1944. A Revision of 

the Mollusca of Indiana. The Amer. Mid. Nat. 32(2): 

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): 


July 14-17. CO. A. (Conchologists of America). 

Sanibel Island, Florida. 
July 19-23. A.M.U. (American Malacological 

Union. New Orleans, Louisiana. 

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. 


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. 


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). 




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 


Vol. 96(2) 

April 21, 1982 





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 


Michael Ross 

Bureau of Fisheries and Aquatic Resources 

Region 7 District Office 

Cebu City, Philippines 


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 

36. Tatsuo Tanaka Memorial Biological 

population explosions of Drupella with 
widespread coral destruction have yet to be 

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 


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 

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 

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). 


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 


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 

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- 


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 


"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. 


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 


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. 


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 


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. 


April 21, 1982 

Vol. 96(2) 

Vol. 96(2) 

April 21, 1982 


Table 1. Composition of dominant corals on 
Mactan Island study sites 


Tambuli Beach 


ingin Beach 













)a laxea 






'oat I lopora 






'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 


'avi tes 








Aaropora formosa 
A. tenuis 
A. florida 
A. palifera 
A. robusta 
A. humilus 

Aaropora hyaointhus 
A. millipora 








Symphy I Ha 





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 


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 


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 

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 


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. 


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. 


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Dorothea S. Franzen 

Illinois Wesleyan University 
Bloomington, Illinois 61701 

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 


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 
^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 


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. 


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 

2. Field No. ALM 1058 (DSF 412); 30 ,Iuly, 1970; 2.5 mi N 
of Mt. Dora, Union County, New Mexico. 


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 

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, 

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 

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 

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 


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. 

No. of 




Width of 

H. Ap 


W. Ap 


W. Ap 

station #1 







H. Shell 

W. Shell 

H. Ap 

Union Co. , NH 

3 1/4 

12.6 mm 

5.45 mm 


b . mm 

4.05 mm 




30 June, 1969 

3 1/4 









3 1/4 









^Range (23 shells) 


3/4 - 3 


6.0- 12.6 

3.3 - 6.3 

.432 - 


3.9 - 8 


2.7 - 5 


.476 - 


.706 - 


.606 - 











Station #4 

Dona Ana Co. , NH 

3 1/4 

12.3 mm 

6.6 mm 


7.0 mm 

4. 7 mm 




3 May, 1966 

3 1/2 









3 1/4 









Range (29 shells) 


- 3 1/2 


3.7 - 6.6 

.514 - 





.541 - 




.606 - 











station #7 

Dtero Co. , NM 

3 1/2 

9.7 mm 

5.2 mm 


5.65 mm 

3. 6 mm 




11 October, 1969 

3 1/2 






.64 5 



3 3/4 









^ange (39 shells) 


1/2 - 3 


4.15 - 9.7 

2.55 - 5.55 

.530 - 


2.9 - 6 


2.0 - 4 


.561 - 


.692 - 


.571 - 











Station #9 

;l Paso Co. , TX 

3 1/4 

12. 1 mm 

6. 3 mm 


6. 75 mm 

4.2 mm 




8 Feb. ; 7 May, 1966 

3 1/2 









3 1/4 








.614 ■ 

iange (58 shells) 


1/2 - 3 


4.2 - 12.1 


.404 - 


2.7 - 6 


1.95 - 4.2 

.528 - 


.637 - 


.573 - 











Station #13 

■Jchiltree Co. , TX 

3 1/2 

11.40 mm 

5.6 mm 


6.85 mm 

4. 55 mm 




23 August, 1968 

3 3/4 









3 1/2 









a<ange (51 shells) 


- 3 3/4 

6.25 - 11.4 

3.65 - 6.10 

.491 - 

64 3 

4.0 - 6 


2.4 - 4 


.557 - 




.60 - 











station #16 

-*'ecos Co . , TX 

3 1/4 

9. 5 mm 

5. 1 mm 


5.8 mm 

3.65 mm 




16 April, 1973 



















arvie (18 shells) 


1/2 - 3 



2.70 - 5.10 

.492 - 


2.85 - 5.8 

1.95 - 3.65 



.629 - 


.60 - 











1 :tation »18 

1 (arris Co. , TX 


7.2 mm 

4-6 mm 


5.6 mm 

3.4 mm 




14 July, 1972 



4. 3 
















;anqe (15 shells) 


1/2 - 3 


2.8 - 4.6 

.542 - 




2.1 - 3 


.661 - 


.657 - 













tjtion #19 

all Co. , OK 


9.8 mm 

6. mm 


7.1 mm 

4. 5 mm 




July, 1968 



















(22 shells) 


1/2 - 3 

5.4 - 9.8 

3.5 - 6.0 

.582 - 


4.0 - 7 


3.0 - 4 


.683 - 


.714 - 


.612 - 


1 Station #19 
larshall Co. , OK 

3 1/4 

9.2 mm 

5. 6 mm 


6. 3 mm 

4.2 mm 




17 July, 1972 



















:ange (18 shells) 


1/2 - 3 



4.0 - 5.6 

.600 - 




3.0 - 4 


.670 - 


.70 - . 


.60 - . 











;tatlon #20 

leaver Co. , OK 

3 1/4 

9.6 mm 

5.0 mm 


5.6 mm 

3.6 mm 




1 August, 1970 

3 1/2 
3 1/2 










ange (19 shells) 


- 3 1/2 

5.J5- 9.6 

3.15 - 5.0 

.493 - 


3.6- 5. 


2.3 - 3. 


.536 - 













tat ion #21 

orton Co. , KS 

3 1/3 

10.5 mm 

5, 7 mm 


6.5 mm 

4. 3 mm 





3 1/4 









3 1/4 









ange (5 shells) 


- 3 1/3 

8.7 - 10.5 

5.0 - 5.7 

.542 - 




3.8- 4. 


.618 - 







9. 3 








■'.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 


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 

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 


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 



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) 


8 - 

13 - 1 - 11 - 8 


8 - 



5 - 

12 - 1 - 16 - 6 


12 - 



13 - 



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 

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). 


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. 


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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Volume 96, number 3 - July 26, 1982 

ISSN 0028-1344 


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, 

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J. Gibson-Smith and W. Gibson-Smith 

An Ancestral Stephopoma (Mollusca: Gastropoda) from the Tertiary 
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K. E. Hoagland and Wesley R. Coe 

Larval Development in Orepidula maculosa (Prosobranchia: Crepidulidae), 

from Florida 122 

Errata 96 


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Vol. 96(3) 

July 16, 1982 



Eileen H. Jokinen 

Ecology Section, U-42 

The Univeristy of Connecticut 

Storrs, CT 06268 

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 



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 


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, 

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, 
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 




29.3 8 6.4 4.5 

74 .059 7.1 2.0 



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 



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 



1 6- 

Q 14 










.4 .6 .8 10 12 14 




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- 

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. 


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. 


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). 


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 

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 


be considered to be intraspecific individual or in- 
terspecific differences awaits more extensive 

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 


Abbott, R. T. 19.50. Snail invaders. Ndtiirnl History 

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. 

.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 

Jokinen, E. H. (In review). Allometric growth of planorbid 

Michelson, E. H. 1970. Aspidogaster conchicola from fresh- 
water gastropods in the United States. Jour. Parasitol. 

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 


July 16, 1982 

Vol. 96(3) 


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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. 


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 


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 

Location Specimens Depth Range 

1 1 1 1 1 1 M 1 1 1 1 1 1 1 1 1 1 1 

; i-?o 



5 1-60 

r< /N 

71- 80 

/ V \ 

N, ' y^ 

9 I-IOO 

\ ( 

M I-I20 



15 1-160 

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) 

Length and Width 

12 X 9 


62-240 m 

12 X 9 


92-137 m 

11.5 X 8.5 
11 X 8.5 
11 X 8 
11 X 8 


33-104 m 

10 X 7.5 
10 X 7.5 


51-102 m 

10 X 7.5 


101-126 m 

10 X 7.5 
10 X 7. 3 
9.5 X 7 


117-146 m 

9.5 X 7 


31-37 m 

9 X 7 
8.5 X 6.5 


15-18 m 

8 X 6.5 


108-130 m 

6 X 5 

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 






































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). 


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 


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- 


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 

Vol. 96(3) 

July 16, 1982 


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 

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). 


Bretton W. Kent 

Department of Zoology 
University of Maryland 
College Park, MD 20742 

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. 


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. 

Vol. 96(3) 

July 16, 1982 


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.) = 

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 


contrarium laeostomum carica 


Canal: Aperture 



.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 


July 16, 1982 

Vol. 96(3) 







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 


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. 

B. laeostomum is also distinct from fossil 
sinistral Busycon, which have either prominant 
shoulder spines, such as B. adversarium Conrad 
1862 and B. obfilosum (Grabau 1903), or strong 
external spiral cords, such as B. obrapum 
(Grabau 1903) and B. perversum robosonense 
Gardner 1948. In B. laeostomum the shoulder is 
tuberculate, but not spiny, and the spiral cords 
are absent qr only very weakly developed even 
in small individuals. The absence of fossil forms 
similar to B. laeostomum suggests that this 
species is relatively recent, possibly a northern. 
Pleistocene offshoot of B. contrarium. 

Conclusions - Hollister (1958) reviewed the 
extant species of Busycon, but made no mention 
of a form resembling B. laeostomum. This omis- 
sion appears to be due to a lack of well-pre- 
served specimens. Except for a few heavily 
beachworn specimens in the Academy of 
Natural Sciences in Philadelphia, which may be 
referable to B. laeostomum. no specimens of B. 
laeostomum appear to have been in museums 
prior to 1967. Beachworn B. laeostomum are dif- 
ficult to separate from B. contrarium and B. 
carica, so in the absence of live-collected 
specimens confusion between the three species 
was not surprising. Only in the past few years as 
live-collected B. laeostomum became available 
for study has it been possible to differentiate 
these three species. 


I would like to express my deepest thanks to 

M. G. Harasewych and E. J. Petuch for their 

numerous contributions. I would also like to 

thank R. S. Houbrick and G. M. Davis for allow- 


July 16, 1982 

Vol. 96(3) 

ing me to examine specimens in the U.S. Na- 
tional Museum and the Academy of Natural 
Sciences, to T. R. Pope for allowing me to ex- 
amine the specimen in her collection, and to G. 
J. Vermeij for his encouragement and helpful 
discussions. Support was provided by a grant 
from the National Science Foundation (number 
OCE-7901806) to the author and G. J. Vermeij. 


Abbott, R. T. 1974. Aynerican Seashells. Van Nostrand/ 

Reinhold Co., New York. N.Y. 66.3 p. 
Gill, T. 1867. On the genus Fulgur and its allies. Amer. Jour. 

Conch. 3:141-152. 
Hollister, S. C. 1958. A review of the genus Busycon and its 

allies. Part I. Paleontographica Amer. 4(28):59-126. 
Pulley, T. E. 1959. Busycon -perversum (Linne) and some 

related species. Rice Inst. Pamphlet 46:70-89. 


M. G. Harasewych 

College of Marine Studies 

University of Delaware 
Newark, Delaware 19711 


Richard E. Petit 

P.O. Box 30 
North Myrtle Beach, 
South Carolina 29582 

Cancellaria reticulata (Linne, 1 767), the type species of the type genus ofCancel- 
lariacea, inhabits subtidal sandy bottoms of the temperate and tropical western 
Atlantic. Externally, the cancellate shell is smooth and well adapted for burrow- 
ing. Internally, it has periodic reinforcing structures analogous to muricid 
varices. With the exception of an enlarged, bilohed left, cephalic tentacle, external 
features, as well as those of the pallial complex and reproductive systems are not 
unlike those of other Neogastropods. The nervou^s system differs from the usual 
rachiglossan arrangement only in the extreme anterior placement of bvx;cal 
ganglia and the resulting length of cerebro-buccal connectives. The cuticularized 
lining of the buccal cavity, highly specialized radula, lack of anterior oesophagus, 
anterior placement of buccal ganglia and simplified alimentary system posterior 
to the valve ofLeiblein are features that distinguish this superfamily from other 
Neogastropods. Although the food and feeding mechanisms remain unknown, the 
functional morphology of the alimentary system suggests a diet of soft tissue or 
fluid. Scanning electron micrographs reveal the complex structure of the radular 
tip, which raises the possibility that the cancellariid radula was formed by fusion 
of central and lateral teeth, rattier than by the loss of lateral teeth, as previously 

The cancellariaceans comprise a small, mono- 
phyietic and morphologically compact group of 
marine neogastropods that inhabit subtidal to 
bathyal sand and mud bottoms of tropical and 
temperate regions. The earliest known fossil 
record is from the Lower Cretaceous (Upper 
Albian) of Austria, while the greatest diversity 
in the Recent fauna occurs along the eastern 

Pacific coast and in the central Indo-Pacific 
area. The relationship between numerous can- 
cellariids from various Cretaceous deposits has 
not been investigated. Several genera based on 
Cretaceous species have been placed in Cancel- 
lariidae by some authors, and elsewhere by 
The taxonomic position of this group has, until 

Vol. 96(3) 

July 16, 1982 


recently, been uncertain. Troschel (1866) placed 
the family Cancellariidae in Toxoglossa, based 
largely on radular characteristics. Thiele (1929) 
and Wenz (1938-1944) included it in the rachi- 
glossan superfamily Volutacea. Olsson (1970), 
also on radular characteristics, erected the 
order Nematoglossa for the grouJD. Ponder 
(1973) showed that this lineage diverged from 
early neogastropod stock with unique modifica- 
tions of. the anterior alimentary system, and 
erected the superfamily Cancellariacea to in- 
clude the families Cancellariidae and the extinct 

Investigations into the anatomy of cancel- 
lariids have been limited, concentrating mostly 
on radular morphology (Troschel, 1865; Barn- 
ard, 1958; Olsson, 1970) and on the fore-gut 
(Graham, 1966; Ponder, 1973). 

The following is an account of the morphology 
of the shell and soft parts of CanceUaria reticu- 
lata (Linne, 1767), the type species of Cancel- 
laria, the type genus of the superfamily. No 
previous description of its anatomy has ap- 
peared, other than several comparative remarks 
by Graham (1966). 

Materials and Methods 

We were fortunate in obtaining a number of 
preserved specimens from the following locali- 

19 and 20", pumped ashore from approxi- 
mately 20 meters depth during a beach nourish- 
ment project, Atlantic Beach, Duval Co., 

19 and 40* dredged in 6 meters, W to SW from 
the southern end of Egmont Key, Florida. 

In addition, a number of shells from the fol- 
lowing localities were used to investigate shell 
structure: 4 specimens dredged in 38 meters, S 
of Sombrero Light, Marathon, Florida. 3 speci- 
mens collected on sand, minus tide, Mangue da 
Olaria, Guarapari, Espirito Santo, Brasil. 2 
specimens from St. Lucie Inlet, Martin Co., 

For anatomical studies, preserved specimens 
were immersed in 10% hydrochloric acid (HCL) 
until shells were dissolved. Soft parts were then 
rinsed and returned to 70% ethanol for dissec- 
tion. Internal shell structure was investigated 

by sectioning dry shells on a diamond saw and 
by fracturing them in a vise. 

Radulae and shell ultrastructure were ex- 
amined on a Novascan 30 Scanning Electron 

a - anus 

ag - albumen gland 

asg - accessory salivary gland 

au - auricle 

ba - bulbus aorta 

be - bursa copulatrix 

bg - buccal ganglion 

cbc - cerebro-buccal connective 

eg - capsule gland 

ct - ctenidium 

cut - cuticularized tube 

dasg - duct accessory salivary gland 

dg - digestive gland 

e - eye 

fo - female opening 

gpd - gonopericardial duct 

hg - hypobranchial gland 

ig - ingesting gland 

k - kidney 

ko - kidney opening 

let - left cephalic tentacle 

m - mouth 

me - mantle edge 

moe - mid-oesophagus 

oaoe - opening of anterior oesophagus 

od - oviduct 

opm - opening of prostrate gland to mantle cavity 

OS - osphradium 

ot - oral tube 

ov - ovary 

p - periostracum 

pc - pericardium 

pd - penial duct 

pen - penis 

pr - prostate gland 

];iro - propodium 

r - rectum 

rme - rear of mantle cavity 

rs - radular sac 

rt -radular teeth 

s - siphon 

sg - salivary gland 

srm - subradular membrane 

sto - stomach 

sv - seminal vesicle 

td - testicular duct 

te - testis 

vd - vas deferens 

ve - ventricle 

vl - valve of Leiblein 

XLC - crossed lamellar aragonite collabral 

XLT - crossed lamellar aragonite transverse 


July 16, 1982 

Vol. 96(3) 

Shell Morphology 

External: Shell fairly large for genus (to 50 
mm), heavy, biconic, and may be pseudo-umbili- 
cate (Fig. 1). Spire sharply conical (spire angle 
56-65°), comprising about 2/5 of total shell 
length. Protoconch paucispiral, of 2 glassy, 
slightly bulbous, amber-colored whorls, unorna- 
mented except by fine pits (Fig. 2). Coiling axis 
of protoconch deviated from that of teleoconch 
by 10-20°. Teleoconch, with up to 7 2/3 convex 
whorls, bears prominent collabral and spiral or- 
nament that intersect to produce the cancellate 
scupture that gives the superfamily its name. 
Suture deeply impressed. Spiral sculpture con- 
sists of 12-16 major cords on body whorl and 
4-6 on penultimate whorl, with 0-2 fine striae 
between major cords. Collabral sculpture of 
markedly prosocline ribs var3ang in number 
from 12-14 on the first post-nuclear whorl to 
28-33 on the sixth post-nuclear whorl. Ribs 
strong and evenly spaced, except for areas 
roughly every 1/3 whorl where they are weak 
and more numerous. Aperture large, hemi-ellip- 
tical, deflected from coiling axis by 22-27°. 
Outer lip with shallow indentation posterior to 
juncture with siphonal canal and 9-13 strong, 
slightly recessed lirae that diminish rapidly. In- 

side I 

reticulata ^Z,iK?i<5| A pert )i ml and right 
sp,. cuium dredged in 6 meters of water, W toSW 

from the xouthem end of Eginnnt Key, Florida. (USNM 
806999). 1 1/2 X. 

ner lip with 2 pronounced columellar folds and a 
distinct siphonal fold. Posterior fold, much the 
largest, distinctly bifurcate. Anterior fold some- 
times subdivided by a shallow furrow, but to a 
much lesser degree. The posterior fold overlies 
the siphonal fasciole. Siphonal canal short 
(about 1/5 aperture length), slightly recurved 
dorsally and to the right. Base color is white, 
with patches of light ginger to chocolate brown 
that are sharply limited by both collabral and 
spiral bands of white. The collabral bands 
overlie areas of weak ribbing and areas midway 
between them. Spiral white bands are situated 
medially and near the siphonal margin. The 
aperture is white. 

Internal: Examination of shells sectioned 
through planes normal to the coiling axis and 
transverse to growth lines revealed that aper- 
tural lirae and columellar folds are not spirally 
continuous, but are prominent only every 1/3 
whorl, and disappear or are greatly reduced be- 
tween (Fig. 3). The spiral lirae of the outer lip 
are produced intermittently, and do not extend 
to the outer lip. The area between the lirae and 
the edge of the outer lip is thin and corresponds 
to the areas of weak collabral ribbing noted pre- 
viously. The columellar folds reach maximum 
extension into the aperture in direct opposition 
to maximum Ural extension, and are reduced 
and appressed to the columella between. 

Ultrastructure: The yellow, unornamented 
periostracum reaches a thickness of 4-6 ^m 
(Figs. 6, 7 P), and overlies a layer of crossed- 
lamellar aragonite (Figs. 4-7, XLC) in which the 
lamellar planes are collabral. As the axial ribs, 
spiral cords and lirae are composed of this layer, 
its thickness is quite variable (from 1 to 3 mm in 
mature specimens). A second crossed-lamellar 
shell layer (Figs. 4-6, XLT), in which the lamel- 
lar planes are roughly perpendicular to those of 
the preceding layer (XLC) as well as to the coil- 
ing axis, lines the inner edge of the shell, and is 
more uniform in thickness (450 to 750 \xm). 

The general form and diversity of sculpturing 
among 73 living species of Cancellariidae have 
been illustrated in color in the forthcoming 
book, the Compendium of Seashells (pp. 
225-231) by Abbott and Dance (1982). Speci- 
mens figured are from the collection of the 

Vol. 96(3) 

July 16, 1982 


FIGS. 2-7. Details of shell structure -j? L ancellana reticulata iLinmi. 2, i ,._,(.,...,„/,. Scak bar = WO tirn. 3, Anterior view of 
shell in which 112 whorl of the outer lip was removed along its midline to show internal structure. Scale bar =5 mm. 4, Fracture 
surface. Plane of fracture parallel to outer lip and passing through apertural lirae. Scale bar =400 tim. 5. Fracture surface. 
Plane of fracture perpendicular to outer lip. Scale bar = UOO urn, 6. Partial fracture through outer, collabral layer but not 
through inner, transverse layer. Scale bar = 100 lAm. 7, Outer edge of fracture surface in a plane parallel to the auier lip. Scale 
bar=i ijm. 


July 16, 1982 

Vol. 96(3) 

junior author and from the type collection at the 
U.S. National Museum. 

Soft-parts Morphology 

External features: The shell-less animal con- 
sists of about 5 whorls, of which the mantle cav- 
ity occupies approximately 2/3 whorl, the volu- 
minous kidney (Fig. 8, k) 1/3 whorl, and the 
large digestive gland (Fig. 8, dg) about 4 whorls. 
Animals were able to retract about 1/2 whorl in- 
side the aperture. The broad attachment area of 
the columellar muscle is situated below the rear 
of the mantle cavity when the animal is with- 
drawn. Preserved animals are uniformly pale 
orange in color, lacking a discernable pattern. 
The mantle edge (Fig. 8, me) is thin and finely 
papillose. The long, narrow, posteriorly taper- 
ing foot has a pronounced propodium (Fig. 8, 
pro) and lacks an operculum. Of the cephalic ten- 
tacles, the left (Fig. 8, let) is longer and broader 
than the right, and bears 2 papillae above the 
eye (Fig. 8, e). The siphon (Fig. 8, s) extends 
slightly beyond the mantle edge and lies over 
the left tentacle. 

Mantle cavity: The mantle cavity is narrow 
and deep, with pallial organs situated as in other 
Neogastropods. The osphradium (Fig. 8, os) is 
short (7 mm) and broad (L/W = 3), bearing about 
70 leaflets per side. There is an unusually large 
space between the osphradium and the cteni- 
dium (Fig. 8, ct) (about 1.5 times the width of 
either organ), so that the ctenidium, which is 
long, slightly narrower than the osphradium and 
consists of about 250 leaflets, is suspended from 
the top of the mantle cavity. On the right side of 
the ctenidium lies the transversely pleated hypo- 
branchial gland (Fig. 8, hg), and to the right of 
it, the rectum (Fig. 8, r) and genital ducts. The 
kidney forms the right rear wall of the mantle 
cavity, and opens into it by a single, vertical slit 
(Fig. 8, ko). The large pericardium (Fig. 8, pc) is 
embedded in the left side of the kidney, and 
forms the left rear wall of the mantle cavity. 

Alimentary system: When fully extended, the 
narrow, pleurombolic proboscis can be pro- 
truded through the rhyncostome to a length 
equal to 1 1/2 times the shell length. It is of the 
same color as the foot, and twice as broad as 
high, being smooth along its curved dorsal sur- 

face and papillose along the entire length of its 
flat ventral surface (Fig. 9). When retracted, the 
proboscis is folded within the proboscis sheath. 
The retractor muscles originate from the body 
wall as well as from the columellar muscle. 
From the mouth (Fig. 9, m) situated terminally 
on the curved anterior of the proboscis, a short 
(750 f^m) oral tube (Fig. 9, ot) leads to a 
cuticularized, ventrally overlapping tube-like 
structure (Fig. 9, cut) [Upper Shield or Mandible 
of Olsson, 1970], which is narrow anteriorly and 
expands posteriorly to enclose the buccal mass. 
The floor of the buccal mass is lined by a broad, 
cuticularized sub-radular membrane (Fig. 9, 
srm) [Inner Shield of Olsson, 1970] which sup- 
ports the radula in a narrow groove. The radula 
(Fig. 12) consists of a ribbon supporting 73-109 
rows of single teeth, of which slightly less than 
half are directed anteriorly, the balance posteri- 
orly. The anteriorly directed teeth may be pro- 
truded slightly beyond the end of the cuticula- 
rized tube. The posteriormost teeth are poorly 
formed, and lie in the radular sac (Fig. 9, rs) 
which extends slightly below the buccal mass. 
The division between the anteriorly and posteri- 
orly directed teeth is slightly anterior to the 
oesophageal opening (Fig. 9, oaoe). Several 
specimens were observed with groups of teeth 
(4-9) coiled in a loop under the oesophagus, indi- 
cating the mechanism by which they are redi- 
rected. Individual teeth are long (1.2-1.9 mm) 
(Fig. 12), narrow (25 ^m at the base) (Fig. 13) 
and extremely flexible. Toward the tip (Figs. 14, 
15), they constrict (20 ^<m) and bear 3 highly 
modified cusps, each with secondary, anteriorly 
directed cusps. The 2 lateral cusps are directed 
inward, and interlock medially with the up- 
wardly directed central cusp. The oesophagus 
opens along the roof of the buccal cavity, where 
it appears to be cuticularized. At the rear of the 
buccal mass is the valve of Leiblein (Fig. 9, vl), 
and beyond it the long, narrow mid-oesophagus 
(Fig. 9, moe) winds backward to the stomach 
(Fig. 8, sto). Just behind the buccal mass lie two 
pairs of salivary glands. One pair (Fig. 9, sg) is 
narrow and long, entering the buccal mass just 
anterior to the oesophageal opening. These are 
homologous to the normal salivary glands of 
other prosobranchs. The second pair (Fig. 9, 

Vol. 96(3) 

July 16, 1982 



'IGS. 8-11. Anatomical features o/Cancellaria reticulata (Linne). 8, Mate specimen removedfrom shell, uncoiled and mnntle 
[Tvity opened mid-darsally to display contents. 9, Dissection of proboscis tip, viewed from left side. 10, Diagrammatic 
Wepresentation of the female genital ducts. 11, Diagrammatic rejjresentation of the male genital diicts. 


July 16, 1982 

Vol. 96(3) 

FIGS. 12-15. Scanning electron micrograpks of raduia o/ Cancellaria reticulata (Linne). 12, Entire radular ribbon with 
suhradular membrane. Scale bar = i.00 \im.. 13, Attachment of basal ends of teeth to radular ribbon. Scale bar=10 nm. 14, Ter- 
minal ends of radular teeth. Scale bar = 10 (j?n. 15, Terminal ends of radular teeth. Scale bar = J, ^im. 

asg) are smaller and sausage-shaped, giving rise 
to ducts (Fig. 9, dasg) that lead anteriorly, 
becoming convoluted and fusing just prior to 
entering the oral tube near the mouth. These ap- 
pear to be homologous to the accessory salivary 
glands of muricids and volutids. At the ventral 
posterior ends of the buccal mass lie the buccal 
ganglia (Fig. 9, bg), and from them the cerebro- 
buc( ' -I'nnectives (Fig. 9, cbc), bound together 
with I ■ poboscis nerves, run medially down 
the len;. ' ^r i!,,i probo.scis. The oesophagus 
passes through il;c )iy^h\y concentrated nerve 

ring and becomes highly convoluted before 
entering the stomach. There is no separate 
gland of Leiblein, but rather a narrow glandular 
strip runs along the mid-oesophagus. The 
stomach (Fig. 8, sto), small, narrow and tube- 
like, runs across the anterior face of the diges- 
tive gland (Fig. 8, dg). The narrow oesophagus 
(1/3 the diameter of the bulbus aortae. Fig. 8, 
ba) enters on the left, near the opening of the 
duct to the digestive gland. Longitudinal folds 
run the length of the stomach into the intestine, 
which continues as the rectum (Fig. 9, r) along 

Vol. 96(3) 

July 16, 1982 


the right side of the roof of the mantle cavity, 
but detaches from it 1-2 mm before ending in 
the anus (Fig. 8, a). There appears to be no anal 

Female reproductive system: The light-yellow 
ovary (Fig. 10, ov) lies at the apex of the diges- 
tive gland, forming the terminal 1 1/2 whorls. It 
is composed of numerous thin ascini that con- 
verge to form the oviduct (Fig. 10, od) that runs 
anteriorly along the right ventral side of the 
visceral mass. The oviduct is joined by the gono- 
pericardial duct (Fig. 10, gpd) just before enter- 
ing the rear of the mantle cavity and joining the 
large, arched albumen gland (Fig. 10, ag). Be- 
tween the albumen gland and the large, rectan- 
gular capsule gland (Fig. 10, eg) is the ingesting 
gland (Fig. 10, ig) that appears as a small mass 
of dark-brown ridged tubules. Its duct serves as 
the seminal receptacle. At the anterior end of 
the capsule gland is the large, squarish bursa 
copulatrix (Fig. 10, be). Cutting through the 
muscular wall reveals an arched, rapidly elarg- 
ing tube with the female opening (Fig. 10, fo) at 
its end. The ventral pedal gland, a 2-3 mm deep 
invagination lined with glandular folds, is situ- 
ated along the midline of the sole of the foot, 
about 1/4 of its length from the anterior edge. 

Male reproductive system: The dark-brown 
testis (Fig. 11, te) consists of a mass of ascinous 
tubules imbedded in the columellar side of the 
digestive gland. These tubules join together to 
form a common testicular duct (Fig. 11, td) 
which runs anteriorly along the right ventral 
side of the visceral mass. When next to the pos- 
terior portion of the kidney, it becomes con- 
voluted, acting as a seminal vesicle (Fig. 11, sv), 
then straightens and enters the rear of the man- 
.tle cavity ventral to the rectum, where it 
enlarges in diameter and again becomes con- 
voluted. These convolutions act as a prostate 
gland (Fig. 11, pr), and have an opening to the 
mantle cavity (Fig. 11, opm). The vas deferens 
(Figs. 8, 11, vd) winds along the base of the man- 
tle cavity from the prostate gland to the base of 
the penis (Figs. 8, 11, pen). The penis is long 
(18-20 mm), narrow (1 1/2-2 mm) and flat, bear- 
ing a terminal papilla (2-2 1/2 mm). Judging by 
the highly coiled penial duct (Fig. 11, pd), the 
penis is capable of considerable extension. 

Nervous system: The cerebral, pleural, pedal 
and suboesophageal ganglia are fused into a 
concentrated nerve ring through which pass the 
oesophagus and the anterior aorta. The buccal 
ganglia are at the base of the buccal mass at the 
tip of the long proboscis, with the cerebro-buccal 
connectives reaching 50 mm in length. The su- 
praoesophageal ganglion lies over the oesopha- 
gus on the right side of the cephalic hemocoel, 
just behind the proboscis sheath, and gives rise 
to the osphradial nerve. The visceral ganglion 
lies in its usual position at the innermost end of 
the floor of the mantle cavity. 


The anteriorly rounded, posteriorly tapering 
shell, short siphonal canal, lack of pronounced 
ornament and smooth surface identify Cancel- 
laria reticulata as a frontally burrowing, in- 
faunal neogastropod. The majority of specimens 
examined (27 of 32) had 1 or more crab breaks 
on the last whorl. These breaks, however, failed 
to penetrate beyond the penultimate apertural 
lirae, suggesting that these serve as shell rein- 
forcements or internal varices. The periodic in- 
crease in the size of the columellar folds in op- 
position to the lirae of the outer lip decreases 
the size of the aperture and serves, at least in 
part, an antipredatory function. This periodicity 
of shell structure indicates that growth is 
episodic and rapid, occurring in increments of 
approximately 1/3 whorl. 

Knight, et al. (1954) suggested that cancel- 
lariids (included in their Volutacea) evolved 
from the Mesozoic Nerineacea, which, along 
with some of the Soleniscinae (Subulitacea), had 
internal folds of considerable complexity. A re- 
examination of these groups for periodicity of 
internal structure may shed light on the origins 
of Cancellariacea. 

Ultrastructural studies reveal that the shell 
consists of a thin periostraclim overlying 2 
orthogonal layers of crossed-lamellar aragonite. 
In the outer layer, which comprises the bulk of 
the shell, the crystal planes are oriented parallel 
to the outer lip, while in the thinner, inner layer, 
which lines the inner surface of the shell, the 
crystal planes are perpendicular to those of the 
outer layer. 


July 16, 1982 

Vol. 96(3) 

The fracture plane of the thicker outer layer 
would cause breaks in the shell to run parallel 
to the outer lip, while the orthogonal inner layer 
greatly increases the mechanical strength of 
the shell. In studies of shell crystallography, 
Boggild (1930) reported that "Different Tertiary 
and Quaternary species of Cancellaria . . . were 
all quite alike and built up of the ordinary three 
layers." [transverse, collabral, transverse], but 
did not indicate where in the shell the sections 
were taken. Along the inner lip, a layer of trans- 
verse crystals is deposited over the outer layer 
of shell, possibly accounting for the discrepancy. 

Cancellaria reticulata is similar to other 
neogastropods in the morphology of its head- 
foot, pallial complex, reproductive and nervous 
systems, with the following noteworthy excep- 
tions. The enlarged and bilobed left cephalic ten- 
tacle was found, in all the preserved specimens 
examined (n = 8), to run along the base of the 
siphon, with the two papillae extending over and 
beyond the opening. It is not known if it func- 
tions together with the siphon, forming a tube, 
or is simply oriented in the anteriormost direc- 
tion. Several other cancellariid species have 
similar, and, in some cases, more elaborate ar- 
rangements (unpublished observations). Of the 
pallial organs, the position of the ctenidium is 
unusual, in that it is displayed to the top of the 
roof of the mantle cavity. The morphology of 
both male and female reproductive systems is 
typically neogastropod (as exemplified by Nu- 
cella lapillus, see Fretter, 1941), with minor 
modifications only in the anteriormost portion 
of each system [bursa copulatrix- female; penial 
papillae -male]. The nervous system differs 
from that of other neogastropods chiefly in the 
location of the buccal ganglia and the extreme 
length of the cerebro-buccal connectives. 

The unique and highly specialized alimentary 
system is the basis for the superfamilial status 
of this group, yet the food and feeding mecha- 
nisms remain unknown for all cancellariids. 
Both Graham (1966) and Olsson (1970) have 
speculated that these animals feed on soft- 
bodied micrncrganisms which are "brushed" into 
the gut by the radular teeth. Like Graham 
(1966), we wt-re unable to find any traces of solid 

food in the alimentary systems of the specimens 
we examined. 

The extreme length of the ventraliy papillose 
proboscis suggests that cancellariids feed at a 
distance on animals or tissues they cannot reach 
directly. The presence of well developed acces- 
sory salivary glands is generally associated with 
a carnivorous rather than a scavanger diet. 

It is currently believed that cancellariid radu- 
lae consist of rows of rachidia, the lateral teeth 
having been lost (Ponder, 1973). Scanning elec- 
tron micrographs of the distal tips of radular 
teeth reveal an elaborate cusp structure, which 
can be interpreted as being a tri-cusped rachi- 
dian in which each of the primary cusps have 
developed secondary cusps. As the lateral cusps 
bear a strong resemblance to rachiglossate 
lateral teeth, we feel an alternative interpreta- 
tion may be that lateral teeth fused with the 
rachidian prior to elongation in the ancestral 

Viewed functionally, the extreme anterior 
placement of the valve of Leiblein would serve 
to minimize the volume of food anterior to it, 
permitting rapid interaction with salivary gland 
secretions, and, perhaps more importantly, to 
prevent the backflow of liquified food posterior 
to it. The lack of a gland of Leiblein, the long, 
thin oesophagus and simple tubular stomach 
suggest that little further treatment of food is 
required prior to digestion. 

Based on certain general similarities between 
the cancellariid alimentary system and those of 
pyramidellids and melanellids, we suggest that 
cancellariids are piercing, fluid feeders rather 
than microphagous grazers. 

We wish to thank William G. Lyons of the 
Department of Natural Resources, State of 
Florida, as well as Dr. Harry G. Lee, Mr. Allan 
Walker and Mrs. Jane Dawley, all of Jackson- 
ville, Florida, for making available the pre- 
served material used in this study. Special 
thanks are due to Mr. L. C. Bailey of the Engi- 
neering Test Center, E. I. du Pont de Nemours 
Co. for his help with scanning electron micro- 
scopy. Critical review of the manuscript by Dr. 
M. R. Carriker, College of Marine Studies, 

Vol. 96(3) 

July 16, 1982 


University of Delaware, is gratefully acknow- 


Abbott, R. Tucker and S. Peter Dance. 1982. Compendium 

ofSeashelh. E. P. Dutton, N.Y. 400 pp., 4,300 color plates. 
Barnard, K. H. 1958. The radula of Cancellaria. Jour. 

Confhology 24(7):243-244. 
Boggild, U. B. 1930. The sheU structure of mollusks. Acad. 

Roy. Sci. Lettres Danemark. Mem. ser. 9, 2:230-325, 

pi. 1-15.- 
Fretter, V. 1941. The genital ducts of some British steno- 

glossan prosobranchs. Jour. Mar. Biol. Assoc. U.K. 


Graham, A. 1966. The fore-gut of some marginellid and 
cancellariid prosobranchs. Stud, Trop. Oceanog. Miami 

Knight, J. B., Bratten, R. L. and Yochelson, E.L. 1954. 
Status of Invertebrate Paleontology, 1953. V. Mollusca: 
Gastropoda. Bull Mus. comp. Zool. Hare. 112(3):173-179. 

Olsson, A. A. 1970. The cancellariid radula and its interpre- 
tation. Palaeontogr. Amer. 7(43):19-27. 

Ponder, W. F. 1973. The origin and evolution of the Neo- 
gastropoda. Malamlogia 12(2):295-338. 

Thiele, J. 1929. Handhuch der systetnatischen Weichtier- 
kunde. (Jena, Gustav Fischer, 1929-1935):1154 p. 

Troschel, F. H. 1856-1893. Das Gebiss der Schnecken zur 
Begriindung einer natiirlichen Classification. 2. Berlin. 

Wenz, W. 1938-1943. Handbuch der Paldozoologie (0. H. 
Schindewolf, ed.). Berlin. 


William Miller, III 

Earth Sciences Dept., Tulane University 
New Orleans, LA 70118 

Donax variabilis is the easily recognized 
tellinacean bivalve that inhabits the high energy 
foreshore zone of ocean beaches from Virginia 
to Texas (Abbott, 1968). This clam is remarkable 
for its abilities as a rapid burrower, for its tidal 
migrations on the beachface, and for its occur- 
rence in densely packed "colonies" in an en- 
vironment from which typically little else is col- 
lected alive (Stanley, 1970). The purpose of this 
note is to describe the distribution of D. 
variabilis, both living populations and dead 
shells, along a section of moderate energy ocean 
beach at Ship Island, Mississippi, a barrier 
island separating Mississippi Sound from the 
Gulf of Mexico. This section of beach was (in 
May, 1980) composed of fine quartz sand with a 
trace of shell fragments, showed no signs of 
pollution, and featured a foreshore slope of only 
a few degrees. Other than infrequent churning 
of the substrate by swimmers using a beach just 
to the west of the study area, the beachface is 
undisturbed by man. 


Sheree Kooser 

148320 Hartwell Terrace 
Novelty, OH 44072 

Beginning 90 m east of the swimming area on 
the western end of Ship Island, 11 samples were 
collected from the middle beachface at 90 m in- 
tervals proceeding east along the beach. At each 
sample station one litre of sand was scooped 
from the beach and wet sieved using nested 
screens having openings of 4.0 and 2.4 mm to 
recover all shells contained in the samples. Live 
individuals, as well as articulated and disarticu- 
lated dead shells, were counted and returned to 
the beachface. The results of the survey are 
summarized in Table 1. 

Nearly all living Z). variabilis recovered were 
about 2 cm in length (adults); only a few in- 
dividuals were larger or smaller. The majority of 
the specimens were alive (86%), and the dead 
shells were mostly disarticulated and abraded 
(84%). The survey appears to have encountered 
at least two "regional aggregations" of living D. 
variabilis in the beachface: one at sample sta- 
tions 3 and 4, and another at stations 7, 8, and 9. 
All sample stations except 2 and 6 encountered 


July 16, 1982 

Vol. 96(3) 

TABLE 1. Distribution o/Donax variabilis at Ship Island. 
Misfdssippi: May 20, 1980. 


Live _ 

Dead Shells 









































N = 629 


N = 86 

x = 57.2 


x = 7.8 

s = 59.0 


s = 7.8 

at least local aggregations. The smaller, local 
aggregations appeared to be clumped together 
forming the larger, regional aggregations. Well- 
defined aggregations were not preserved 
among the dead shells. The largest local aggre- 
gations of living clams occurred near the end of 
the transect farthest from the public swimming 

The distribution of Donax is of interest to 
paleontologists because of its use as an indicator 
of proximity to ancient shorelines in paleoenvi- 
ronmentai reconstructions (for example, see 
Mixon and Pilkey, 1976). However, as shown in 
this study, relatively few dead shells of D. 
variabilis were found in situ in the beachface at 
Ship Island. It seems that after individuals die 
their shells are temporarily concentrated at the 
plunge step just offshore from the beachface 
(see Davis, 1978). Tides and currents probably 

redistribute the dead shells to various areas in 
the nearshore zone and other parts of the inner 
shelf. This transport of D. variabilis shells out 
onto the adjacent shelf explains the admixture 
of small numbers of these swash zone inhabi- 
tants in Pleistocene fossil assemblages of sub- 
tidal marine origin with which we are familiar, 
and argues against the use of Donax as an une- 
quivocal indicator of proximity to ancient shore- 
lines. Moreover, tropical storm overwash and 
the migration of tidal inlets tend to recycle 
Donax shells into backshore, interdune, and 
even backbarrier environments. Shells re- 
worked from relict sediments compound the 
problem (see Pilkey et al.. 1969). In conclusion, 
paleoenvironmental interpretations involving 
Donax always should take into account the 
taphonomic history and paleosynecologic prop- 
erties of fossil deposits, and not focus on a few 
selected taxa as "reliable" indicators of environ- 


Abbott, R. T. 1968. Seashells of North America. Golden 
Press, New York, 280 pp. 

Davis, R. A. 1978. Beach and nearshore zone. In. R. k. Davis 
(ed.), Coastal Sedimentary Environments: Springer- 
Verlag, New York, pp. 237-285. 

Mixon, R. B. and 0. H. Pilkey. 1976. Reconnaissance geo- 
logy of the submerged and emerged Coastal Plain Prov- 
ince, Cape Lookout area, North Carolina. U. S. Geol. 
Survey, Pi-oJ. Paper 859, 45 pp. 

Pilkey, 0. H.,'b. W. Blackwelder, L. J. Doyle. E. Estes and 
P. M. Terlecky. 1969. Aspects of carbonate sedimentation 
on the Atlantic continental Shelf off the southern United 
States. Jour. Sed. Petrology 39:744-768. 

Stanley, S. M. 1970. Relation of shell form to life habits of 
the Bivalvia (Mollusca). Geol. Soc. Am. Memoir 125. 
296 pp. 


SEA SHELLS P. Q. Box 83 

Glen Ellen, CA 95442 




Specmen Shells 

Offering microscopic and miniature (to '/. inch) shells 
from the Florida Keys, vrith accurate locality data Also 
unsorted grunge; u/rite for list. 

Margaret Teskey 

P.O. Box 273 

Big Pine Key. Fl. SSOiS 

Vol. 96(3) 

July 16, 1982 




J. Gibson-Smith and W. Gibson-Smith 

Escuela de Geologia, Minas y Geofisica, Universidad Central de Venezuela, 
Apartado 47.351, Caracas 1041 A, Venezuela 

Lima albicoma Dall. 1886. from the Caribbean Recent. i,s reassigned to 
Divarilima Powell. 1958. a Recent taxon from the Western Pacific, the ancestor 
probably being the genus Badiotella Bittner. 1890. known only from the middle 
Trias of Europe. The new subspecies D. albicoma handini, is described from the 
Venezuelan Recent. 

Lima albicola Dall, 1886, was found off Cuba 
(type locality) and off Barbados at a depth of 
about 200 m; it has been re-figxired by Abbott 
(1974:454:5249). This small shell (8 mm') with its 
long, straight, anterodorsal margin is similar in 
outline to the large, coarsely sculptured Lima 
S.S., but resembles Ctenoides Morch, 1853, in 
possessing a divaricate sculpture. It differs from 
all Western Atlantic limids in having a very 
deeply excavated lunule, or "truncature", as Dall 
(1886:225) called it. A similar Recent species 
from off eastern Australia and Kermadec 
Island, Lima sidneyensis Hedley, 1904, was 
made the type of Divarilima Powell, 1958 (Cox 
& Hertlein, in Moore 1969:N389), thus drawing 
attention to the divaricate sculpture, a feature 
of Ctenoides, rather than to the characteristic 
lunule. Lima albicola can be assigned to 
Divarilima. The same lunule is present in 
another small genus, Badiotella Bittner, 1890 
(Cox & Hertlein, in Moore 1969:N386), known 
only from the middle Trias of Europe which, 
although lacking the divaricate sculpture, can 
probably be considered ancestral to the present 
day forms of Divarilima from the Western 
Pacific and Western Atlantic, forms to be 
regarded as Tethyan relicts. 

From the north coast of Venezuela comes a 
second Caribbean form, here named: Divari- 
lima albicola handini, n. subsp. 

Divarilima albicola handini 

Gibson-Smith & Gibson-Smith, n. subsp. 

Figures 1,2. 
Description - Shell small, trigonally ovate. 
Anterodorsal margin nearly straight, extended, 
coinciding with a strong umbonal ridge along 
which runs a fine keel. Posterior margin almost 
straight above, these two margins subtending 
an acute angle at the umbo. Ventral margin 
deeply rounded. Shell almost equilateral. Poste- 

FIGS. 1-3. 1 and 2, Divarilima albicoma handini n. subsp. 
Holotype, external and internal views, right valve, height 
10.0 mm, length 8.2 mm. Recent of Venezuela. USNM no. 
784699. 3, Divarilima albicoma Dall. 1886. Paratype. inter- 
nal view, left valve, height .V.S mm. length 3.S mm. Caribbean 
Recent. MCZ 7829. 


July 16, 1982 

Vol. 96(3) 

rior auricle moderate, anterior absent. Cardinal 
area triangular with a narrow ligament pit over- 
hung slightly by the small sharp umbo; hinge 
edentulous. Lunule triangular, concave, both 
borders angulate (90°), sculptured with a weak 
radial riblet and weaker, irregular, radial wrin- 
kles. Ornament of fine, close, punctate grooves, 
divaricate along the median line from to umbo to 
ventral margin; angle of divarication very acute 
and hardly to be seen. Several strongly marked 
growth stages stepping down across the disc. 
Color a very pale brown, translucent; growth 
stages marked by narrow, opaque white, con- 
centric bands. Remnant of pale brown perio- 
stracum within the lunule. 

Holotype - USNM No. 784699. A right valve, 
height 10.0 mm, length 8.2 mm, semi-diameter 
2.5 mm. 

Type locality - Recent, Chichiriviche de la 
Costa, Federal District, Venezuela. 

Remarks - Apart from the holotype there are 
two, small paratypes each measuring approx- 
imately: height 3.0 mm, length 2.75 mm. All are 
right valves and came from a sand sample col- 
lected at 30 m by SCUBA-diver Alan Handin. A 
figure of a paratype of D. albicoma (courtesy Dr. 
Kenneth J. Boss, MCZ) is shown for comparison 
(Fig. 3). 

Comparisons - Divarilima albicoma handini 

n. subsp. differs from D. albicoma Dall in being 
less produced anteriorily and posteriorily, re- 
sulting in a narrower, more equilateral shell. In 
handini the anterodorsal and posterior margins 
subtend an acute angle at the umbo; in albicoma 
the angle is obtuse. The narrowness of handini 
is reflected in the much more acute angle of 
divarication along the median line. Further- 
more, albicomn is described as having two lines 
of divarication (the figure even shows three), in 
handini there is only one. Whether the depth 
ranges of the two forms overlap remains to be 
seen; it may be significant that neither taxon 
was found in seafloor samples from nine off- 
shore drilling locations, in water depths from 55 
to 150 m. 


Abbott, R. T. 1974. Armrican Seashells. 2nd ed. Van 
Nostrand Reinhold Co., New York; 663 pp., 4000+ text 
figs.; 24 pis. (in color). 

Dall, W. H. 1886. Reports on the results of dredging, under 
the supervision of Alexander Agassiz, in the Gulf of 
Mexico (1877-78) and in the Caribbean Sea (1879-80) 
by the U.S. Coast Survey Steamer "Blake" Bull. Mus. 
Comp, Zool., Vol. 12, pp. 153-318, 9 pis. Harvard Univ., 
Cambridge, Massachusetts, U.S.A. 

Moore, R. C. 1969. Treatise on Invertebrate Paleontology, 
Part N, vol. 1, Mollusca 6, Bivalvia, pp. i-xxxviii + 
N1-N489, text figs. Geol. Soc. Amer. Inc. & Univ. Kansas. 


J. Gibson-Smith and W. Gibson-Smith 

Escuela de Geologia, Minas y Geofisica, Universidad Central de Venezuela 
Apartado 47.351, Caracas 1041 A, Venezuela 

The presence of two, new melampid species in the Venezv£lan Recent i§ re- 
ported: Detracia roquesana n. sp. and Tralia venezUelana n. sp., the latter occur- 
ring also in the early Miocene Cantaure Formation, Paraguand Peninsula, and 
the late Pliocene Mare Formation, Cabo Blanco. T. venezuelana is only the second 
species o/ Tralia to be recorded from the Western Atlantic. 

The pulmonate genus Melampus is repre- 
sented in the Venezuelan Recent by Melampus 

(Melampus) coffea (Linnaeus, 1758) (Fig. 1) and 
Melampus (Pira) monilis (Bruguiere, 1789) 

Vol. 96(3) 

July 16, 1982 


Figs. 2, 3). These two taxa range from Florida 
ind Bermuda to Brazil (Abbott 1974:331). The 
former is generally known as cojfeus, but it has 
Deen pointed out by Altena (1975:86) that being 
1 noun, it is correctly, coffea. Two other 
aulmonate genera present in the Recent of 
Venezuela are Detracia Gray in Turton, 1840, 
ind Tralia Gray in Turton, 1840. The former is 
-epresented in Florida and the northern Carib- 
bean by the type-species D. bullaeoides 
Montagu, 1808), D. floridana (Pfeiffer, 1856) 
md D. clarki Morrison, 1951, but in Suriname 
Altena, 1975:86) and Brazil (Marcus & Marcus, 
1965:42) the representative is D. parana 
Viorrison, 1951, the type locality being Para 
= Belem), Brazil. In Venezuela, Detracia has 
lot been found on the mainland but is present on 
;he islands of Los Roques where it is repre- 
sented by a new species here identified as 
letracia roquesana n. sp. The genus Tralia is 
cnown in the Recent of the Western Atlantic by 
I single species, T. ovula (Bruguiere, 1789), 
vhich ranges from southeast Florida and Ber- 
nuda to Barbados (Abbott 1974:33). It has been 
■eported also from the Caribbean coast of 
^anama (Olsson & McGinty 1958:19) and from 
];uracao (Coomans 1958:103), in both instances 
IS Tralia pusilla (Gmelin, 1791), a synonym. It 
)ccurs also in Venezuela (Figs. 4, 5) where, 
lowever, a second, more common form is pre- 
sent which is identified here as, Tralia 
jenezuelayia n. sp. The ancestor of the latter is 
)resent in the early Miocene (Burdigalian) Can- 
a.ure Formation of the Paraguana Peninsula; it 
vas referred to, Tralia cf. oimla, by Gibson- 
Smith & Gibson-Smith (1979:22), but is now 
bund to be identical with T. venezuelana n. sp.; 
■t occurs also in the late Pliocene Mare Forma- 
tion, Cabo Blanco. The Melampinae are hardly 
mown as fossils in the region, one other record 
)eing of Tralia vetula Woodring, 1928, from the 
^liocene Bowden Formation, Jamaica. 

Detracia roquesana 

Gibson-Smith & Gibson-Smith, n. sp. 
Fig. 6 

Description - Shell small, about 10 mm in 

height, obovate, widest at the middle. Whorls 
adpressed, about 11 in all, the bodywhorl about 
two-thirds of the shell height. Protoconch at 
right angles to shell axis, one-half turn only visi- 
ble, brown. The shell lacks sculpture and is 
spirally banded in shades of brown. Aperture 
narrow, the outer lip not flared. Inner lip with 
heavy callus, a strong fold at the base of the col- 
umella and a weak fold, well within the aper- 
ture, at the middle of the parietal area. Above 
the latter the parietal area is weakly excavated 
and lacks callus, with the result that the lower 
edge of the area becomes a broad, low fold, or 
pseudo-fold. Within the outer lip are 3 to 8 lirae, 
the lowest and largest projects towards the col- 
umella fold, leaving only a narrow gap between. 
Opposite the parietal fold and pseudo-fold, the 
gap is again narrowed by two strengthened lirae 
lying opposite within the outer lip. 

Holotype - USNM 784718. Height 10.6 mm, 
diameter 5.8 mm. 

Locality - Recent, Islas Los Roques, Vene- 

Paratypes - British Museum (Natural His- 
tory) and the Natural History Museum Basel. 
Remainder in the collections of the authors. 

Remarks - The 6 specimens were collected in 
beach drift and the exact ecological niche is 
unknown, but is, presumably, above high tide 
level as for other members of the family. The 
islands are fringed by coral reefs and man- 
groves abound. There is no obvious variation in 
shell morphology. 

Comparisons - Detracia roquesana n. sp. dif- 
fers from all the four known Western Atlantic 
species; D. bullaeoides is more elongate, lacks a 
parietal fold and lirae within the outer lip, and 
the protoconch -is white; D.floridana is smaller, 
more globose and the lirae within the outer lip 
are not enlarged opposite the columella and 
parietal folds; D. clarki has 2 lower columella 
folds and is sculptured with spiral, incised lines 
above the shoulder and around the base and, 
lastly, D. parana is smaller, more rotund and 
normally possesses only a single, outer lip lira, 
many shells lacking even that. 


July 16, 1982 

Vol. 96(3) 

FIGS 1-9. The subfamily Melampinae {n Venezuela (for explanations, see opposite). 

Vol. 96(3) 

July 16, 1982 


Tralia venezuelana 

Gibson-Smith & Gibsoyi-Stiiith, n. sp. 
Figs. 7, 8 9 

Tralia cf. otmla (Bruguiere, 1789), Gibson-Smith & Gibson- 
Smith, 1979, GEOS No. 24, p. 22. 

Description - Shell small up to about 15 mm 
in height, obovate, greatest diameter about the 
middle. Protoconch small, pimple-like, at right 
angles to shell axis, about one-half turn only visi- 
ble. Teleoconch of 8 whorls, the weakly shoul- 
dered bodywhorl comprising three-quarters of 
the shell. Shell glossy, color dark brown with 
faint spiral and axial banding in shades of 
brown. Sculpture of up to 9, spiral, pitted 
grooves above the shoulder, marking the rows 
of setae of the periostracum; remainder of 
surface covered with crowded, microscopic 
scratches, stronger around the base. Shell sur- 
face interrupted by growth incrementals. Aper- 
ture wider and flaring below, restricted at the 
middle by a heavy spiral thickening within the 
outer lip. Columella with a strong basal fold and 
a stronger one above at the base of the parietal 
area with, on the parietal area proper, weaker 
third and fourth folds, the uppermost the weak- 
est, lying well inside the aperture and more 
easily seen in juvenile specimens. 

Holotype - USNM 784719. Height 12.7 mm, 
diameter 7.2 mm. 

Locality - Recent, Borburata, Falcon State, 
Venezuela. Living also at other north coast lo- 
;alities from the Paraguana Peninsula to 
Carenero, Miranda State, and on the islands of 
Los Roques, Tortuga and Margarita. As a fossil 
it occurs in the early Miocene Cantaure Forma- 
tion, Paraguana Peninsula, and in the late Plio- 
cene Mare Formation, Cabo Blanco. 

Paratypes - British Museum (Natural His- 
l;ory) and the Natural History Museum Basel. 
Remainder in the collections of the authors. 

Remarks - The material available consists of 
87 specimens from the various localities; it is 
from beach drift only, the animal living, pre- 
sumably, just above high tide level as do other 
members of the family. There is some variation 
in globosity. In Venezuela, Tralia ovula is large- 
ly replaced by T. venezuelana n. sp. only 3 
specimens of the former having been found, two 
at Borburata, Carababo State and one from 
Islas Los Roques (Figs. 4, 5), two localities 
where T. venezuelana also occurs. From the 
Cantaure Formation one juvenile (Fig. 9) and 
one spire have been recovered and from the 
Mare Formation a spire only with pitted 
grooves. T. venezuelana is only the second Re- 
cent species to be reported from the region. Its 
presence at Cantaure and Mare helps to confirm 
the shallow water nature of those faunas. 

Comparisons - The fourth inner lip fold dis- 
tinguishes T. venezuelana from T. ovula and 
from the west coast forms T. payiamensis (C. B. 
Adams, 1852) and T. vanderbilti Schwengel, 
1938. T. ovula, furthermore, lacks the pitted 
spiral grooves of T. venezuelana. The only other 
fossil from the region is T. vetula Woodring, 
1928, from the Pliocene Bowden Formation, 
Jamaica; it also has pitted spiral grooves at the 
top of the whorl, but lacks the fourth, inner lip 
fold and it is smaller and slimmer. 


Abbott, R. T. 1974. American Seashells. Second Edition. 

Van Nostrand Reinhold Company, New York, 663 pages. 
Altena, C. 0. van Regteren. 1975. The marine Mollusca of 

Suriname (Dutch Guiana) Holocene and Recent, Part 3: 

Gastropoda and Cephalopoda. Zoo/. Verhand. No. 139, 

104 pp., 43 Text figs., 11 pis. 
Coomans, H. E. 1958. A survey of the littoral Gastropoda of 

the Netherlands Antilles and other Caribbean Islands. 

Studs. Fauna Curacao, No. 31, pp. 42-111, 16 pis. 

^IG. 1. Melampus (Melampus) coffea (Linnaeus, 1758). Re- 

■ent, Carenero, Miranda State, Venezuela. Height 9.5 mm. 

iiameter 6.0 mm. 

^'IGS. 2-3. Melampus (Pira) monilis (Bruguiere. 1789). Re- 

■ent. Borburata, Carabobo State, Venezuela. 2, height 12.1 

nm. diameter 7.9 mm. 3, whitened, height 8.9 mm, diameter 

).l mm. 

""IGS. 4-5. Tralia ovula (Bruguiere, 1789). 4, Recent. 

uvenile. Borburata, Carabobo State. Venezuela, whitened. 

wight U.6 mm, diameter 2.J, mm. 5, Recent, Islas Los Roques, 

Venezuela, height 10.1 mm., diameter 5.0 mm. 
FIG. 6. Detracia roquesana n. sp., holotype, USNM 781,718, 
height 10.6 mm. diameter 5.8 mm. Recent, Islas Los Roques, 

FIGS. 7-9. Tralia venezuelana n. sp. 7, holotype, USNM 
784719, height 12.7 mm, diameter 7.2 mm. Recent, Bor- 
burata, Carabobo State, Venezuela. 8, paratype, whitened, 
height 7.8 mm, diameter J,.6 mm, ibid. 9, Cantaure Forma- 
tion, Paraguana Peninsula, Venezuela, height .3.8 mm. 
diameter 2.2 mm. 


July 16, 1982 

Vol. 96(3) 

Gibson-Smith, J. and W. Gibson-Smith. 1979. The genus 
Ardnella (Moilusca: Bivalvia) in Venezuela and some 
associated faunas. GEOS No. 24, pp. 11-32, 3 pis. 

Marcus E. and E. Marcus. 1965. On BraziHan supratidal and 
estuarine snails. Bol. Fax:. Fit Cien. Letr. Univ. S. Paulo 

No. 287, Zoologia No. 25, pp. 19-82, 10 pis. 
Olsson, A. A. and T. McGinty. 1958. Recent marine mol- 
lusks from the Caribbean coast of Panama with the de- 
scription of some new genera and species. Bull. Amer. 
Paleont. 39(177):58, 5 pis. 


J. Gibson-Smith and W. Gibson-Smith 

Escuela de Geologia, Minas y Geofisica, Universidad Central de Venezuela, 
Apartado 47.351, Caracas 1041 A, Venezuela 

Stephopoma pennatum Morch, I860, from the Panamic Province and not pre- 
viously knovm as a fossil, has been found in the early Miocene Cantaure Forma- 
tion, Venezuela. The only other species in the region is S. myrakeenae Olsson & 
McGinty, 1958, from the Caribbean adjacent to the exit of the Panama canal. S. 
myrakeenae has not been shovm to differ from S. pennatum and this raises the 
question of whether the former is a Miocene relict, or whether the sessile pennatum 
has reached the Caribbean shore via the canal, attached to the hulls of ships. 

Stephopoma pennatum Morch, 1860, lives in 
shallow water in the Eastern Pacific from 
Nicaragua to Peru. Keen (1971:396:450) places 
the genus in the Vermiculariinae, probably 
because of its turretellid operculum edged with 
bristles. On the other hand, Abbott (1974:101) 
assigns it to the Vermetidae. The most striking 
feature is the protoconch which is neither tur- 
ritellid nor vermetid. It is described by Keen 
(loc. cit.) as follows: "The initial whorls are flat, 
resembling a small Heliacus, lighter-colored 
than the adult shell, studded with radial rows of 
pustules". Olsson & McGinty (1958:35) identified 
a second species, but from the Caribbean coast 
of Panama: Stephopoma myrakeenae. They 
assigned it also to the Vermetidae. 

Stephopoma Morch, 1860, the type of which is 
Vermetu.s roseum Quoy & Gaimard, 1832, from 
the Recent of New Zealand, is not known as a 
fossil. However, from the early Miocene (Burdi- 
galian) Cantaure Formation, Paraguana Penin- 
sula, Venezuela, twelve protoconchs have been 
collected with short lengths of the disjunctly- 
coiled shell attached (Fig. 1). The specimen 
micrographed (SEM) was inadvertently in- 

verted when mounted; but no matter. The 
planorbid protoconch consists of 1 1/4 rapidly 
expanding whorls, the first half-turn smooth, 
followed by the pustulose decoration. The ir- 
regular adult tube, subquadrate to subcircular in 
section, is smooth and glossy within; externally, 
it is closely, longitudinally ribbed, the surface 
roughened by sinuous growth incrementals, 
some of which are scaly, and the longest tube 
measures about 15 mm with a diameter of 2 mm. 
All this is much as in S. pennatum, to which the 
shell is referred. 

The Caribbean 5. myrakeenae was not dif- 
ferentiated from S. pennatum and although it 
was not described as being ribbed, it was said 
that most of the adult tubes were very worn. 
Otherwise, it is indistinguishable and should, 
perhaps, be added to the short list of species oc- 
curring on either side of the Isthmus of Panama 
(Radwin 1969:234, 235; Vermeij 1978:213, 269). 
S. myrakeenae was found at Colon and Bocas del 
Toro Island, some 250 km to the west. The ques- 
tion has to be asked, therefore, whether it is 
pure coincidence that this Miocene relict found 
refuge adjacent to the exit of the Panama canal. 

Vol. 96(3) 

July 16, 1982 


FIG. 1. Stephupoma pennatum Miij-r/i. ISHO. Entiy Miocene 
Cantaure Formation, Venezuela. SEM mirnigrnph (x 55). 
Ventral view of protoconck. 

There is the possibility that it is a case of 
recolonization by pennatum, this sessile animal 
navigating the canal attached to the hulls of 
ships. The collecting dates were 1917, 1920 and 
1953; the canal opening was in 1914, but it is not 
known whether tuyrukeenae was present at 
Bocas del Tore Island on the earlier visits. If 
present only in 1953, just to reach the island 
would require a migration rate of some 6 km per 
annum, assisted by the prevailing currents. It 
might be significant if it were to be found that to 
the east of Colon, against the prevailing cur- 
rents, there was a not too distant limit to the oc- 
currence of rnyrakeenae. It has been reported 
from Payardi Island (Radwin 1969:230), but that 
lies very close to Colon. 

If it is a case of re-colonization, then Sfepho- 
poma joins the list of "paciphile" genera (Wood- 
ring 1966:426). In that case, its extinction in the 
Caribbean could not have been due to the 
change in environment induced by the severing 
of the Atlantic-Pacific connection, an event 
believed to have resulted in many extinctions 

(Woodring loc. cit.). Had it been so, it could not 
have re-colonized. If extinction occurred, there- 
fore, it was more likely due to the drop in sea- 
level and temperature during the Pleistocene 
glaciations (Olsson 1961:7). 


Abbott. R. T. 1974. Ameriran Sea-shells. 2nd ed., Van 
Nostrand Reinhold Co., New York; 663 pp., 4000 -^ text 
figs. 24 pis. 

Keen, A. M. 1971. Sea shells of tropical West Amei-ica: 
marine mollusks from Baja California to Peru, 2nd ed., 
Stanford Univ. Press, Stanford, Calif, i-xiv + 1064 pp., 
ca. 4000 text figs., 22 col. pis. 

Olsson, A. A. 1961. Mollusks of the tropical Eastern Pacific: 
Panamie- Pacific Pelecypoda. PRI, Ithaca, N.Y., 574 pp., 
86 pis. 

Olsson, A. A. and T. L. McGint.v. 1958. Recent marine mol- 
lusks from the Caribbean coast of Panama with the de- 
scription of some new genera and species. Bull. Amer. 
Paleont. 39(177):52, 5 pis. 

Riidwin, G. E. 1969. A Recent molluscan fauna from the 
Caribl>ean coast of Panama. Trans. San Diego Soc. Nat. 
Hist. 15(14):229-236, 1 fig. 

Vermeij, G. .J. 1978. Biogeograpliy and adaptation: patterns 
of life. Harvard Univ. Press, Cambridge, Massachusetts 
and London, England, i-xi -i- 332 pp., 50 text figs. 

Woodring, W. P. 1966. The Panama land bridge as a sea 
barrier. Pine. Amer. Phil Soc. 110(6):425-433, 3 figs. 

Rare and Exotic Spex:imen Shells 
for the discriminating collector 

Free price list 



946 Ralph Avenue 

Brooklyn, New York 11236 USA 

(212) i85-S550 


July 16, 1982 

Vol. 96(3) 


K. E. Hoagland and 

Lehigh University 

Bethlehem, PA 


Academy of Natural Sciences 

Philadelphia, PA 19103 

Mr. Dale Stingley recently forwarded to The 
Nautilus a letter he found in his files dated 
October 14, 1952, from Wesley R. Coe, now 
deceased. The letter contained unpublished 
information on the type of eggs produced by 
Crepidula maculosa Conrad, 1846. Stingley had 
sent two or three specimens of C. maculosa^ col- 
lected at Sanibel Island in May 1952 to Dr. Coe, 
who had already published several papers on 
developmental mode and size of eggs in species 
of the genus Crepidula (Coe, 1949). It was 
Stingley (1952) who recognized that C. maculosa 
was, indeed, a valid species distinct from C. for- 
nicata with which it had been synonymized. 

Coe's remarks to Stingley are quoted below. 
They are significant because of renewed interest 
in the evolutionary ecology of egg type and lar- 
val development in mollusks (Pechenik, 1979; 
Caswell, 1981). The verification of complete 
brooding in Crepidula maculosa compared with 
mixed development (brooding followed by plank- 
totrophy) in C. fomicata, a larger but similar 
species, provides another case of divergent 
modes of egg development in two similar con- 
geners. As found by Gallardo (1977, 1979) for C. 
dilatata and C.fecunda in Chile and as discussed 
by Hoagland (1975, 1977, p. 403) for the genus 
as a whole, the smaller species of Crepidula tend 
to have direct development of a few yolky eggs 
while the larger species release numerous plank- 
totrophic veligers that develop from small eggs. 

Caswell (1981), uninformed of Gallardo (1979), 
erroneously stated that individuals of Crepidula 
dilatata could produce one of two types of eggs: 
those that metamorphose within egg capsules 
and those that hatch into planktonic larvae. No 
known species of Crepidula can produce both 

Wesley R. Coe 


'The name Calyptraeidae Blainville, 1824, is also used for 
Crepidulidae Fleming, 1 S22, but would need to be made a 
nomen conservandum -bd\tor. 

types of eggs. The works of Stingley and Coe on 
C. maculosa confirm this fact and add to the 
foundation of systematics and life history re- 
quired for incorporating Crepidula into models 
of evolutionary ecology. 

Remarks on the Eggs of Crepidula maculosa 
by W. R. Coe, 1952 were: ". . . you may add that 
a superficial examination of the egg cluster will 
show a marked distinction from C fomicata. 
Your collection shows that C. maculosa pro- 
duces at each ovulation about 10 to 12 gelatin- 
ous capsules each containing 8 to 10 large ova 
(0.44 mm in diameter) or about 100 in all. C.for- 
nicata on the contrary deposits 40 to 70 capsules 
each with 120 to 150 small ova (0.17 mm dia- 
meter) or 10,000 to 13,000 at one ovulation. The 
ova of C. maculosa (we now know) are com- 
pletely incubated beneath the parent's foot, 
while those of C. fomicata hatch early into free- 
swimming veligers." 


Caswell, H. 1981. The evolution of "mixed" life histories in 
marine invertebrates and elsewhere. Amcr. Naturalist 

Coe, W. R. 1949. Divergent methods of development in 
morphologically similar species of prosobranch gastro- 
pods. Joun. ofMorphoL 84:383-400. 

Gallardo, C. S. 1977. Two modes of development in the 
morphospecies Crepidula dilatata (Gastropoda: Calyp- 
traeidae) from Southern Chile. Marine Biol. 39:241-251. 

1979. Especies gemelas del genero Crepidula 

en la costa de Chile; una redescripcion de C dilatata 
Lamarck y descripcion de C fecunda n. sp. Studies on 
Neotropical Fauna and Environment 14(4):216-227. 

Hoagland, K. E. 1975. Reproductive strategies and evolu- 
tion in the genus Crepidula (Gastropoda: Calyptraeidae). 
PhD. Diss., Harvard University. 360 pp. 

1977. Systematic review of fossil and recent 

Crepidula and discussion of evolution of the Calyptraei- 
dae. Malacologia 16(2):353-420. 

Pechenik, J. A. 1979. Role of encapsulation in invertebrate 
life histories. Amer. Naturalist 114(6):859-870. 

Stingley, D. V. 1952. Crepidula maculosa Conrad. The 
Nautihis 65:83-85. pi. 2. 


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October 29, 1982 

ISSN 0028-1344 


Jules R. DuBar and Susan S. DuBar 

A Sinistral Specimen of Scaphella from the 

Waccamaw Formation (Early Pleistocene), South Carolina. 

Michael A. Zeto 

Notes on the Freshwater Mussels (Unionidae) of the 
Upper Monongahela River Basin, West Virginia 



Edward M. Stern and Cheryl A. Vander Weit 

Helix aspersa and Polygyra cereolus. Two Gastropods Introduced into Wisconsin . 

Russell J. Barber 

The Indigenous Distribution of Elliptio complanata 

in Ohio: Archaeological Evidence 



C. John Finlay and Danker L. N. Vink 

New Records of Cymatiidae (Gastropoda) in the Western Atlantic . 


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The Occurrence and Spread of the Introduced Asiatic Freshwater Clam, 
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Hermaphroditism, Sexuality and Sex Ratio in the Surf Clam, 
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Billy G. Isom and Robert G. Hudson 

In Vitro Culture of Parasitic Freshwater Mussell Glochidia. 


Becky A. Houck 

Temporal Spacing in the Activity Patterns of Three Hawaiian Shallow- water Octopods. 

Barry Roth and William K. Emerson 

Rediscovery of the Marginellid Gastropod Persicula 

tessellata (Lamarck, 1822) on the Pacific Coast of Panama 

Notices 161 

Recent Deaths. 




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Vol. 96(4) 

October 29, 1982 




Jules R. DuBar' and Susan S. DuBar 

Geoscience Department 

Morehead State University 

Morehead, Kentucky 


A sinistral specimen of the volut id gastropod Scaphella floridana brennmortoni 
Olsson and Petit was collected from the Waccamaw Formation (Early 
Pleistocene) of northeastern South Carolina. This appears to be the first sinistral 
specimen of the genus to be recorded froyn the Neogene of Southeastern United 
States. The associated shallow water marine faunal assetnblage includes 
numerous semi-tropical and tropical species rare elsewhere in the Waccamaw, or 
previously knoum only from late Neogene deposits of southern Florida. 

A sinistral specimen of the gastropod 
Scaphella floridana brennmortoni Olsson and 
Petit was collected from the Waccamaw Forma- 
tion (Early Pleistocene) of Horry County, South 
Carolina (Plate 1, Figures 1, 2). This is the first 
recorded occurrence of a sinistral specimen of 
the genus Scaphella from the Waccamaw, and to 
the authors' knowledge, the first recorded from 
the Neogene (post-Oligocene) of eastern United 
States. At least three sinistral specimens of the 
closely related Scaphella junonia (Lamarck, 
1844) are known from the Recent (Abbott, p. 
244, 1974). 

The sinistral specimen was found in place in 
the basal one foot of the Waccamaw where the 
formation is exposed along the right bank of the 
Intracoastal Waterway, approximately 14.0 
miles northeast of Myrtle Beach, South Carolina 
(Fig. 1). The enclosing sediment is a slightly in- 
durated admixture of very fine to fine quartz 
sand (17% by weight), silt and clay (3% by 
weight), and calcareous shells (80% by weight). 

The lower part of the Waccamaw at this locali- 

FIGS. 1-4. Scaphella floridana brennmortoni Olsson and 
Petit. 196J,. locality WA 56A--2. Waccamaw Formation (Ear- 
ly Pleistocene), Horry County, South Carolina. 1 and 2, 
sinistral specimen (height restored: 69.2 mm): 3 and 4, dex- 
tral specimen (height 91.2 mm). 

\-aj|r •' . 

'New address: Bureau of Economic Geology, University Sta- 
tion, Box X, Austin, TX 78712. 


October 29, 1982 

Vol. 96(4) 


'6 ' r-iT 
10 Mile 


FIG. 1. Map showing locality from which sinistral Scaphella 
specimen was collected. 

ty was deposited below wave base, several miles 
offshore. The sedimentation rate was relatively 
slow, and turbidity was generally very low. 
SaHnity was normal for the open ocean and bot- 
tom water temperatures were in the range of 
those prevailing today off Southern Florida. 

The associated faunal assemblage yielded 30 
dextral specimens oi Scaphella (Aurinia) obttisa 
(Emmons) and 49 dextral specimens of 
Scaphella Jloridana brennmortoni Olsson and 
Petit (Plate 1, figures 3, 4). In addition, the 
assemblage included numerous semi-tropical 
and tropical species rare elsewhere in the Wac- 
camaw or previously known only from late 
Neogene deposits of southern Florida (Table 1). 
The total macrofauna exceeds 250 species 
(DuBar, 1965, 1971). 

Generally the morphological features of the 
sinistral specimen lie well within the observed 
range of variation for the species. It is noted, 
however, that the sutural slope angle is slightly 
greater than that of any associated dextral 

The height of the sinistral specimen is 62.6 
mm, however, approximately 7.0 mm of the 
anterior extremity of the columella has been 
broken away. The original height of 69.2 mm 
places it near the median for all associated 

specimens (15.5 mm to 138.9 mm) from the 
same locality. 

TABLE 1. Invertebrate species associated with sinistral 
speciriieii of Scaphella floridana brenmorloni Olsson and 


Anadara rustica (Tuomey and Holmes) 

Anadara scalarina (Heilprin) 

Antigona rugatina (Heilprin) 

Area wagneriana Dall 

Chama. (Psev-dochama) caloosana Dall 

Glycymeris americana (DeFrance) 

Laevicardium laevigatum wagnerianum 

Olsson and Harbison 
Semele leayia (Dall) 


Aesopns slearnsi (Tyron) 
Callio.stoma willcoxiayium Dall 
Cancellaria venusta Tuomey and Holmes 
Diodora caloosaensis (Dall) 
Diodora carditella Dall 
Fngciolaria cronleyensis Gardner 
Fusinus caloosaensis carolinensis Dall 
Murex (Chicoreus) floridanus (E. Yokes) 
Marex {Facartia) macgintyi (M. Smith) 
Murex (Phyllonotus) globosu.s (Emmons) 
Plerorhytis conradi Dall 
Scaphella floridana brennmortoni 

Olsson and Petit (dextral specimens) 
Scaphella (Aurinia) obtiisa (Emmons) 


Septastraea crassa (Holmes) 


Arbiiciii wacca/naw Cooke 

I'lypeaster crassus Kier 

Encope michelini imperforata Kier 

Rhyricholampus sp. cf. R. ayresi Kier 


Abbott, R. T. 1974. American SeaJiells. 2n(l Ed.. Van 

Nostrand Reinbold Co., N.Y., 663 p. 
DuBar, J. R. 1971. Neogene stratigraphy of the lower 

Coastal Plain of the Carolinas. Atlantic Coastal Plain 

Geo!. Assoc, 13th Ann, Field Conf., Guidebook: 1-128. 
and H. W. C. Furbunch, 1965. The Waccamaw 

Formation (Pliocene?) and its macrofauna, Intracoastal 

Waterway, Horry County, South Carolina. Geologic Notes. 

Div. Geol., South Carolina Development Board, (^ilumbia. 

S. C. 9(l):l-24. 
Olsson, A. A., and Petit, R. E.. 1964. Some Neogene 

Mollusca from Florida and the Carolinas, Bull. Am. 

Paleoiit. 47 (217);.i09-.574; 7 pis. 

Vol. 96(4) 

Oct(.l>er 29, 1982 





Michael A. Zeto 

West Virginia Department of Natural Resources 

Division of Water Resources 

350 North Vance Drive 

Beckley, WV 25801 


During the spring and summer months of 1980, a preliminary survey of the 
freshwater mussels of the Upper Monongahela River Basin was conducted. A total 
of fifteen species of the family Uyiionidae were found in various tributaries of the 
Monongahela River, with only the Asiatic clam, Corbicula fluminea being found 
in the mainstem. of the river. Many of the species recorded represent new records 
for this watershed. 

Information on the extant mussel populations 
in West Virginia is extremely limited at this 
time. Recent studies in the state were done by 
Taylor and Hughart (1981), Morris and Taylor 
(1978), and Taylor (1980). These studies were 
conducted on the Elk, Kanawha, and Ohio 
Rivers, respectively. Bates (1971) attempted a 
statewide survey, which represents the only re- 
cent study performed on the upper Mononga- 
hela River Basin. Earlier studies were per- 
formed on the Monongahela River by Rhoads 
(1899) and Ortmann (1909). However, these 
earlier works were concerned with mussel 
populations of the Monongahela River and 
tributaries in Pennsylvania. 

Study Area 

The Monongahela River is formed by the con- 
fluence of the West Fork and Tygart Valley 
Rivers at Fairmont in Marion County, in north 
central West Virginia. The river meanders 
northward through Marion and Monongalia 
counties over a course of 60.5 kilometers before 
exiting the state into Pennsylvania. 

Site 1 is located on the West Fork River off 
county route 25/3 approximately 1.60 kilo- 
meters northeast of West Milford in Harrison 
County. The river originates in southwestern 
Upshur County and flows in a generally norther- 
ly course through Lewis, Harrison, and Marion 

counties. The river is 166 kilometers long and 
falls at an average of 2.13 meters per kilometer. 

Site 2 is situated near the mouth of Hackers 
Creek at U.S. route 19 bridge approximately 
3.20 kilometers southeast of Goodhope in the 
southern portion of Harrison County. Hackers 
Creek is a tributary of the West Fork River and 
is 37.19 kilometers long. The stream has a 
drainage area of 150 square kilometers over its 
course through Upshur, Lewis, and Harrison 

Site 3 is located on Buffalo Creek off county 
route 1 near Mannington in western Marion 
County. Buffalo Creek is 48.90 kilometers long 
and has a drainage area of 323.75 square 
kilometers. The entire drainage is located within 
Marion County and falls at an average of 7.30 
meters per kilometer. The stream is the largest 
tributary of the Monongahela River in West 
Virginia, with the confluence located in North 

Site 4 is situated on Dunkard Creek off county 
route 7/28, approximately 1.70 kilometers 
northeast of Fentress in northern Monongalia 
County. The stream is nearly 56.45 kilometers 
in length as it meanders and crosses the state 
line six times before it joins the Monongahela 
River in Pennsylvania. The stream has a total 
drainage area of 588 square kilometers of which 
272 are located in West Virginia. 


October 29, 1982 

Vol. 96(4) 

All stream descriptions are taken from the 
West Virginia Department of Natural Re- 
sources Monongahela River Basin Plan (1981). 


The specimens used in this study were col- 
lected by wading the shallow riffles and pools 
and hand picking. Empty mussel shells were 
also collected in this fashion from sand bars and 
the banks of the streams. The shells collected 
appeared to have been recently discarded. Each 
site was visited at least twice during the study 
period in the spring and summer months of 
1980. The Monongahela River was brailed in the 
fall of 1981 at several locations. No Unionid 
mussels were obtained nor were there any signs 
of shells along the river banks. Identification of 
the specimens collected was aided by the use of 
Burch (1975) and were verified by Dr. David 
Stansbery, The Ohio State University. 


A total of fifteen Unionid species were col- 
lected during this study (Table 1). The greatest 
diversity was found in Dunkard Creek (site 4) 
which supports twelve species. The West Fork 
River (site 1) and Hackers Creek (site 2) also had 

TABLE 1. MusseLs of the Upper Monongahela Rii'er Basin, 





2 3 


Anodonta q. grand is (Say. 18291 



Strophitus u. undulatus (Say,18J7) 




Sinpsonaias ambiqua (Sav. 18251 


Lasmiqona costata (Raf..l820) 


X X 


Tritoqonia verrucosa (Raf..l82n) 




/Imblera t^ plicaU (Say. 1317) 


Fuscofiaia flava (naf.. 18201 



PleurobcTO clava (lam.. 1819) 


Pleurobema sintoxia (Raf..I320) 


Elliptio dilauta (l!af.,lS20) 




Ptychobranchus fascioUrls (Raf..l820) 



Obovarla subrotunda (Raf..l020) 


X X 
X X 

Lampsilis radiata luteola (Lam.. 1819) 


Lampsllls ventricosa (Carne5.1B23) 




EpIoblasiM triQuetra (Raf.. 18201 



Corblcula fluminca (Muller.1774) 

at Morqantown 

• also present at tlononqahcla River 

a fairly diverse population. Freshly discarded 
shells oi Corbicula Jlurninea were noted below 
the locks and dam at Morgan town. This species 
was also flourishing in the West Fork River. 

Fusconaia flava represents a new record from 
the West Fork River, as it was not reported by 
Bates (1971) and there are no records at the ma- 
jor museums in the eastern United States (pers. 
comm. Taylor, 1981). 

All of the mussels collected on Hackers Creek 
represent first records, as there are no records 
of collection on this watershed nor were any of 
the species reported by Bates (1971). This state- 
ment would also apply to the three species col- 
lected on Buffalo Creek (site 3). 

Many of the species taken from Dunkard 
Creek are also new records for this watershed in 
West Virginia. The following species were not 
reported by Ortmann (1909) nor Bates (1971): 
Amblema. p. plicata, Simpsonaias ambigua, 
Epioblasma triquetra, Fusconaia flava, Pleuro- 
bema sintoxia. Elliptio dilatata, and Tritogonia 
verrucosa. Other species collected at this site 
had been recorded by Ortmann (1909). 

It should be noted that two of the species col- 
lected appear on the list of Rare and En- 
dangered Mollusks of the United States (USFW, 
1971). These species are Pleurobema clava from 
site 2 and Simpsonaias (formerly Simpsonicon- 
cha) ambigua from site 4. 

Representatives of the mussels collected in 
this survey have been accessioned in The Ohio 
State University Museum of Zoology as voucher 

I would like to extend my great appreciation 
to Dr. David Stansbery and Dr. Ralph Taylor for 
their valuable time and assistance in identifica- 
tion. I would also like to thank the following in- 
dividuals for their assistance in collecting: 
Denzil Courtney, Sheila Kelley, Jack Mumaw, 
Judy Milne Ricketts, and Sheila Zeto. 


Bates, ,1. M. 1971. Mussel Investigations State of West 
Virginia, Part I -Section I. U.S. Bureau of Commercial 
Fisheries, 91 pp. 

Burch, .J. B. 197.5. Freshwater Unionacean C7am,< of North 

Vol. 96(4) 

October 29, 1982 


America. Malacologica! Publications, Hamburg, Michigan. 

l:()4 pp. 
■Morris, J. S., and R. W. Taylor 1978. A survey of the fresh- 
water mussels of the Kanawha River of West Virginia. The 

(.)rtmann, A. E. 1909. Unionidae from an Indian garbage 

heap. The Nautiluf: 23(l):ll-l.o. 
Rhoads, S. N. 1899. On a recent collection of Pennsylvanian 

mollusks from the Ohio River system below Pittsburg. The 

Nautilus 12(12):133-137. 
Taylor, R. W. 1980. A Survey of the Freshwater Mussels of 

the Ohio River from Greenup Locks and Dam to Pitts- 

burgh, Pa. L'.S. Army C'orjis of Engineers, Huntington/ 

Pittsburgh Districts. 71 pp. 
Taylor, R. W.. and R. C. llughart 19X1. The freshwater 

naia<ls of Elk River, West Virginia with a comparison of 

earlier collections. The Ndutihia 95(l):21-25. 
llnited States Fish and Wildlife Service, Department of the 

Interior 1971. Proceedings of a Symposium on Rare and 

Endangered Mollusks (Naiads) of the U.S. 79 pp. 
West Virginia Department of Natural Resources, Water 

Resources Division 1981. Monongahela River Basin Plan. 

414 pp. 


Edward M. Stern and Cheryl A. Vandei Weit 

Department of Biolog'y 

University of Wisconsin 

Stevens Point, Wisconsin 54481 


Two species of land snails. Helix aspersa and Polygyra cereolus, are reported 
for the first time from Wisconsin. Both species were introduced through the com- 
■mercial shipment of plants. 

Numerous individuals have reported the 
spread of both native and exotic mollusks to new 
localities throughout the United States via com- 
mercially shipped plants. Hanna (1966) summa- 
rized molluscan introductions into western 
North America while more recently Dundee 
(1974) catalogued introduced mollusks of east- 
ern North America. Abbott (1950) gave distribu- 
tional maps of introduced land and freshwater 
mollusks, including Helix aspersa. 

During August, 1980, while examining plants 
at a local grocery store, the junior author 
observed a single subadult Helix aspersa Miiller 
crawling on the foliage of the creeping fig, Ficus 
pumila. The store manager indicated that all of 
their plants were purchased from a single sup- 
plier located in Wausau, Wisconsin. It was not 
possible to verify conclusively that the snail had 
been transported on the fig plant and the owner 
of the involved nursery firm noted that occa- 

sionally snails and slugs were observed on 
plants shipped from California, Texas, and 
Florida. Helix aspersa is established and/or has 
been intercepted in the latter two states 
(Dundee, 1974) and a similar introduction of H. 
a.spersa with container grown nursery stock was 
reported in neighboring Michigan (Hanna, 
1969). With the cooperation of the wholesaler, 
the senior author examined the warehouse for 
additional specimens. No H. aspersa were 
found. Unlike H. pomatia Linnaeus, which at 
last report (McClary, 1965) was still established 
near Milwaukee, Wisconsin some 25 years after 
its introduction (Washburn, 1941), there is no 
indication that H. a.'ipersa has become estab- 
lished as a breeding population within central 

However, several individuals of a second 
species, Polygyra cereolus (Muhlfeld), were col- 
lected within the greenhouse. Unlike//, aspersa. 


October 29, 1982 

Vol. 96(4) 

the natural distribution of P. cereolus is rather 
restricted geographically. It inhabits the Florida 
Keys and the rim of the peninsula (Pilsbry, 
1940). Although the spread of P. cereolus may 
not be of economic or regulatory concern and no 
established colonies of H. aspersa are known to 
occur in Wisconsin, they well illustrate the 
dispersal of mollusks is a continuing problem. 

Voucher specimens of both species are de- 
posited in the University of Wisconsin-Stevens 
Point Museum of Natural History. 

Abbott, R. Tucker 1950. Snail Invaders. Natural History 

Dundee, D. S. 1974. Catalog of introduced molluscs of east- 
ern North America (North of Mexico). Sterkiana 55:1-37. 
Hanna, G. D. 1966. Introduced mollusks of western North 

America. Occ. Papers Calif. Acad. Sci. 48:1-108. 
Hanna, M. 1969. Dispersal of Helix aspersa with container 

grown nursery stock. The Nautilus 82:145. 
McClary, A. 1965. Helix pomatia in Wisconsin. The Nautilus 

Pilsbry, H. A. 1940. Land Molluscaof North America (North 

of Mexico). Acad. Nat. Sci. Phila. Monog. 3. Vol. 1, Pt. 2, 

pp. 575-994. 
Washburn, R. G. 1941. A Wisconsin colony oi Helix pomatia. 

The Nautilus 5i:145. 


Russell J. Barber 

Institute for Conservation Archaeology 

Peabody Museum, Harvard University 

Cambridge, MA 02138 

It generally has been concluded that extra-limital records of the freshwater 
mussel, Elliptio complanata, in Ohio resulted from species dispersal allowed by 
the nineteenth century construction of canals. Faunal remains from a prehistoric 
archaeological site suggest that an earlier mechanism of dispersal was in opera- 
tion. The potential valv£ of archaeological evidence for zoogeographical studies is 

The freshwater mussel Elliptio complanata 
(Solander in Lightfoot) is distributed widely in 
the rivers of the Atlantic Seaboard of North 
America, in the rivers and lakes of the Great 
Lakes basin, and in a few rivers flowing into the 
extreme eastern portion of the Gulf of Mexico. 
It generally is excluded, however, from the In- 
terior Basin of the Ohio and Mississippi Valleys 
(Matteson 1948; LaRocque 1967: 171-172). 

In Ohio, Elliptio complanata has been 
reported from two localities: Grand River, 
Ashtabula County (Ortmann 1919: 103) and 

Tuscarawas River, Tuscarawas County (Sterki 
1907: 393). The Grand River is part of the Great 
Lakes drainage system; the Tuscarawas River is 
part of the Ohio River drainage system. The oc- 
currence of the species in the Tuscarawas River, 
presently unconnected with other rivers from 
which Elliptio complanata is known, requires 

Sterki (1907: 393) suggested that the species 
migrated to the Tuscarawas River from Lake 
Erie via the Ohio Canal. While recognizing the 
possibility of "a very complex and rather un- 

Vol. 96(4) 

October 29. 1982 

THE NAl'TI I. rs 131 

likely introduction through drainage changes in 
the river system," LaR<icque(1967: 172) favored 
the explanation of introduction through nine- 
teenth century canals. 

These interpretations of the meager evidence 
lare hampered by limited time depth for distribu- 
tional data. Molluscan remains from prehistoric 
archaeological sites, tyj^ically collected and iden- 
tified during the course of archaeological 
research, can extend that time depth backward 

The Anderson Village site {33-Wa-4) is located 
in Warren County, Ohio, on the banks of the Lit- 
tle Miami River, a tributary of the Ohio River. 
During the years around 1350 A.D., it was oc- 
cupied by Indians of the Fort Ancient Culture. 
Excavation at the site by Patricia S. Essenpreis 
in 1976 disclosed a series of refuse pits 
;"features"), some of which contained several 
hundred freshwater bivalve shells, apparently 
the refuse remaining from human meals. These 
remains subsequently were analyzed and 
reported by Barber (1978). 

The Anderson Village site produced 10 valves 
of Ell iptio complanata, 2 from Feature 10 and 8 
from Feature 11. Seven of the 10 valves were 
whole or nearly so and all included both hinge 
and umbo portions. Preservation of shell was 
good and the identifications were routine and 

These remains of ElUptio complanata might 
be explained in two ways. First, they might be 
claimed to have been imported from the Great 
Lakes drainage basin. The distance involved, 
however, is one of over 100 kilometers and suc- 
cessfully transporting a perishable foodstuff 
such a distance would have been impossible. The 
shells were found with domestic refuse in- 
dicating simple cooking; there was no evidence 

of the use of shells for industrial or otlu'r pur- 

Second, the ElUptio complataita might have 
been living in the Little Miami River during the 
fourteenth century and might have been 
gathered locally. This explanation is consistent 
with present knowledge of human resource ex- 
ploitation and is favored here. 

The Anderson Village site remains, thei'el'ore, 
suggest that the Euro-American canals might 
not have been the mechanism of introduction of 
ElUptio complanata to the Ohio Valley, or at 
least not the earliest mechanism. Complex river 
system changes or even fortuitous transporta- 
tion by mobile animals indeed may have been 

Archaeologists long have sought the advice of 
zoologists in the interpretation of the faunal re- 
mains they encounter. Though largely untap- 
ped, archaeological remains form a data base 
which zoologists profitably could use, especially 
in zoogeographical studies. Hopefully the future 
will demonstrate the mutual benefit of increased 
information exchange between the disciplines. 

Identification of the mussel was confirmed by 
Richard I. Johnson. 


Barl)er, K. J. 197S. Kep<jrt iin iho molluscan fauna from the 
Anderson Village site(33-Wa-4). Report on file, Ohio State 
Historical Society. 

LaKocijue. A. 1967. Pleistocene Mollusca of Ohio. Bulletin. 
Ohio Gi'otoyicat Surety. 62(2). 

Matteson, M. K. 1948. The taxonomic and distril)Utional 
history of the fresh-water mussel Elliptio complanatu.s 
(Dillwyn, 1817). The Nautilus ^l(A):\21-\Z2, 62(1):13-17. 

Ortman, A. E. 1919. A monograph of the Najades of Penn- 
sylvania. Part III: Systematic account of the genera and 
species. Carnegie Museum Mevwirx 8(1). 

Sterki, V. 1907. A preliminary catalogue of the land and 
fresh-water Mollusca of Oliio. Prdceedim/s of the Ohio 
State Academy of Science 4(8):3()7-402. 




5EA SHELLS p. O. Box 83 

Glen Ellen, CA 95442 


!()()K LISTS 

Specimen Shells 

Offering microscopic and miniature (to '/» inch) shells 
from the Florida Keys, unlh accurate locality data. Also 
unsorted grunge: urrite for list. 

Margaret Teskey 


Big Pme Key. Fl. SSOUS 


October 29, 1982 

Vol. 96(4) 


C. John Finlay 

116 Tanglewood Lane 

Newark, Delaware 19711 



Danker L. N. Vink 

Prinsenweg 73 

2242 EC Wassenaar 


Since the coverage of the family Cymatiidae /or the Western Atlantic by Clench 
and Turner in 1957, and subsequently by Abbott in 197^. a number of new records 
in the genera Argobuccinum, Ranella, Charonia, Sassia and Cymatium have 
turned up in the Caribbean marine province. 

Argobuccinum gemmiferum (Euthyme, 1889) 
Cuba- An immature worn specimen was 
taken by Jose Fernandez Milera after a severe 
winter storm in 1952, washed up on a rocky 
shoreline, approximately one mile west of the 
fishing village of Camarioca, Matanzas, not far 
from Varadero Beach, Cuba. 

Brazil-Dr. Luiz Roberto Tostes kindly sent 
us an adult specimen taken by a diver alive on 
sand, near rocks, at a depth of 40-50 metres, 
near Cabo Frio, Ilha dos Pagagaios, State of Rio 
de Janeiro. A total of four specimens were taken 
near Cabo Frio, the largest measuring 126.5 
mm in length, and the smallest 101.4 mm. Two 
of these shells are in the collection of Dr. Tostes, 
one in the collection of the Museu Oceanografico 
de Rio Grande, and the other in the Vink collec- 

FIGS. 3 and 4. Argobuccinum gemmiferum (Euthyme, 1889). 
Jeffrey's Bay. South Africa, Length: 123 mm. 

FIGS. 1 and 2. Argobuccinum gemmiferum (Euthyme, 1889). 
Avar Camarioca., Matanzas Prov., Cuba. Length: 62 mm. 

F'IGS. h anil ti. Argoliuccinum genunifei-um /£."w//iy///< . ISSU). 
Near Cabo Frio. Ilha dos Pagagaios, R. J.. Brazil. Length: 
101 mm. 

Vol. 96(4) 

October 29, 1982 


tion. A fifth specimen from Marica, State of Rio 
de Janeiro, measuring 1 10.0 mm, was tal<en by a 
fisherman in 70-80 metres, and is also in the col- 
lection of Dr. Tostes. 

Ranella olearium (Linne, 1758) 

Brazil-ln June 1973, Matthews, Rios and 
Coelho, described a new Brazilian species of 
Bur^ifi as 5. harcellosi. The new species was con- 
sidered by the authors to be closely related to B. 
multinodosa Bucknill, 1927, found in New Zea- 
land waters. Certain differing characteristics 
however were cited to support the new species, 
the most important of which was the lack of nod- 
ules on the early post nuclear whorls of B. multi- 
nodosa. From a recent examination of the New 
Zealand taxon in the collection of the Delaware 
Museum of Natural History, co-author Finlay 
has been able to determine that the four ex- 
amples at the museum revealed nodules on all of 
the early post nuclear whorls. As only one 
specimen of B. multinodosa was apparently ex- 
amined, it could be assumed that that specimen 
was worn and the nodules eroded. Later, Rios, 
in 1975, considered B. barcellosi as a synonym of 
Ranella olearium. Although not so stated by 
Rios, it would seem that B. multinodosa must 
also appear in the synonomy of olearium.. 

Eight type specimens of B. barcellosi (i.e., 
Ranella olearium) were dredged in depths from 
100 to 280 metres, by the vessels "Mestre 
Jeronimo" and "Kosei Maru" from a muddy sand 
substrate, at stations off Torres, Solidao, Con- 
ceigao and Albardao, all in the State of Rio 
Grande do Sul. One of the paratypes was do- 
nated to the United States National Museum, 
Smithsonian Institution, Washington, D.C. 

Bermuda -In recent years. Jack Lightbourn 
and Arthur Guest of Bermuda, have taken 
several specimens of this species in specially de- 
vised deep water traps, set in depths of 120 to 
140 fathoms (219-256 metres), VU miles off Cas- 
tle Roads, S. E. Bermuda (all shells were dead, 
occupied by hermit crabs.) A specimen of the 
species has been deposited in the collection of 
the Delaware Museum of Natural History. This 
species has a worldwide distribution, occurring 
in the Mediterranean, West Africa, South 
Africa, Australia and New Zealand. 

Charonia rubicunda (Perry, 1811) 

Brazil -In 1981, Rios and Tostes reported the 
presence of Charonia. rubicunda in Brazilian 
waters. A total of 18 records were cited, from 
Itajai, State of Santa Catarina, to Sao Mateus, 
State of Espirito Santo, between 8 and 50 m in 
depth. The majority of the specimens, collected 
in 1976 and 1977, occurred off Cabo Frio, State 
of Rio de Janeiro. Divers reported that the spec- 
imens were taken on a sandy bottom near rocks. 
This species occurs commonly in Australian 

Cymatium pharcidum (Dall, 1889) 

Surinam and Bermuda -DaW described this 
species as new in the "Blake Report" from a 
specimen dredged off Barbados in 82 fathoms 
(150 metres) on a sandy bottom. Clench and 
Turner, 1957, considered the species as a syno- 
nym of Cymatium krebsi Morch, 1877. Abbott, 
1974, suggested that the species might be a 
"high spired, finely beaded form of Cymatium 
vespaceum (Lamarck, 1822) or may be a valid 
species". It is the considered opinion of the 
writers that the species is valid. Dr. E. Gitten- 
berger of the Rijksmuseum van Natuurlijke 
Historie, Leiden, Holland, and D. W. van Veen, 
presently have a publication in press redescrib- 
ing and reporting this species from 104 to 130 m 
off Surinam, some 100 miles north of the 
Marowijne river. They compared their speci- 
mens with the holotype of C. pharcidum. The 
type is illustrated in the forthcoming Compen- 
dium ofSeashells by Abbott and Dance (E. P. 
DuttonCo., 1982). 

Specimens from Bermuda, recently trapped in 
120-140 fathoms (219-256 m) I'A miles off Castle 
Roads have been taken by Jack Lightbourn. 
Specimens from this same source are also in the 
Delaware Museum of Natural History and 
Finlay collections. 

Cymatium trigonum (Gmelin. 1791) 

Venezuela -Th\s species had heretofore been 
known only from the Eastern Atlantic (Cape 
Verde and Senegal, to Benguela (Angola)). J. W. 
and F. Gibson-Smith (1970) reported this species 
washed ashore after a storm in 1967 near 


October 29, 1982 

Vol. 96(4) 

Caniuri Grande, Federal District, on the north 
coast of Venezuela. 

Sassia lewisi Harasewych and Petuch, 1980 

Mexico and Barbados -Specimens of this 
recentl}' described species, were originally 
dredged off Contoy Island, Quintana Roo, Mex- 
ico, in 60-80 metres, and off St. James, Bar- 
bados in 140 metres. 


Abbott, R. Tucker, 1974. American Seashells. 2nd edition. 

Van Nostrand Reinhold, N.Y. 
Clench, William J. and Turner, Ruth D., 1957. The family 

Cvmatiidae in the Western Atlantic. Johnsonia 3(36): 

Finlay, C. John 1978. Review of the genus Bursa in the 

Western Atlantic. The Nautilus 92(4), 147-150. 
Gibson-Smith. J. W. and F. Another species of the genus 

Cymatium swims the Atlantic. Boletin Informativo. Asoc. 

Venezolana de Geologia. Minera y Petroleo. Sept. 1970, 

Vol. 13 No. 9. 
Harasewych, M. G. and Petuch, E. J. Sassia leufisi. a new 

Cymatiid gastropod from the Caribbean Sea. The Nautilus 

Matthews, H. R., Rios, E. C. and Coelho, A. C, 1973. Super- 

familia Tonnacea do Brasil. V-Nova especie de genero 

Bursa Roding, 1798 (Mollusca: Gastropoda) Arq. Cien. 

Mar. 13(l):51-56, Junho 1973, Fortaleza, Ceara, Brasil. 
Rios, E. C. and Tostes, L. R. Nota sobre a presenija de 

Charonia rubicunda em aguas brasileiras (Mollusca, 

Gastropoda, Cymatiidae) Lab. de Biologia Marinha, Acad. 

Brasileira Ciencia, Rio de Janeiro, 1981. 




Robert F. McMahon 

Department of Biology 

The University of Texas at Arlington 

University Box 19498 

Arlington, Texas 76019 

The distribution and dispersal of Corbicula fluminea (MiUler) in North 
America were examined on the basis of all available published reports of new 
populations from 192Jt through 1982. The dispersal of C. fluminea in North 
America has proceeded from two epicenters of artificial introduction, the first a 
result of an introduction from southeast Asia into the no7-thwestern United 
States, and the second from a population in the Ohio River (probably a result of 
introduction of specimens from populations earlier established in western United 
States). The dispersal from these two sites of introduction has been marked by a 
progressive invasion of adjacent drainage systems, high downstream rates of 
dispersal and inability to breach barriers to dispersal such as mountain ranges. 
The very high natural dispersal powers of C. fluminea appear to be associated 
with its invasive nature and have been previously overlooked as the basis for this 
species' extraordinarily rapid invasion of North American fresh waters. 

Corbicula fluminea (Miiller) is a freshwater 
clam (Family Corbiculidae) (Newell, 1969) 
endemic to southeast Asia (Morton, 1979). This 

species was introduced to North America near 
the end of the 19th century. It was first re- 
corded from the northwest corner of the United 

Vol. 96(4) 

October 29, 1982 


States and has since spread rapidly over many 
parts of the United States (Cherry, et nl. 1980a, 
1980b). C. fluminea has now been reported in 35 
of the continental states, the exceptions being 
Montana, Wyoming, North Dakota, South 
Dakota, Nebraska, Kansas, New York, Connec- 
ticut, Rhode Island, New Hampshire, Maine, 
Massachusetts, and Vermont (Cherry, et ai. 
1980a, 1980b). It has also been reported from 
fresh waters in northern Mexico (Taylor, 1981; 
B. Morton, personal communication). 

C. fluminea has become an important member 
of many benthic communities in North Ameri- 
can fresh waters. Its high growth rates and ex- 
traordinary reproductive capacity (Aldridge and 
McMahon, 1978) allow it to reach very high den- 
sities soon after establishment of a new popula- 
tion (McMahon, 1983). 

By sheer weight of numbers C. fluminea has 
become one of the most important aquatic pest 
species in the United States. Its shells accumu- 
late in and obstruct irrigation canals and under- 
ground pipes (Eng, 1979; Fitch, 1953; Ingram, 
1959; Prokopovich, 1969; Prokopovich and 
Hebert, 1965). It enters and fouls municipal 
water treatment facilities (Ingram, 1959; 
Sinclair, 1974; Smith, et ai. 1979). C. fluminea 
also renders dredged river sand and gravel unfit 
for use as aggregation material in cement 
(Sinclair and Isom, 1961, 1963). This species has 
also been reported to outcompete native unionid 
and sphaeriid bivalves, many of which are 
presently endangered (Boozer and Mirkes, 1979; 
Cooper and Johnson, 1980; Fuller and Imlay, 
1976; Gardner, et ai. 1976; van der Schalie. 
1973; Sickel, 1973; Taylor and Hughart, 1981). 
The most serious aspect of C. fluminea as a pest 
species is its ability to foul the service water 
systems and steam condensers of electrical 
generating stations, causing extensive damage 
to equipment and expensive shut-downs for 
manual removal of shells (Boozer and Mirkes, 
1979; Cherry, et a/., 1980a, 1980b; Diaz, 1974; 
Goss and Cain, 1977; Goss, et a/., 1979; Harvey, 
1981; Ingram, 1959; McMahon, 1977; Mattice, 
1979; Sinclair and Isom, 1961, 1963; Smith, et 
'//.. 1979; Thomas and MacKenthum, 1964; 
Thomerson and Myer, 1970). Perhaps of even 
greater concern, C. fluminea has not only been 

reported to foul the steam condensers of nuclear 
power stations, but, also, to enter and obstruct 
the emergency reactor cooling systems of these 
facilities (Parsons, 1980). 

While there have been several recently pub- 
lished accounts of the distribution of C. fluminea 
in the United States (Britton and Morton, 1982; 
Cherry, et a/.. 1980a, 1980b), there is little 
published information regarding the actual pat- 
tern and pace of the spread of this important 
pest species in North American fresh waters. 
Since its probable introduction by Chinese im- 
migrants who may have carried it to North 
America as a conveniently transported food 
item (Britton and Morton, i979; Counts, 1981), 
it has been presumed by the majority of investi- 
gators that the spectacularly rapid spread of 
this species through North American fresh 
waters has been the result of human activities 
including; its use as fish bait (Ingram, 1959); 
utilization by tropical fish hobbyists as an 
aquarium specimen (Abbott. 1975); transport of 
juveniles in the bilge water of pleasure craft or 
as a tourist curiosity (Britton and Morton, 
1979); and, perhaps, unknowingly by fish stock- 
ing programs and with transported river sand 
and gravel used as cement aggregation material 
(Britton and Morton, 1979; Sinclair and Isom, 
1961, 1963). 

While human mediated dispersal of C. flu- 
minea certainly has occurred, investigators ap- 
peared to have generally overlooked this 
species' great inherent natural powers of disper- 
sal which are associated with its invasive habit 
and are an apparent adaptation to its preferred 
disturbed, highly variable, temporally unstable, 
lotic habitats, from which most unionids and 
sphaeriids are excluded (Kraemer, 1979; 
McMahon, 1983). Indeed, the high dispersal 
powers of the genus, Corbicula, are illustrated 
by the fact that it has a recent fossil history in 
North America and by fossil record and amino- 
stratigraphic shell dating evidence that indicate 
that a species of Corbicula has reinvaded the 
fresh waters of southeastern England during 
each of the last three or four interglacial periods 
(Miller, et ai. 1978). 

If the spread of C. fluminea in North America 
was primarily the result of human activity one 


October 29, 1982 

Vol. 96(4) 

would expect that the appearance of new popu- 
lations would have a largely random component, 
with upstream invasion rates being equal to or 
greater than downstream rates, with reports of 
new populations occurring in habitats great dis- 
tances outside the previously known species' 
range and with a marked dispersal of new popu- 
lations across natural barriers, as mountain 
ranges, that bisect drainage systems. On the 
other hand, if the majority of dispersal in this 
species was by natural means then dispersal 
would be characterized by a steady, progressive 
expansion of its range from one closely adjacent 
drainage system to the next, with no anomalous 
reports of high upstream invasion rates or the 
breaching by new populations of natural bar- 
riers to dispersal. 

In order to better understand the pace, pat- 
tern and modes of dispersal of C. fluminea in 

North America all available literature on this 
species was surveyed for reports of new popula- 
tions (primarily in The Nautilus) as part of a 
more extensive review of the biology of this 
species in North America (McMahon, 1983). The 
location of each new population and the date on 
which it was first observed were recorded on an 
outline map of the United States (Fig. 1). This 
information was then utilized in the following 
analysis of the distribution and spread of C. 
fluminea in North American fresh waters. 

The first specimens of C. fluminea found in 
North America were recorded as empty shells in 
Namaino, Vancouver Island, British Columbia 
in 1924 (Counts, 1981). The first living popula- 
tion was discovered near the mouth of the Co- 
lumbia River separating Washington and 
Oregon in 1938 (Burch, 1944) (Fig. 1). From this 
apparent point of original introduction, C. 

FIG. 1. Occurrence and spread o/Corbicula fluminea in North America from 1924 to 1982. Published population records for C. 
fluminea ore indicated by the stars and dates on which they were first obsemed. Probable patterns of dispersal by natural 
rneanx are indicated by solid arrows while those pattenm of dispersal axsociated unth human activity and vectors are indicated 
hy dashed arrows. 

,/ol. 96(4) 

October 29, 1982 


luminea dispersed throughout North American 
fresh waters at an extraordinary rate, primarily 
oy successive invasion of adjacent drainage 
ystems (a pattern characteristic of dispersal by 
aatural means). 

C.fluminea spread southwards in West Coast 
drainage systems from the Columbia River, 
•eaching the San Joaquin River Drainage 
System near San Francisco, California, by 1946 
Ingram, 1948). By 1952 it was recorded from 
Irainage systems in extreme southern Califor- 
lia (Fitch, 1953) (Fig. 1). During this period of 
southward expansion on the West Coast no C. 
luminea populations were reported from east of 
,he Rocky Mountains, which formed an effective 
larrier to its dispersal (Fig. 1). Only after 
•eaching the southern portions of California, did 
l.fluminea spread to the east, probably through 
i network of interconnecting irrigation canals 
nto the Colorado River Drainage System where 
t was first recorded in irrigation canals near 
Phoenix, Arizona, as early as 1956 (Dundee and 
Dundee, 1958). From the Colorado Drainage 
System Corhicula spread to the adjacent 
irainage system of the Rio Grande River, the 
most likely mode of dispersal again being 
through irrigation canal systems, where it was 
first observed in the Rio Grande River near El 
Paso, Texas, in 1964 (Metcalf, 1966) (Fig. 1). C. 
tluminea quickly extended its range to the 
downstream portions of the Rio Grande, being 
reported near the mouth of the river at 
Brownsville, Texas, in 1969 (Murray, 1971). 
From the Rio Grande the range of C. fluminea 
expanded rapidly northwards through several 
closely adjacent Texan river drainage systems 
reaching northeastern Texas by 1972 (Britton 
and Murphy, 1977; Aldridge and McMahon, 
1978; Pool and McCullough, 1979) (Fig. 1). 

While the range of C. fluminea was expanding 
into southern California on the West Coast of 
the United States, a second, unexpected infesta- 
tion of this species was recorded in 1957 in the 
Ohio River, near Paducah, Kentucky (Sinclair 
and Isom, 1961, 1963). This anomalous popula- 
tion was recorded at a time when the range of C. 
fluminea in the western United States had ap- 
parently, extended no further east than the Col- 
orado Drainage System in Arizona, 3100 km 

southwest of Paducah (Fig. 1). As such, it 
almost certainly represents an unnatural exten- 
sion of the range of C. fluminea in North 
America, associated with an, as yet, uniden- 
tified human vect(M-. It is highly unlikely that 
this new infestation was the result of specimens 
being carried to the Ohio River from western 
populations by migratory waterfowl as C. 
fluminea cannot survive in their digestive tracts 
(Dreier, 1977; Thompson and Sparks, 1977). 

This artificially introduced population of C. 
fluminea in the Ohio River formed the epicenter 
of a second major expansion of this species' 
range in the eastern half of the United States 
(Fig. 1). From Paducah it spread rapidly down- 
stream in the Mississippi River, reaching the 
lower portions of the river itself and adjacent 
Gulf Coastal drainage systems extending into 
western Florida as early as 1960 (Dundee and 
Harman, 1963; Schneider, 1967) (Fig. 1). The 
rapid downstream advance oi C. fluminea in the 
Ohio and Mississippi Rivers was almost certain- 
ly associated with the ability of its small juvenile 
stage (~ 0.2 mm in shell length, Aldridge and 
McMahon, 1978) to be passively transported by 
water currents (Goss and Cain, 1977; Goss, et 
al.. 1979; Sickel, 1979; Sinclair, 1964; Sinclair 
and Isom, 1961, 1963; Smith, et al, 1979). As 
did the Rocky Mountains on the West Coast, the 
Appalachian Mountains seemed to have formed 
an effective barrier to the expansion of C. flu- 
minea into the drainage systems of the Atlantic 
seaboard (Fig. 1). Instead, only after it reached 
the lower portions of the Mississippi River, did 
the range of C. fluminea expand greatly in an 
easterly direction, progressing successively 
across the closely adjacent drainage systems of 
the lower elevations of the Southern Coastal 
Plain, extending into extreme southern Florida, 
by 1969 (Clench, 1970). During this period of 
range expansion across the Gulf Coastal States, 
a second anomalous range extension of C. 
tluminea was reported in Lake Overholser, 
Oklahoma, in 1969 (Clench, 1972) (Fig. 1). Speci- 
mens were taken from this lake long before C. 
fluminea had extended into north central Texas 
and either represent an incredibly rapid up- 
stream expansion in the Arkansas River Drain- 
age System, or, as appears more likely, an artifi- 


October 29, 1982 

Vol. 96(4) 

cial introduction by a liuman vector (Fig. 1). 

From the drainage systems of the eastern 
Gulf Coastal Plain, C. Jluminea extended into 
the adjacent fresh waters of the Atlantic 
Coastal States, being reported from the Atlantic 
drainage system of Georgia by 1971 (Fuller and 
Powell, 1973) and reaching as far north as 
Virginia and New Jersey by 1972 (Diaz, 1974; 
Fuller and Powell, 1973). Thereafter, the north- 
ern expansion of C. Jluminea slowed greatly 
with the species still reported to be slowly ex- 
tending its range in New Jersey (Trama, 1982). 
The present northern limit of C. jluminea in the 
eastern United States is marked by a marginal 
population in Lake Erie, Michigan, first re- 
corded in 1980 (Clarke, 1981) (Fig. 1). 

C Jluminea has now probably reached the ex- 
tent of its northern distribution in North 
America. It is apparently excluded from more 
northern fresh waters by its intolerance of low 
winter temperatures. Laboratory studies have 
shown the absolute lower thermal limit of C. 
Jluminea to be 2°C (Mattice and Dye, 1976), a 
minimum temperature exceeded by most bodies 
of water in the higher latitudes of North 
America for several months during the winter. 
Indeed, massive mid-winter mortalities induced 
by unusually cold ambient water temperatures 
(< 2°C) have been reported for a C. Jluminea 
population in the Ohio River near Cincinnati 
(Horning and Keup, 1964). In this regard, it is 
highly significant that populations on the north- 
ern edge of the range of C. Jluminea in the 
eastern United States appear to be restricted to 
and to only survive in areas receiving thermal 
discharges from power stations during winter 
months (Dreier and Tranquilli, 1981; Eckbald, 
1975; Rodgers, et al, 1979; Thomas and 
MacKenthum, 1964). 

The upstream invasion rate of C. Jluminea ap- 
pears to be far slower than downstream rates in 
the major drainage systems in which it has 
become established (Fig. 1). Of particular in- 
terest is the slow upstream expansion of C. 
Jluminea in the Ohio River from its point of 
probable origin near Paducah, Kentucky, 
around 1957, from which it reached the up- 
stream limit of its range in the Ohio River 
Drainage System in the Kanawha River, West 

Virginia, by 1963 (Thomas and MacKenthum, 
1964) (Fig. 1). During the same period this 
species spread downstream a much greater 
distance into Louisiana, Mississippi and 
Alabama (Fig. 1). The upstream dispersal of C. 
Jluminea in the Mississippi River was similarly 
slow, with populations reaching Allamakee 
County, Iowa, in the river proper by 1974 
(Eckbald, 1975) and Lake Sangchris, Illinois, by 
1973 (Dreier and Tranquilli, 1981) (Fig. 1). 

If the majority of dispersal of C. Jluminea in 
the United States was mediated by human vec- 
tors the upstream dispersal rate should be 
similar to the downstream rate. Instead, the 
downstream rate is many times that of the 
upstream rate in most drainage systems, 
presumally the result of passive downstream 
transport of newly released juveniles over 
relatively large distances on water currents. 
However, rapid upstream invasion has been 
recorded for C. Jluminea in the Tennessee 
River, where it was found in the river's upper 
reaches by 1959, only two years after it was 
recorded at the river's mouth in the Ohio River 
in 1957 (Sinclair and Isom, 1961, 1963), in- 
dicating that human activity may have been im- 
plicated in its dispersal in this drainage system. 

It appears that much of the dispersal of C. 
Jluminea in North America has occurred by 
natural means. It can be demonstrated that 
mountain ranges appear to form effective bar- 
riers to the extension of the range of C. flu- 
minea, and that downstream dispersal rates are 
generally much more rapid than upstream rates, 
two factors that would not be characteristic of 
the more random dispersal patterns that should 
be associated with human vectors. Upstream 
dispersal may be the result of transport of speci- 
mens in fish digestive tracts as many species are 
known to feed on smaller (shell length <5.0 mm) 
individuals of C. Jluminea (Britton and Murphy, 
1977; Dreier, 1977; Grantham, 1967; Ingram, 
1959; Rinne, 1974; Sinclair and Isom, 1961, 
1963). A more likely vector for the transport of 
C. Jluminea are wading shore birds and water- 
fowl. Juveniles of C. Jluminea (<5.0 mm shell 
length) have been observed to produce a muci- 
lagenous byssal thread with which they attach 
themselves to sand grains (Kraemer, 1979) and 

Vol. 96(4) 

October 29. ]!t,S2 


particularly to filamentous green algae 
(McMahon. unpulilished observations). Such 
sand grains and especially filamentous green 
algae could easily become attached to or en- 
tangled in the feet of wading birds or ducks 
(many species of which feed on C. fluminea. 
Dreier, 1977; Thompson and Sparks, 1977, 
1978) and be subsecjuently transported into 
upstream habitats. Obviously, it is also highly 
likely that such transport by bird vectors may be 
the major mode of dispersal between closely ad- 
jacent drainage systems and could, therefore, 
along with passive dispersal of juveniles by 
water currents, account for most of the rapid 
range expansion of this species in North 
American freshwaters. In contrast, longer-term 
migratory flights would expose juveniles to 
lethal levels of desiccation and would, therefore, 
prevent transportation across mountain ranges. 

Consequently, it appears that while there have 
been several obvious instances of human 
mediated introductions of C. fluminea into 
, North American freshwater habitats, the ma- 
jority of the dispersal of this species appears to 
have been by natural means from perhaps two 
major epicenters of artificial introduction, one 
in the Northwest corner of the United States 
and a second in the Ohio River. The dispersal of 
C. fluminea from these two sites of introduction 
has been marked by a steady progression of its 
range through adjacent drainage systems, 
higher downstream than upstream invasion 
rates and the obstruction of its dispersal by 
mountain ranges, a pattern of range expansion 
generally associated with natural modes of 

Of apparent high importance in the rapid ex- 
pansion of the range of C. fluminea in North 
America is the passive transport of its juvenile 
stage over large distances on water currents. In 
this regard, human activities in the navigable 
waterways of the United States as dredging and 
canalization not only appear to jeopardize native 
unionid and sphaeriid species (Kraemer, 1979; 
McMahon, 1983) but, also, to optimize condi- 
tions for passive dispersal of C. fluminea on 
water currents. 

Certainly, the extraordinary rate at which C. 
fluminea has spread through North American 

fresh waters (Fig. 1) no longer requires explana- 
tions based solely on human vectors. Rather, it 
may have occurred primarily by natural means 
that are clearly associated with the highly inva- 
sive nature of this species and with the remark- 
able capacities for dispersal, as evidenced in the 
fossil record (Miller, et al. 1979), of freshwater 
members of the genus, Corhicida. as a whole. 

I wish to express my gratitude to Lori 
McDowell and Colette (J'Byrne-McMahon for 
their assistance with the preparation of this 
manuscript. This research was supported by 
grants from the Texas Electric Service Com- 
pany and Organized Research Funds of The 
University of Texas at Arlington to R. F. 


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October 29, 1982 


Rodgers, J. H., Jr., D. S. Cherry, K. L. Dick.son and J. 
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fluminea in the Raritan River. The Nautilus 96:6-8. 


John W. Ropes 

National Marine F'isheries Service 

Northeast Fisheries Center 

Woods Hole Laboratory 

Woods Hole, Massachusetts 02543 

In the surf clam, Spisula solidissima and soft-ahMl clam. Mya arenaria, herma- 
phroditism is an anomalous condition of eery loiv prevalence (ea. 0.13% for surf 
clams and 0.35% for soft-shell clams). No conclusive evidence was found that en- 
xnronmental stress caused the condition or influenced the sex ratio, which inis ap- 
proximately equal for both .^fiecies. 

Accidental functional hermaphroditism in the 
dioecious surf clam, Spisula solidissima 
Dillwyn, was reported l)y Ropes (1968a) from a 

single individual taken from off P'alse Cape, 
North Carolina. He pointed out that this anom- 
aly is rare, in part because the ^tmadal tissues 


October 29, 1982 

Vol. 96(4) 

must be prepared histologically and examined 
microscopically to detect the condition, but 
more importantly because the condition is an 
unusual deviation from normal gametogenesis 
in pelecypods(Pelseneer, 1894; Coe, 1943, 1944; 
Galtsoff, 1961). Its rarity suggests that two 
additional specimens found recently be docu- 

The discovery of accidental functional herma- 
phrodites in species thought to be normally of 
separate sexes (unisexual, dioecious, or gono- 
choristic) has resulted in speculation about its 
cause. A discussion of the condition in surf 
clams and soft-shell clams (My a arenaria) is in- 
cluded relating possible causes to sexuality and 
the sex ratio of the two species. 


The preparation of gonadal tissues for micro- 
scopic examination was by standard histological 
methods outlined in Ropes and Stickney (1965). 


The first hermaphrodite was among 14 dredge 
samples and 132 clams collected from off Point 
Pleasant, New Jersey, during January to 
November 1968 (Fig. 1 A-B). It was found with 
four females and five males sampled on August 
7 from a 68 ft (ca. 20.7 m) depth about 10 n mi 
(ca. 18.5 km) east-southeast of the Manasquan 
Inlet entrance (40°00'N Lat., 73°53'W Long.). 

The second hermaphrodite was among nine 
dredge samples and 87 clams collected during a 
June 20 to July 2, 1969 assessment survey (Fig. 
1 C-F). It was found with nine females and one 
male sampled on June 22 at a 76 ft (ca. 23.2 m) 
depth about 24 n mi (ca. 44.5 km) east of Little 
Egg Inlet (39°35'N Lat., 73°47'W Long.). 

Both specimens were bilateral hermaphro- 
dites, as was the earlier example, and gameto- 
genesis in testicular and ovarian alveoli was 
much like that described for the earlier speci- 
men (Ropes, 1968a). A major difference in the 
clams reported herein is that the gonads also 
contained a trematode parasite of uncertain 
identity (Yancey and Welch, 1968). The parasite 
can effectively castrate infected clams by in- 
filtrating the alveoli. In the hermaphrodites 
uninfected alveoli undergoing apparently nor- 

mal gametogenesis comprised about 20% of the 
gonad; the parasites filled the lumina of remain- 
ing alveoli. 


Hermaphroditism may be a positive evolu- 
tionary response by some bivalve species, per- 
mitting survival under unfavorable environmen- 
tal stress, such as low salinity or fresh water 
conditions (Fretter and Graham, 1964). Purchon 
(1968) postulated that a delicate balance exists 
for the expression of the gonochoristic or her- 
maphroditic state in mollusks, and considered 
the former a more primitive condition. Further, 
he felt that young mollusks have a latent capa- 
city to develop in either state; the transition 
possibly effected by genetic or environmental 

Direct causality between specific intrinsic or 
extrinsic factors and the expression of her- 
maphroditism has not been found. Castration 
from parasites invading the gonad may upset 
the balance between male and female poten- 
tialities, but the mechanism is not understood 
(Noble and Noble, 1961). Malek and Cheng 
(1974) noted that the evidence for true sex 
reversal in mollusks suffering from parasitic 
castration was inconclusive, because the gonia 
for one sex may not be completely destroyed by 
parasitic infection. This was the case in the two 
infected hermaphroditic surf clams from New 
Jersey; cells denoting male and female sex were 
observed in development (Figure 1). The 
parasite was found in other surf clams that 
showed no apparent signs of bisexuality. 

Environmental stress has recently been cited 
as the possible cause of accidental functional 
bisexuality in the soft-shell clam, Mya arenaria 
(Otto, 1973). Ten hermaphroditic clams were 
found in 1,311 examined from low salinity 
(3.0-13.8''/oo) areas of Chesapeake Bay, a 
prevalence rate of 0.76% (Otto, 1973). The 
Chesapeake clams which are near the southern 
limit of their geographic range (Hanks, 1963), 
were seriously affected by dilution of the Bay 
water after a 1972 hurricane (Shaw and 
Hamons, 1974). The hermaphroditic soft-shell 
clams studied by Otto (1973) ranged from 50 to 
70 mm in shell length and were considered 

v'ol. 96(4) 


October 29, 1982 


. 100 ^m 





]0 ^m 

' * 

to tj.f" 

FIG 1 SecUon. of the gonad, ofhennaph nMl<lir .urf clams, Spisula solidissima. M Trst^cular (a) and ova-nan 
(9) alveoli and duct coviairwm, parasites (p); and (B) parasite tissues and spermatogeritc cells m the darn col- 
lected on August 7. 1968. (C) Testicular (^1 and ovarian (9) alveoli and duct contaming parasites (p: (D) 
parasite tissues and spermatogenic cells: (E) oogenic cells; and (F) spermatogemc cells in the rl,u„ collected on 
June 22. 1969. A scale of magnification is in the left hand comer of each photograph. 


October 29, 1982 

Vol. 96(4) 

adults (Otto, personal communication). Coe and 
Turner (1938) found "no evidence of protandry 
or change of sex" in the species, but did find 
three hermaphrodites in over 1,000 examined 
from New Haven, Connecticut, and reported a 
prevalence value of less than 0.5% (Table 1); 
Shaw (1970) found one in 36 from Umpqua Bay, 
Oregon, a prevalence of 2.8%; and Porter (1974) 
found one in 1,785 from Skagit Bay, Washing- 
ton, a prevalence of 0.00056%. Other investiga- 
tors have examined many (ca. 5,600) soft-shell 
clam gonads without finding any hermaphro- 
dites (Ropes and Stickney, 1965; Pfitzenmeyer, 
1965; Shaw, 1962, 1965; Brousseau, 1978; Table 
1). The prevalence of hermaphroditism from all 
these observations is 0.35%. 

Although soft-shell clams are generally con- 
sidered to be a gonochoristic species, Coe and 
Turner (1938) observed that "the differentiated 
young gonad shows some indications of bisex- 
uality" and their figure 17 includes "ovocyte- 
like" cells in the spermary of a young clam. If, in 
fact, the potentialities for both sexes exist in the 
gonads of young soft-shell clams and extrinsic 
conditions cause hermaphroditism, then it can 
be theorized that the population sex ratio might 
also be affected. The sex ratios of gonochoristic 
species are generally thought to approximate 
equality and, thus, the observed frequency in 

samples can be subjected to goodness of fit 

The highest per-sample prevalence (4-8%) oj 
hermaphroditic soft-shell clams was reported bj 
Otto (1973); a sex ratio of 1:0.82 males tc 
females was noted in these samples. No signifi- 
cant differences (P>0.05) in the sex ratio, 
however, were detected in any of the sets ol 
samples. Thus, an imbalance in the sexuality ol 
these clams was not observed. 

The hermaphroditic surf clam found in 196J 
and 1969 were from the environmentallj 
stressed New York Bight area, which has and i; 
being intensively studied to identify th« 
ecological effects of waste disposal practices 
(Gross et al., 1976). Although the impact o: 
these spoils on the surf clam resource in this 
region is not well understood, sublethal effects 
are suspected to have important impact on som( 
or all life stages of marine animals, possibly in 
fluencing survival, normal growth, and othei 
physiological processes related to disease anc 
parasitism (Sindermann, 1976). High incidences 
of chromosome or mitotic irregularities hav( 
been found in developing mackerel eggs fron 
the area (Longwell, 1976). Surf clams are non 
migratory, sedentary, and infaunal creatures 
They cannot avoid the impact of contaminants 

Environmental stress has occurred in th( 

TABLE 1. Observations of henna pli >•< ifli f ism in tht> soJl-shi'lJ clum. Mya arenaria. 


No. clams 

No. hermaphro- 


New Haven, Conn. 
Tred Avon R. , Md. 
Patuxent R. , Md. 
Maine to Mass . 
Umpqua Bay, Oreg. 
ChQ,sapeake Bay 
Skagit Bay, Wash. 
W. Gloucester, Mass. 



Coe S Turner, 1938 


Shaw, 1965 

> 700 

Pfitzenmeyer, 1965 


Ropes 5 Stickney, 1965 



Shaw, 1970 



Otto, 1973 



Porter, 1974 


Brousseau, 1978 

3l. 96(4) 

October 29, 1982 


ea. Anoxic conditions off the New Jersey 
ast in 1976 resulted in the loss of a larjje por- 
)n (144,672 metric tons of meats and 61.5%) of 
e surf clam biomass (Ropes et al. 1979). Ogreii 
d Chess (1969) observed mortalities of surf 
ims and other marine animals related to anox- 

around wrecks and reefs off New Jersey in 


Environmental stress affecting the survival of 

rf clams in the New York Bight might cause 

jtagenesis and sex ratio imbalances. Data 

)m observations on 2,307 gonadal tissues of 

rf clams were available from several sources: 

samples taken during 1962-1965 to study the 

reproductive cycle of the clam in offshore New 
Jersey waters (Ropes, 1968b); and from loca- 
tions off Long Island, NY, to off Virginia, 42 
samples taken during surveys in the spring and 
fall of 1965; 35 samples taken in 1968; and 9 
samjiles taken in 1969 (Table 2). In general, the 
hyjHithesis of an equality in the .sex ratio of surf 
clams can be accepted, based on the combined 
totals and chi-square test results of all but three 
1969 samples. These latter, taken off I'oint 
Pleasant, NJ, are an exception, since the dif- 
ferences in sex were highly significant 
(X^= 10.36; df=2; P>.01)-78.6%' of the clams 
were females; 17.9% were males; and 3.6% her- 

TABLE 2. The sex ratio ofsurfrliniis (Spi.sula S(]li(li.ssiiiia). 

Number of 










Long Island 






























0. 1498 

Pt. Pleasant, NJ 
























2. 1304 





















-) -1 








Cape May-Wildwood, 











36 ^ 















Delmarva Peninsula 







5 Virginia 

























Grand Total 






1/ Includes a hermaphrodite in the total. 


October 29, 1982 

Vol. 96(4) 

niaplirodites. However, the sample size may 
have been too small to make the test statistically 
conclusive (Dixon and Massey, 1957). The 
prevalence of hermaphroditism in all of the 
samples was a low 0.13%; the sex ratio was 
1:1.0645 males to females. 

I wish to thank Drs. Diane J. Brousseau of 
Fairfield University, Fairfield, Connecticut, 
Saul B. Saila of the University of Rhode Island, 
KinjTston, Rhode Island, and Fredric M. Ser- 
chuk of the National Marine Fisheries Service, 
Northeast Fisheries Center, Woods Hole, Mass- 
achusetts, for their critical reviews and helpful 


Brousseau, D. J. 1978. Spawning cycle, fecundity and 
recruitment in a population of Mya arenaria (soft-shell 
clam) from Cape Ann, Massachusetts. U.S. Fish. Bull. 

Coe, W. R., and H. J. Turner, Jr. 1938. Development of the 
gonads and gametes in the soft-shell chim (Mya iirenariiil. 
Jour. Morjih. 62:91-111. 

Coe, W. R. 1943. Se.xual differentiation in mollusks. I. Pole- 
cypods. Quart. Rev. Biol. 18:1.54-l(i4. 

1944. Sexual differentiation in mollusks. II. Gas- 
tropods, Amphineurans, Scaphopods, and Cephalopods. 
Qtiart. Rn: Biol. 19:8r)-97. 

Dixon, W. J., and F. .1. Massey, .Jr. UI.ST. Iiilroi/iirlinn hi 
Statistical Amily.'<is. McGraw-Hill [?ook Co.. Inc., N.Y. 
488 p. 

Fretter, V., and A. (Jraham 1964. Reproduction. In: Wilbur. 
K. M. and C. M. Yonge (eds.) Physiology of Mollusca. 
Academic Press, N. Y. Vol. 1, p. 127-164. 

Galtsoff, P. S. 1961. Physiology' of reproduction in molluscs. 
Amer. Zool. 1:273-289. 

Gross, M. G., R. L. Swan.son, and H. M. Stanford. 1976. 
Man's impact on the middle Atlantic continental shelf and 
the New York Bight- Symposium summary. In: (iross, M. 
G. (ed.) Middle Atlantic Continental Shelf and the New 
YiTk Bight. Amer. Soc. Limnol. Ocfanogr.. Inc.. Spec. 
Symp. Vol. 2, p. 1-13. 

Hanks, R. W. 1963. The soft-shell clam. r..S. Fish and Wihll. 
ScTiK. Circ. 162, 16 p. 

Longwell, A. C. 1976. C'hroniosome mutagenesis in develop- 
ing mackerel eggs sampled from the New York Bight. In: 
Gross, M. G. (ed.). Middle Atlantic Continental Shelf and 
the New York Bight. Amer. Sue. Limnol. Oceanoyr.. Inc.. 
Spec. Symp.. Vol. 2, p. 337-339 (Abstract). 

Maiek, E. A., and T. C. Cheng. 1974. Medical and Economic 
Malacology. Academic Press, N.Y. 398 p. 

Nohle, E. R., and G. A. Noble. 1961. Parasitology. The 
Biology of Animal Parasites. Lea and Febiger, Phila. 
767 p. 

Ogren, L., and ,J. Chess. 1969. A marine kill on New Jersey 
wrecks. Underwater Naturalist. 6(2):4-12. 

Otto, S. V. 1973. Hermaphroditism in two species of 
pelecypod mollusks. Proc. Natl. Shellfish. Assoc. 

Pelseneer. P. 1894. Hermaphroditism in molluscs. Quurl. 
.lour. Micros. Soc. 37:19-46. 

Pfitzenmeyer, H. T. 1965. Annual cycle of gametogenesi.^ ■ ; 
the soft-shell c\am,Mya arenaria. at Solomons. Maryland. 
Chesapeake Sci. 6:.52-.59. 

Porter, R. G. 1974. Reproductive cycle of the soft-shell clam, 
Mya arenaria. at Skagit Bay, Washington. F(sA. Bull. 72: 

Purchon, R. D. 1968. The Biology of the Mollusca. Pergamon 
Press, Oxford. 560 p. 

Ropes, J. W., and A. P. Stickney. 1965. Reproductive cycle 
of Mya arenaria in New England. Biol. Bull. 

Ropes, J. W. 1968a. Hermaphroditism in the surf clam, 
Spisula solidissinia. Proc. Natl. Shellfi.'<h. Assih-. 

1968b. Reproductive cycle of the surf clam. 

Spisula solidissinia. in offshore New Jersey. Biol. Bull. 

Ropes, J. W., A. S. Merrill, S. A. Murawski, S. Chang, and 
C. L. MacKenzie, Jr. 1979. Chap. 11. Impact on clams and 
scallops. Part 1. Field survey assessments. In: Swanson, 
R. L., and C. J. Sindermann (eds.) Oxygen depletion and 
associated mass mortalities in the New York Bight. 
A'0.4.4. Prof Paper 11, p. 263-275. 

Sindermann, C. J. 1976. Effects of coastal pollution on fish 
and fisheries- with particular reference to the Middle 
Atlantic Bight. In: Gross, M. G. (ed.). Middle Atlantic Con 
tinental Shelf and the New York Bight, Amer. Soc. Lim- 
nol. Oceanogr.. Inc.. Spec. Symp.. \'ol. 2. p. 281-301. 

Shaw. W. N. 1962. Seasonal gonadal changes in female sol't- 
shell clams, Mya arenaria. in the Tred Avon Ri\i-i-, 
Maryland. Proc. Natl. Shcllfi.'^h. .4.s.soc. 53:121-132. 

1965. Seasonal gonadal cycle of the male soft- 
shell clam, Mya arenaria, in M,-u-yland. L'.S. Fish Wildl. 
Serv., Spec. Sci. Rep. Fish. 508, 5 p. 

1970. A hermaphroditic soft-shell clam. .Mn" 

arenaria. from IJmpqua Bay. Oregon. Res. Rcpts.. Fi.^h. 

Comm. Oregon. 2:100-102. 
Shaw. W. N. and F. Hamons 1974. The present status of tlic 

soft-shell clam in Maryland. Proc. Natl. Shellfish. Asso,: 

Yancey. R. M.. and W. K. Welch 19(;s. Tlu' Atlantic coasi 

surf clam- with ,a partial liililiivgra|)hy. r..S. Fish Wihll. 

Sere. Circ. 288, 14 pp. 

Vol. 96(4) 

October 29, 1982 



Billy G. Isom 

Division of Water Resources 

Office of Natural Resources 

Tennessee Valley Authority 

Muscle Shoals, AL 3566o" 


Robert G. Hudson 

Associate Professor of Biology 

Department of Biology 

Presbyterian College 

Clinton, SC 29325 


Ellis and Ellis (1926 and 1930) reported fnnusformation of fresh water mussel 
glochidia in vitro culture. However, their methodology was never published. This 
report gives a new method for the in vitro culture of mussel glochidia to juveniles 
rather than by their natural fish host encystmmt. The medium consists of 
physiological salts, amino acids, glucose, vitamins, antibiotics, and a nonspecific 
component offish blood plasma. The relative concentration offish plasma re- 
quired for optimum, results was S3 percent. In vitro culture may prove very 
beneficial in reestablishing the populations of endangered mussels, management 
of species used by the pearl culture industry, and culturing of stocks for bioassays, 
genetic studies, or other uses. 

Application of in vitro culture for the purpose 
3f sustaining endangered mussels of commercial 
mportance is obvious, since usually an unknown 
'ish host is required for larval development in 
lature. The female mussel has her eggs fertil- 
zed by sperm cells which are released into the 
vater by a male mussel and travel into the 
'emale with water taken in during filter feeding. 
The fertilized eggs, contained in the female's 
jills, develop into simple glochidia, or larvae, 
■onsisting primarily of two shells, mantle cells, 
ind one adductor muscle. The glochidia are 
•eleased and encyst in a specific fish tissue, 
isually the gills. Development of glochidia into 
uveniles (transformation) in an artificial 
nedium would help endangered mussels, since 
he fish host required for most of these species is 
mknown and would be difficult to determine, 
^he fish host of some species may even be extir- 
lated. Use of in vitro culture could sustain a 
pecies until its fish host could be found or the 
tbsence of a host availability determined. Even 
nussels with a known fish host could be reared 
or commercial or scientific purposes. 

Freshwater mussels have been used commer- 
ially for manufacturing buttons and presently 
is nuclei for cultured marine pearls. J'hey are 

also currently being used for the culture of 
freshwater pearls in the United States. Over- 
harvesting as a result of demand for shells has 
contributed significantly to the decline of 
freshwater mussel resources in the United 
States over the past three decades. Historically, 
over 500 of the more than 1000 world species of 
freshwater mussels occurred in the United 

8n-211-200X LIGUHIA RECTA 

FK;. 1. Cliii-lniliiil n}icU of tkrj're.'ih water miissti l.iuuniia 
recta (Iciiijtii JfiO \ini). 


October 29, 1982 

Vol. 96(4) 

States. Many species are now thought to be ex- 
tinct; others are listed as threatened or en- 
dangered by individual states or the United 
States Government. 

Ellis and Ellis reported in 1926 that they had 
successfully obtained transformation of parasi- 
tic glochidia of freshwater mussels in physio- 
logical nutrient solutions. However, neither the 
composition of the media nor the process used 
was ever published. In addition, unlike the pre- 
sent study, Ellis and Ellis excised the glochidia 
they used from a known fish host, an action 
which undoubtedly contributed to their reported 

Composition of our artificial medium include 
salts which are modified from the "unionid 
Ringers" solution proposed by Ellis et al. (1930) 
and are shown in Table 1. The essential modifi- 
cation of the solution includes the deletion of 
K^HPO. and the addition of 2.2 gm NaHCOj per 
1000 milliliters of solution. The NaHCOj was re- 
quired in order to regulate the pH with varying 
atmospheric CO2 concentrations. 

TABLE 1. Sdlt.'i contained in stock artificial glochidial 








The amino acids in the artificial medium are 
the same as those used by Eagle (1959) for cell 
and tissue cultures with the exception of the ad- 
dition of taurine and ornithine which are consti- 
tuents of fish blood. The essential amino acids 
are shown in Table 2, the nonessential amino 
acids in Table 3. 

The vitamins present in the complex artificial 
growth medium, the same as those used by 
Eagle (1959) for cell and tissue cultures, are 
shown below in Table 4. The antibiotics and 
antimycotic used in the artificial medium are 
listed in Table 5; other compounds are shown in 
Table 6. 

Plasma was obtained from fish blood which 

TABLE 2. Essential amino acids contained in stock artifi- 
cial gluchiiiial medium. 


TABLE 3. Nonessential amino acids contained in stock ar- 
tificial glochidial medium. 


m pound 


■ arginine 


■ cystine 


■ histidine 


■ isoleucine 


■ leucine 


- lysine 


■ methionine 


■ phenylalanine 


■ threonine 


■ tryptophane 


■ tyrosine 


■ valine 


L - alanine 

L - aspara^ne 

L - aspartic acid 


L - glutamic acid 

L ■ proline 

L - serine 


L - ornithine 

(mg/L I 



TABLE 4. Vitamins contained in stock artificial glochidial 


choline chloride 

folic acid 



calcium pantohthenate 





was removed from the fish by heart puncture 
with a sterile heparinized syringe (the syringe 
chamber was coated with a sodium heparin solu- 

Vol. 96(4) 

October 29, 1982 

TABLE 5. The antibiotics and antimycotic contained in 
stock artificial glochidial medium. 



Gentamicin sulfate 


Amphotericin B 


100 uglml 
100 f/g/ml 
100 i^glm] 

5 ng/ml 

TABLE 6. Other compounds contained in stork artificial 
glochidial medium. 



Phenol red (optional) 


tion of 1000 U/ml) having an 18 gauge needle. 
The blood was centrifuged at 1000 rpm in a 
refrigerated centrifuge for 10 minutes followed 
by 3000 rpm for 10 minutes, decanted into 
sterile centrifuge tubes, and centrifuged for 
another 10 minutes. The plasma was removed 
by aspiration, then frozen and subsequently 
sterile filtered. The antibiotics and antimycotic 
were added to the plasma in the same concen- 
tration as shown in Table 5, at time of use. 

Glochidia from gravid female mussels were 
obtained by utilizing sterile surgical instruments 
to remove the gills containing glochidia, then 
the gills were placed in sterile deionized water. 
rhe glochidia were excised from the gills by sec- 
tioning and then swirling the gills in the deio- 
aized water. 

The glochidia were washed several times with 
ieionized water by decanting or aspirating the 
A'ater following swirling of the beaker contents. 
The healthy glochidia settled, allowing the 
-emoval of tissue debris, dead glochidia, 
jacteria, and protozoa. The glochidia were left 
n the last rinse water until they were removed 
vith a Pasteur pipette and placed in the growth 
nedium. A representative scanning electron 
photomicrograph of a glochidium of Ligumia 
•ecta (Lamarck, 1819) is shown in Figure 1. 

Different components and concentrations of 
he growth medium were tested to determine 
vhich combination would stimulate and best 



FIG. 2. Transformed juvenile of the freshwater mussel 
Ligumia recta, 39 days old, in lake water. Note the pro- 
truding ciliated foot. 

FIG. 3. Growth of juvenile Ligumia recta, 75 days in 
culture, 13 days in water (dark field), a total of 28 days. End 
view showing protruding foot in center. 

FIG. 4. Growth of juvenile Lampsilis ovata, 22 days culture, 
9 days in lake water (dark field), a total of 31 days old. 


October 29, 1982 

Vol. 96(4) 

support glochidial transformation. Nuniorical 
comparisons of different cultures were difficult 
to evaluate because the varying stages of devel- 
opment in, as well as the number of, glochidia. 
The presence of bacteria in initial cultures in 
1981 further confounded the accuracy of numer- 
ical culture evaluations. Many cultures failed to 
yield fully transformed glochidia because of 
bacterial infections, even though most glochidia 
had good initial development prior to their con- 
tamination. Experiments in 1982 with the new- 
antibiotics shown in Table 5 have been much 
more successful. 

Only two different salt solutions, unionid 
Ringers (Table 1) and Earles' balanced salt solu- 
tion (Earle, 1943), were compared in experi- 
ments. Glochidia transformed in both salts, even 
though the amino acids used in conjunction w ith 
each varied slightly. Because parasitic glochidia 
are surrounded by fish cells, and since Earles' is 
the salt solution most commonly used in fish cell 
culture medium (Wolf and Quimby, 1969), glo- 
chidal adaptation for life in Earles' salt concen- 
trations as well as in unionid Ringers would be 

Different groups of amino acids were tested in 
cultures to get transformation, but elimination 
of nonessential, single, amino acids has not been 
attempted to date. In earlier experiments with 
the medium being 20 percent artificial, the 
amino acid content seemed less important, prob- 
ably because the 80 percent plasma had the nec- 
essary concentrations of amino acids. The artifi- 
cial growth medium was formulated to include 
all free amino acids which had been found in 
three species of catfish (Iddlurus furcates, I. 
punctatus. and Pylodictis olivaris) (Johnson, 
1971). All of the amino acids which had been 
found in fish blood could also be found in equal 
or higher concentrations in Eagles' essential 
and nonessential amino acids if taurine and 
L-ornithine were added. The only amino acid 
found in Eagles' amino acid group which was 
not found in the catfish blood was L-trypto- 
phane. Since Eagles' amino acids could be pur- 
chased premixed, this combination was used 
along with taurine and L-ornithine in the suc- 
cessful in vitro medium. Medium 199, which also 
has 21 amino acids (Morgan et al., 1950), was 

sufficient for glochidial transformation. Medium 
199 has the same 12 essential amino acids as 
Eagles' medium; however, medium 199 is lack- 
ing 4 of the 9 nonessential amino acids which 
had been used. This would indicate that these 
four (L-alanine, L-asparagine, L-ornithine, and 
taurine) were not necessary for glochidial 
development. Even though development occur- 
red in both media, the development rate was 
faster in Eagles' amino acids with taurine and 
L-ornithine than in medium 199. 

Fish blood plasma was found to be necessary 
to stimulate development in all mussel species 
tested. Other massive protein components, 
specifically fetal bovine serum, bovine serum, 
and lobster hemolymph, were tested and found 
unsuccessful. Although earlier tests indicated a 
positive correlation between percent transfor- 
mation and percent fish blood plasma in the 
medium, 33-1/3 percent blood plasma was 
adopted in our standard medium as producing 
satisfactory yields at a more economical level. 

Earlier cultures of Legumia recta were placed 
in unfiltered plasma which contained several 
blood cells. First observations seemed to in- 
dicate that development was enhanced by the 
presence of these cells. The glochidia were held 
in a matrix of cells and fibrin on the dish and 
developed rapidly. Subsequent media changes 
were made using filtered plasma to prevent a 
buildup of cells in the culture. Conversely, later 
cultures with Fusconaia ebena (Lea, 1831) did 
not show any indication of enhancement by the 
presence of these cells. In fact, cultures with un- 
filtered plasma suffered a higher rate of con- 
tamination which resulted in a lower yield. More 
work is needed in this area, but the presence of 
the fish cells does not appear to be necessary for 
glochidial development. 

The pH of the medium varied from ca. 7.3 to 
8. 1; however, the lower pH promoted best trans- 
formation in at least one species. Cultures of 
Fui^conaia ebena developed much better in a pH 
range of 7.3 to 7.4 rather than 7.8 to 8.0. For ex- 
ample, in eight different cultures of F. ebena. 16 
of 26 culture dishes contained transformed 
juveniles after 18 days at a pH of ca. 7.3, where- 
as none of an identical set of 26 cultures were 
alive at 11 days when cultured at a pH of ca. 7.9. 


(MoIht 29, 1982 


Bacterial infection was present in cultuix's at 
both pH levels and was responsible for much vi' 
this loss; however, many of the higher pH 
cultures died even when bacterial action was 
minimal, while infections in the lower pH 
cultures did not result in death of the entire 
culture. The lower pH level appeared to either 
make it easier to control bacterial contamina- 
tion or minimize the detrimental effects of the 
bacteria. Subsequent experiments have corro- 
borated that a pH of 7.3 is optimum. 

The use of antibiotics and their concentrations 
varied with the presence or absence of bacteria 
in the medium. €arbenicillin, usually added to 
yield lOU i^glm\, can be increased to as high as 
500 ^iglm\ without apparent inhibition to the 
glochidial development. Gentamicin sulfate and 
Rifampin can also be increased from lOU f.ig/ml 
to 500 figlml safely, and the antimycotic, Am- 
photericin B (Fungizone) from 5 to 25 Mg/'ii'- 
These antibiotics and antimycotic, along with 
rinsing and changing infected glochidia to new 
media and dishes, usually would control con- 
tamination successfully. Older glochidia could 
withstand infection much longer than younger 

The medium was kept at 23°C but cultures 
were successful, to a lesser extent, even at 
28°C. Higher temperature did reduce the yield, 
as shown by the comparison of glochidia of 
Ligurnia recta which developed in 23°C and 
28°C. Using six culture dishes grown in 60 per- 
cent plasma, three grown in 23°C had an 
average yield of advanced glochidial develop- 
ment equal to 48.8 percent (Sx = 8.7), while 
those grown at 28°C averaged 18.8 percent 
(Sx = 3.3). A transformed glochidium of Ligurnia 
recta is shown in Figure 2. 

During the spawning season of 1982, these 
variables have been refined to a greater degree, 
resulting in successful mass culturing of 
glochidia (some cultures averaging as high as 80 
percent transformation). These successes will 
provide opportunity for conservation of many 
mussel species including endangered species, 
and the management of others. Kfficacy of the 
process to date includes transformation of 
glochidia of six genera and species representing 
two subfamilies of Unionidae, the Unioninae 
and Lampsilinae. 

A growth stage of a juvenile Ligmia recta is 
shown in Figure 3 and that of a Lampsilis ovata 
in Figure 4. 


Eagle, H. 1959. Amino acid metabolism in mammalian coll 
cultures. Science 130:432-437. 

Earle, W. R. 1943. Production of malignancy in vitro. IV. 
The mouse fibroblast cultures and changes seen in the liv- 
ing cells. Jour. National Cancer Institute 4:165-212. 

Ellis. M. M. and M. D. Ellis. 1926. Growth and transforma- 
tion of parasitic glochidia in physiological nutrient solu- 
tions. Science 54(1667):579-58(l. 

Ellis, M. M., A. D. Merrick and M. 1). f:ilis. 1930. The blood 
of North American freshwater mussels under normal and 
adverse conditions. Bulletin of the Bureau of Fisheries 
56, Bur, of Fisheries Doc. No. 1097:509-.542. 

.Icihnson, C. L. Unpublished. U.S. Dept. of the Interior, 
Bureau of Sport Fisheries and Wildlife, Cook, Wash., 
1971. In: Biology Data Book. P. L. Altman and D. S. 
Dittmer (eds.). Federation of American Societies for 
Experimental Biology. Bethesda, Maryland, 1974. 

Morgan, J. F., H. J. Morton and R. C. Parker. 1950. Nutri- 
tion of animal cells in tissue culture. I. Initial studies on 
a synthetic medium. Proc. Soc. Exper. Biol. Medicine 

Wfilf. K. and M. C. Quimby. 1969. Fish cells in tissue culture. 
//(. Fish Physiology. W. S. Hoar and D. J. Randall (eds.). 
\(il. Ill, Academic Press. New York. pp. 253-301. 

Recent Deaths 

We regret to announce the death of our dear 
friend, Katherine Van Winkle Palmer, former 
Director of the Paleontological Research Insti- 
tution, of Sept. 12, 1982. at the age of 87. Con- 
tributions to her memorial fund for P.R.I, may 
be sent to Dr. Raymond Van Houtte, Tompkins 
County Trust Co., Ithaca, NY 14850. An 
obituary is in prepartion. 


The Eighth International Malacological Con- 
gress, sponsored by the Unitas Malacologica, 
will be held in Budapest, Hungary, in 1983, from 
August 29 - September 3. Further information 
may be obtained by writing Dr. Laszlo Pinter, 
Natural History Museum, Baross u. 13, H-1088 
Budapest, Hungary. 


October 29, 1982 

Vol. 96(4) 


Becky A. Houck 

Dept. of Physical and Life Sciences 

University of Portland 

5000 N. Willamette Boulevard 

Portland, OR 97203 

Th-ee species of Haw ana h, octopods show distinct iocoinotor activity rhythms in 
the laboratory and in the field. The periods of peak activity are species-specific 
even in isolation. This temporal spacing ayid differences in preferred 
microhabitats during periods of inactivity ynay limit interspecific competition 
for food and home sites and reduce the incidence of predation. Inactive Octopus 
ornatus superficially behave like vertebrates in sleep. 

Animals occupying similar niches in the same 
geographical area may reduce competition by a 
variety of spacing mechanisms. Three species of 
shallow-water octopods can be collected on the 
same fringing reef flats off the south shore of 
Oahu, Hawaii, throughout the year. Data are 
presented indicating that these three species 
show temporal spacing in locomotor activity 
patterns. Observations in the field and in 
aquaria also suggest utilization of different 
microhabitats on the reef flat for use as refuges 
during periods of inactivity. 


Animals were collected by netting during low 
tides on shallow reef flats off the south shore of 
Oahu, Hawaii. 0. cyanea Gray was collected 
before dusk and transported to the Hawaii In- 
stitute of Marine Biology, Kaneohe Bay, Oahu. 
Specimens of 0. ornatus Gould and the "cres- 
cent octopus," a small undescribed Hawaiian 
octopod (Houck, 1977; reference specimens: 
Department of Invertebrate Zoology, Santa 
Barbara Museum of Natural History) were col- 
lected at night and maintained at the Bekesy 
Laboratory of Neurobiology, University of 

Locomotor activity rhythms of four (). orna- 
tus and twelve crescent otopuses were moni- 
tored in isolation under controlled laboratory 
conditions for periods of one to six months per 

animal. Locomotor acitivity rhythms of each 
animal were recorded under a variety of light- 
dark regimes. Only typical activity patterns 
under light-dark conditions similar to timing of 
field illumination are reported here. An 
ultrasonic activity monitor allowed continuous 
recording of all movement of an animal within a 
light-tight aquarium. Data were read by a DEC 
PDP 11/10 computer and stored on cassette 
tape for later analysis. A complete description 
of the monitoring system is provided elsewhere 
(Akaka and Houck, 1980). The animal within the 
aquarium was exposed to an artifical light il- 
lumination with light intensity of 7.5 watts/m^ 
an average value at 0630 hours in a typical oc- 
topus habitat (Franzisket, 1969). Hourly activity 
values were computed as a percentage of the 
mean hourly activity for a monitoring run. 

The larger size of specimens of 0. cyanea did 
not permit use of the ultrasonic monitoring 
system since the animals were too large for the 
aquaria. Activity of 0. cyanea was recorded by 
providing the animal with a single cinderblock 
"refuge" within a 6 foot trough functioning as 
part of an open sea water system at the Hawaii 
Institute of Marine Biology. The cinderblock 
"refuge" was equipped with a trap door bottom 
that was activated when the animal retreated in- 
to the depression of the block. A multichannel 
event recorder documented time spent inactive 
within the refuge. Hourly activity values were 

Vol. 96(4) 

Ort()l)er 29, 1982 


calculated as a percentage of the mean hdurly 
activity, with time spent out of the refuge con- 
sidered activity. Specimens of 0. cyanea were 
exposed to a diffused natural light regime. 


0. cyanea is reported to be active during the 
day and inactive at night (Van Heukelem, 1966; 
Yarnall, 1969) while 0. omatus is considered by 
local fisherman to be night active. No previous 
data on locomotor activity or behavior of the 
crescent octopus have been published. The loco- 
motor activity rhythms demonstrated in this 

study were species-specific in pattern, and peak 
activity of each species was distinct from {)eak 
activity of the other two. 

In this as in previously published studies, O. 
cyanea showed diurnal activity. Maximum loco- 
motor activity, as measured by time spent out of 
the prepared "refuge", occurred just after dawn 
at 7 am (activity from 0600 to 0700 hours) and in 
a broad peak between 1300 and 1800 hours. Ac- 
tivity was maintained throughout daylight 
hours. Van Heukelem (1966) and Yarnall (1969) 
reported 0. cyanea to be diurnally active, with 
crepuscular peaks in activity. This diurnal pat- 







300 1 T 



0. cyane a 
0. ornatus 
crescent octopus 


Time (Hrs.) 

FIG. 1. Locomotor activity patterns for three species of shallow water Hawaiian octopods. Octopus ornatus 
record is based on. 6i continuous hourly values. 0. cyanea record is based on 21, continuous hourly values, and 
the "crescent octopus" record is based on 91 continuous hourly values. 


October 29, 1982 

Vol. 96(4) 

tern is confirmed in tiie record of locomotor ac- 
tivity seen in Fig. 1. 

0. oniatu.'^ demonstrated a nocturnal pattern 
of overall activity in the aquarium as recorded 
by the ultrasonic monitoring system. With a 
light regime of twelve hours of light and twelve 
hours of darkness, peak activity occurred in the 
middle of the lights off period. A dark period 
from 6 pm (1800 hours) to 6 am corresponded to 
natural conditions on the reefs in Hawaii (US 
Department of Commerce, 1944); under this 
light regime, 0. ornatus was maximally active 
from 2100 hours to 0200 hours. 

The crescent octopus also demonstrated noc- 
turnal activity in the aquarium as recorded by 
the ultrasonic monitor, but maximum activity of 
the crescent octopus occurred during those 
hours of darkness when 0. ornatu>^ showed 
reduced locomotor activity. Under the light 
regime in which the lights off period extended 
from 1800 hours to 0600 hours, the crescent oc- 
topus was most active in the hour immediately 
after artifical dusk (1800 to 1900 hours) and im- 
mediately before dawn (0500 to 0600 hour-s). 

Periods of inactivity in all three species were 
characterized by an almost complete lack of 
movement, other than regular mantle respira- 
tory movements. The ultrasonic monitor was es- 
pecially useful for examination of these periods 
of inactivity, since it recorded even small twitch- 
ing of a single arm. Numerous observations 
were made of 0. or-ymtus in the aquarium during 
the inactive period. During these periods, the 
octopus was found at the bottom of the tank 
with its eyelids closed, its arm wrapped about its 
head and mantle, a white translucent color to its 
skin that was quite distinct from the bright red 
and white banded appearance of an active ani- 
mal of this species. Respiratory movements 
were regular, and sensory perception was suffi- 
ciently suppressed for the author to be able to 
move a small net about the tank vigorously with- 
out the animal responding in any way. When the 
animal was touched, it reacted violently with 
what could be called a startled reaction. It 
opened the eyelids, rapidly spread its arms and 
frequently expelled water through its funnel at 
the source of the intrusion. This behavior super- 
ficially resembles sleep in vertebrates during 

periods of inactivity. Other marine inverte- 
brates show periods of heightened and reduced 
locomotor activity (DeCoursey, 1976) but no 
other marine invertebrate has been reported to 
exhibit such pronounced sleeplike behavior. 

The patterns of locomotor activity demon- 
strated in aquaria by the three species were 
reflected in observations in the field. When div- 
ing on the reefs of Oahu during the day, 0. 
cyanea was the only species commonly sited. At 
night it was unusual to find this species on the 
reef. Yarnall (1969) and Van Heukelem (1966) 
report that 0. cyanea retreats to its den at 
night, often blocking the entrance with stones. 
0. oniutua was never observed by the author on 
the reef during the day, and the crescent octo- 
pus was rarely seen. 0. ornatus was most easily 
collected after 9 pm (2100 hours); when low 
tides occurred in the middle the night, 0. or- 
nntu^ was frequently seen fully exposed and 
moving freely across the reef flat or on sand and 
gravel substrates. The crescent octopus was 
routinely collected after dusk in shallow tide 
pools where the animals positioned themselves 
under rock ledges at or near the air-water inter- 

Although all three species of octopods can be 
found within 100 feet of each other on a shallow 
reef flat, there are differences in the substrates 
of the inicroenvironments favored by each 
species. 0. ornatus cannot be observed on the 
reef during inactive periods. When provided 
with a gravel or coral substrate in an aquarium, 
0. 07~n.a,tus will burrow under the gravel and 
disappear from sight during the lights on period. 
The crescent octopus, as mentioned above, fre- 
quents the undersurface of ledges at the aii-- 
water interface in tidepools. In an aquarium, the 
crescent octopus assumes a position at this in- 
terface, often with several arms extended over 
its head, the suckers resting at the water sui-- 
face. 0. cyanea has been located in sites under 
large coral boulders or holes at the base of con- 
crete walls (Van Heukelem, 1966; Yarnall, 
1969). Yarnall reports use of the same "refuge" 
by several 0. cyanea in succession. In an 
aciuarium, (). cyanea will readily inhabit a ccm- 
Crete block or a length of PVC tubing. 

Vol. !)fi(4) 

October 2i). 1982 



Octopods demonstrate a variety of locomotor 
activity rhythms in the aquarium and in the field 
(Woods, i965; Van Heukelem, 1966, 1976; 
Altman, 1967; Yarnall, 1969; Kayes, 1974; 
Houck, 1977). The three species of Hawaiian oc- 
topods in this study show species-specific tem- 
poral spacing in locomotor activity rhythms. 
This s])acing may influence the diet of each 
species. In the laboratory, all three species will 
readily accept much the same variety of grapsid 
crabs and shrimp species as food. Crustaceans 
show locomotor activity patterns of their own, 
often entrained by photoperiod and tidal 
rhythms (DeCoursey, 1976). In the field, hunt- 
ing activity by an octopus at a specific time of 
day may restrict the types of crustaceans eaten, 
reducing interspecific competition for food. 

The distinct locomotor activity rhythms of the 
three species of octopods, coupled with the dif- 
ferences in habitat and substrate preference, 
minimize interspecific contact in the field. Com- 
petition for food may not be the only factor im- 
pacted. Octopods are predators on other mem- 
bers of the genus (Van Heukelem, 1976); larger 
animals will readily feed upon small individuals. 
The three species of this study vary dramatically 
in size. 0. cyanea is the largest, with maximum 
weight over 5 kilograms. 0. ornatui< has a mid- 
dle size range, with mature individuals of 500 
grams common. The crescent octopus is the 
smallest species. Maximum size of any in- 
dividual collected by the author was 90 grams. 
The vast size difference may encourage the 
smaller species, especially the crescent octopus, 
to limit activity to periods when contact with the 
larger individuals of other species is unlikely. It 
should be noted, however, that activity rhythms 
were recorded and maintained in the laboratory 
in isolation, so visual observation of individuals 
of the same or other species is not required for 
demonstration of locomotor patterns. Boyle 
(1980) indicates that a larger octopus may take 
food forcefully from a smaller octopus, so tem- 
poral spacing may be especially beneficial to 
smaller species. 

Competition for home sites in the aquarium 
has been documented in OcfopM.s vulgaris Cuvier 
(Boyle, 1980). On the reef flats on the south 

shore of Oahu, tide pools, coral heads and coral 
rubble are found near each other. By inhabiting 
one of these microhabitats over others, the 
three species of Hawaiian octopods in this study 
may minimize interspecific contact and reduce 
competition for home sites. When this habitat 
selection is coupled with the temporal spacing 
evident in activity periods, contact between 
species may be effectively reduced. 

Thus, Octopus cyanea. 0. ornatus and the 
"crescent octopus" show distinct species-specific 
patterns of locomotor activity in the laboratory 
under light regimes similar to those in the field. 
The three species are active on the same reefs 
on the south shore of Oahu at different times of 
the day. These times correspond to the periods 
of maximum activity recorded in the laboratory. 

The author thanks A. N. Popper, I. M. Cooke 
and R. E. Young for their advice and encourage- 
ment; W. H. A.kaka, R. Shoemaker and M. 
Valdez for help in design and construction of the 
acoustic monitor, and W. van Heukelem for 
sharing his extensive knowledge of cephalopod 
biology. She is also appreciative of editorial 
assistance from F. G. Hochberg and R. T. 
Abbott. The research was supported in part by 
an NIH predoctoral fellowship granted the 


Akaka, W. H. and Houck, B. A. 1980. The use of an 
ultrasonic monitor for recording locomotor activity. 
Eehnv. Ret^. Methods and Infttr. 12:514-516. 

Altman, J. S. 1967. The behavior of Octopus vulgaris Lam. 
in its natural habitat: a pilot study. Undenvater Ass. Rep. 

Boyle. P. R. 1980. Home occupancy by male Octopus cul- 
garis in a large seawater tank. Animal Behavior 

UeCoursey, P. J. 1976. Builogiral Rtiylhmx in the Marine 
Enviromnent. Columbia: Univ. of South Carolina Press. 

Franzisket, L. 1969. The ratio of photosynthesis to respira- 
tion of reef building corals during a 24 hour period. Forma 
et Functio 1:153-158. 

Houck, B. A. 1977. A morphological and behavioral study of 
an extra-ocular photoreceptor in octopods. Ph.D. disser- 
tation. University of Hawaii. 

Kayes, R. J. 1974. The daily activity pattern of Octopus 
vulgaris in a natural habitat. Marine Behac. Physiol. 


October 2y, 1982 

Vol. 96(4) 

U.S. Dt'partmenl of ('omnierce 1944. Sunshine tables. (npuf: cyaiipa and Octopnn 111111/11. I'h.l). tlissiTlal imi. 

Weather Bur. No. 805. Washington, D.C. University of Hawaii. 

Van Heukelem, W. F. 1966. Some aspects of the ecologi,' and Woods, J. 1965. Octopus-watching off Capri. Aninidls 

ethology of Octopus cyanea Gray. M. S. thesis. University 7:324-327. 

of Hawaii. Yarnall, J. L. 1969. Aspects of the behavior of thinijus 
1976. Growth, bioenergetics and life-span of Or- ciiancd Gray. Ariiiiidl Behariar 17:747-754. 




Barry Roth 

California Academy of Sciences 
San Francisco, California 94118 

William K. Emerson 

and American Museum of Natural History 

New York, New York 10U24 

Specimens of Persicula tessellata (Lamarck, 1822), a species originally de- 
scribed from an unknoum province, are here recorded from the Pacific coast of 
Panama and are compared with Persicula accola (Roth and Coan, 1968) from the 
same area. The identity of Persicula porcellana (Gmelin, 1791), a Caribbean 
species that ha.s been confused with P. tessellata, is reevaluated. 

Live-collected specimens of a Persicula from 
Pacific Panamic waters submitted to the Ameri- 
can Museum of Natural History by James 
Ernest agree well with the holotype of Mar- 
(jincllii fessellata Lamarck, 1822, described from 
an unknown locality. These specimens permit a 
reevaluation of the synonymy oi Marginella tes- 
sellaia with Voluta porcellana Gmelin, 1791, 
first advanced by Reeve (1846) and perpetuated 
in the west American literature (Keen, 1958; 
Coan and Roth, 1966; Roth and Coan, 1968). 
Persicula tessellata (Lamarck) is here removed 
from the synonymy of P. porcellana (Gmelin, 
1791) and is recognized as an inhabitant of 
tropical west American waters. The Pacific 
\'-diYc\m\c Persirula areola Roth and Coan, 1968 
is compared with Lamarck's P. tessellata, and 
the identity of Gnu'lin's P. porcellana from the 
trojiical western .'\tlantic is reconsidered. 

Institutional Abbreviation.s 

AMNH = American Museum of Natural History 
ANSP = Academy of Natural Sciences of 

CAS = California Academy of Sciences 
MCZ = Museum of Comparative Zoology, 
Harvard University 

USNM = National Museum of Natural History. 
Smithsonian I nstitutic )n 

Family Marginellidae Fleming, 1828 
Genus Persicula Schumacher, 1817 
Type species by monotypy: Persicula vari- 
abilis Schumacher, 1817 {= Voluta persicula 
Linnaeus, 1758), Recent, west Africa. Medium- 
sized to small marginellids with flat spire usually 
concealed l:)y callus; aperture narrow through- 
out its length; anterior end strongly notched; 
most species strikingly marked or colored. 
Eocene to Recent, Old and New World tropics. 

Persicula tessellata (Lamarck, 1822) 

(Figs. 1-6) 
MiirijiiwUd lessctlala Lamarck, 1822:361-362. Kiener. 

1834:24, pi. 5, fig. 20. 
Maryinetla parri'ltrina (Gmelin), Reeve, 1864:pl. 13, figs. 

53a, 53b. 

Vol. 96(4) 

October 2[). 1982 

THENAlTlLrS 157 

"IGS. 1-6. Persicula tessellatafLar/iarcA:, 18-J'^). 1-4, specimens from Lm Zurrones, Isia Cehaeo, Panama, ex Ernest C.ullec- 
'11)11. 1. 2. (AMNH 208760). S. U. (AMNH 208761); x2. 5, 6. holotype o/Marginella tessellata, Lamarck Collection. Mus. Hist. 
Vat. Geneva, courtesy of Dr. E. Binder; x2.1. 7, 8. Persicula accola Roth and Coan (1968). Isla Perida. Panama. (AMNH 
102770); X 2. 9-16. Persicula porcellana fGme/in, 1791). 9, 10. reproduttion of Chemnitz (1788) figures 11,19 and U20. Spe^igler 
)iUt-ction; x i. 11, 12, heach specimen from "Venezuela" (AMNH 4961,3. ex Constable Collection, received in 1901); x2. 13, 14, 
Specimen, from Puerto La Cruz. Venezuela (AMNH 20301,5), x 2. 15, 16, specimen from "St. Martha, " Colombia in the Redfield 
Collection (ANSP 29116). labeled typological lot o/Marginella obesa Redfield; x2. 

'-'i rsicula porcellana (Gmelin), Jousseaume, 1875:260. Keen, 
1958:436, fig. 680. Coan and Roth, 1966:282-283 (m 
part), pi. 48, figs. 14, 15 (non figs. 16, 17, which = Persi- 
i-iila accola Roth and Coan, 1968). Roth and Coan, 
1968:63, pi. 7, figs. 5, 6, 

\nn Valuta porcellana Gmelin, 1791:3449. 

\on Marginella tessellata Wood, 1828:42. pi. 3, fig. 31 
I =P. areola Roth and Coan. 1968]. 

Original description - "24. Marginelle par- 
quetee. Marginella tessellata. M. testa obovatd, 
apice retusa, albidd, punctis rufis quadratis 
transversim seriatis tessellata: seriis cofertis; 
columella plicis praecipuis quinis instructd: 
supra aliis duohus seu tribus minimis; labro in- 
tus crenulato. An valuta porcellana? Chemn. 


October 29. 1982 

Vol. 96(4) 

Conch. 10. t. 150. f. 1419, 1420. C.mel. p. 3449. 
no. 139." (Lamarck, 1822, p. 3(il). 

Tyijc locality - Unknown. Here de-sij^nated, 
off Isla Cebaco, west Panama. 

Supplementary deseription - The following- 
description is based on seven specimens dredged 
in about 30 meters at Los Zurrones, Isla Cebaco, 
Panama (7°40'10"N. 81°30'10"W) by James 
Ernest, 1982 (6 specimens, AMNH 208761; 1 
specimen CAS 032387). Shell large for the 
genus, solid, ovate, diameter 0.61-0.67 times 
length; narrower anteriorly, greatest width at 
60-64 percent of distance from anterior end; 
surface polished, unscuptured e.xcept for a few 
raised incremental lines. Color white with 9-11 
spiral rows of grayish brown, subrectangular 
spots covering 60-80 percent of shell surface; 
most posterior row just below suture; most 
anterior row running along margin of anterior 
canal; spots irregularly spaced, not tending to 
align longitudinally, mostly wider than interven- 
ing spaces in a row; spiral rows added by fission 
of an existing row; incompletely split spots 
usually present. Spots well defined or slightly 
blurred (particularly on trailing edge) by overly- 
ing clear to translucent whitish callus. Spire 
covered by a Hat or slightly projecting apical 
pad of white callus, circled by a ring of more or 
less fused reddish brown blotches; spacing of 
these blotches independent of pattern on body 
whorl. Outer lip strongly thickened by callus, 
not extending beyond apex, white, tinged along 
outer margin with reddish brown, finely denti- 
culate on apertural edge. Urate within. Aperture 
narrow, almost even but slightly wider anterior- 
ly, white inside, with acute posterior sulcus and 
deep, narrow, oblique anterior canal. Canal set 
off from body whorl by raised fasciolar Hange; 
columellar side produced dorsally into a rounded 
spur, ticked posteriorly with reddish brown. 
Parietal wall moderately convex, shallowly ex- 
cavated anterior to center; with thick white 
callus along its length, merging posteriorly with 
apical callus; left edge of callus a raised rim 
paralleling aperture. Columella with 8 folds in- 
cluding one at base of columella, decreasing in 
strength posteriorly; posterior fold sometimes 
barely perceptible; second fold from anterior 
end large, bifid. 

Dimension.^ a/the seven specimens examined- 
Largest specimen: length 18.4 mm, diameter 
12.3 mm; smallest specimen: length 16.1 mm, 
diameter 10.6 mm; average for length: 17.5 mm; 
average for diameter: 11.1 mm. 


Comparison of the seven specimens from 
James Ernest with the holotype of Persicula 
lessellata (Figures 5, 6) leaves little doubt that 
they are conspecific. The holotype is 16.5 mm in 
length and 10 mm in diameter. It has 9 subequal 
rows of spots on the back of the body whorl; fis- 
sion increases these to 12 by the last quarter of 
the whorl. The apical callus pad projects slight- 
ly. Interspaces in a row of spots are somewhat 
variable, particularly around the middle of the 
whorl. The thick parietal callus pad and shallow 
excavation of the parietal wall are exactly like 
those of the Ernest specimens (Figures 1-4). 

The holotype is a slightly worn specimen. Its 
pattern is warm brownish orange. The super- 
ficial wash of milky callus that covers the shell in 
many Persicula species (e.g. P. bandera Coan 
and Roth, 1966; P. hilli (Smith, 1950); P. accola 
Roth and Coan, 1968; and the present species) 
imparts a grayish cast to the dark patterning 
underneath. When this layer is removed by ero- 
sion, the remaining pattern shows as a warm 
brownish red or orange, as in the holotype of P. 
tessellata. In the Ernest sjiecimens, the differ- 
ence in hue between the hody-whoi'l pattern and 
the brown tinting on spire, outer liji, and canal is 
probably not the ]:>roduct of separate pigments, 
but the result of the body-whorl patterns being- 
seen through the translucent wash of callus. 

The Ernest specimens of P. tessellata differ in 
several ways from Persicula accola Roth and 
Coan (1968, pi. 7, figs. 7, 8). P. accola is smaller 
(range of 20 adult specimens examined: lenglli 
11.2-14.6 mm, diameter 7.0-9.1 mm). The rows 
of brown spots cover about 90 percent of the 
shell surface, the interspaces between them be- 
ing narrow and uniform (Figures 7, 8); as in P. 
tessellata, the basic number of rows seems to be 
nine, with higher nimibers always the result i)t' 
fission of one or more rows. The sfiire is tiat to 
very low, not projecting as in somi' /'. ttsscllnln. 

Vol. 96(4) 

October 29, 1982 


and the apical callus is ringed by a dense, solid 
brown band or else (less commonly) entirely 
brown. The brown ticking on the anterior 
fasciolar flange is darker and more extensive 
than on P. tessellata; the brown patch on the 
outer margin of the thickened outer lip darker, 
longer, and more distinct. Most P. accola have 
seven columellar folds; occasionally a faint 
eighth fold is perceptible posteriorly. Specimens 
with five and six columellar folds occur. 

All known occurrences of Persicula accola are 
on the Pacific coast of Panama between Punta 
Burica (8°02'N, 82°52'W) and Punta Mariato 
(7°12'N, 80°53'W), intertidally (Coan and Roth 
in Keen, 1971) to nine meters (AMNH 202770, 
AMNH 208759). Isla Cebaco, the source of the 
P. tessellata specimens, is also in this area. 
James Ernest (m litt. to Emerson, 18 May 1982) 
reports that P. accola is collected intertidally in 
sand mixed with mud, while P. tessellata is 
found in "deeper water [to about 30 meters] and 
white sand, with very clear water." 

Gmelin (1791) described Valuta porcellana 
with reference to two figures in Chemnitz (1788) 
(Figures 9, 10) depicting a broadly ovate Persi- 
cula with 16 rows of small, irregularly shaped 
spots. The locality was cited as Indian Ocean, 
undoubtedly speculative since Gmelin probably 
had no firsthand knowledge of the source of the 
Chemnitz specimen. No occurrence of any Per- 
sicula resembling P. porcellana in the Indian 
Ocean has been confirmed in the many interven- 
ing years. (Nevertheless, a few poorly grounded 
citations in the later literature still place P. 
porcellana in the Indian Ocean; cf. Dodge's 
[1955, p. 85-86] speculative remarks.) 

In describing Marginella tessellata, Lamarck 
(1822, p. 361-362) "cited the same figure in 
Chemnitz with an interrogation mark, then 
went on to comment, about the specimen in his 
own collection ("Mon cab."), "Ses points ne sont 
pas sagittes comme dans le figure citee de 
Chemniz, mais carres." Most later authors, 
beginning with Reeve (1864) have interpreted 
Lamarck's species and Gmelin's as synonymous, 
perhaps mainly because both authors cited the 
same Chemnitz figure. Dodge (1955, p. 86) inter- 
preted Lamarck's comment about the shape of 
the spots as a criticism of the fidelity of Chem- 

nitz's illustration. It can equally well be seen as a 
statement of contrast between the Chemnitz 
specimen and Lamarck's own, which we accept 
as the holotypic specimen (Figures 5, 6). 

Kiener (1834, pi. 5, fig. 20) illustrated as Mar- 
ginella tessellata a specimen with eleven rows of 
subrectangular, brownish orange spots and a 
moderately projecting apical callus. At 26.5 mm 
in length, the figure is certainly enlarged; but 
with the allowance for a little bit of artistic 
license, it could practically be Lamarck's 

Under the name Marginella tessellata, Sower- 
by (1846) illustrated three different specimens. 
His figure 195 is evidently Persicula chryso- 
melina (Redfield, 1848). Figure 194 shows the 
dorsal view of a Persicula 11.1 mm in length 
with nine spiral rows of variously sagittate, 
subrectangular and irregular spots. Figures 
196 and 197 show dorsal and ventral views of a 
similar Persicula 14.8 mm in length with 13 
rows of irregular to sagittate spots and a 
maculated outer lip. Sowerby's (1846, p. 395) 
description of the species characterizes the col- 
umella as white, swollen in the middle, and 
elevated into a swollen varix anteriorly. Sower- 
by was also the first to associate a locality with 
the name M. tessellata, stating that it was a 
common species from Venezuela. 

The lack of precision in the Sowerby illustra- 
tions makes them somewhat difficult to inter- 
pret. However, the characters of maculation on 
the outer lip (visible in ventral view) and a tumid 
parietal callus rising to a ridge or varix anterior 
to the middle are strongly suggestive of an At- 
lantic species variously identified in museum 
collections as Persicula obesa (Redfield, 1846) 
and P. porcellana (Gmelin. 1791). Most of these 
specimens are from older collections and the lo- 
calities are indefinite: Venezuela (AMNH 49643, 
ex James Arnold Constable collection; CAS 
030298, ex Ruth Coats collection), Honduras 
(USNM 19617), "West Indies" (MCZ 265502). 
They have 14-16 spiral rows of spots. The spots 
are usually punctate, sagittate, or dash-shaped 
rather than rectangular; there is typically a zone 
of larger spots over the broadest part of the 
body whorl, and another, less pronounced, about 
one-fourth of the distance posterior to the 


October 29, 1982 

Vol. 96(4) 

anterior end. The apical callus is flat or slightly 
projecting and strongly tinged with brown. The 
callus thickening of the outer lip is whitish, often 
with brown dashes echoing the color pattern of 
the body whorl. None of these specimens show 
the longitudinal brown patch of P. tessellata and 
P. accola. Authenticated modern records of this 
species in the western Atlantic are: Puerto La 
Cruz, Anzoategui, Venezuela, three specimens 
(AMNH 203045, ex F. J. Fernandez H. collec- 
tion, 1976) (Figures 13, 14), Tobago, West In- 
dies, two specimens (AMNH 141495, ex Mrs. 
Stuart Brown collection, 1967). Rios (1970, 
1975) records this species (as P. obesa) from 
Brazil, but the cited specimens are actually from 
Venezuela {teste Rios, 1982). 

The hoiotype of Valuta porcellana Gmelin, 
formerly in the Lorenz Spengler (1720-1807) 
collection of the University of Copenhagen, is 
lost (Coan and Roth, 1966). The Chemnitz draw- 
ings (cf. Figures 9, 10 with 11, 12) cited by 
Gmelin for his species agree more closely with 
the Caribbean species than with the Pacific Per- 
sicula tessellata. Voluta porcellana is the oldest 
name for the Caribbean species, with Voluta 
albida Bosc, 1801 (cites Chemn. figs. 1419 and 
1420) and Margimlla obesa Redfield, 1846 (not 
Sowerby, 1846) both junior synonyms. Redfield's 
taxon was based on specimens provided by W. 
W. Whitney from the "Caribbean Sea at Cartha- 
gena, SA" [Colombia] that were deposited in the 
"Cabinet of the Lyceum" [of Natural History, 
New York City]. The collection of the Lyceum 
was destroyed by a fire in 1866, (Fairchild, 
1887). There are, however, four specimens 
(ANSP 29116) from the collection of John H. 
Redfield (1815-1895) labeled as "types". None of 
these specimens match the measurements or 
figures cited in the original description (Red- 
field, 1846, p. 164, 165, pi. 10, figs. 5a, b). Fur- 
thermore, the locality for this lot is given as "St. 
Martha, S.A." [ = Santa Marta, Colombia]. Al- 
though this may not represent the type lot, the 
specimens agree well with Redfield's description 
and illustration (see figures 15, 16). Three topo- 
typical specimens of this taxon (ANSP 29386) 
are in the Robert Swift collection (1796-1872). 
Marginella similis Sowerby, 1846, from 
"Brazil", is a possible additional junior synonym 

of P. porcellana. Redfield (1848) and others have 
referred Sowerby's taxon to the synonymy of 
Marginella obesa Redfield, but we have not ex- 
amined the types of M. similis. which we 
presume are in the British Museum (Natural 

Coan and Roth (1966) designated the hoiotype 
(which they called a lectotype) of Marginella 
tessellata to be a neotype for Voluta porcellana. 
Their aim was to cement the synonymy, gener- 
ally accepted up to that time, of M. tessellata 
and V. porcellana. In light of the foregoing 
demonstration that Persicula tessellata is a 
Pacific species, and Persicula porcellana is a 
consistently differing Atlantic species, the Coan 
and Roth neotype designation fails to meet the 
provisions of ICZN Article 75 (c) (4), which 
states that to be validly designated a neotype 
must be published with "evidence that the 
neotype is consistent with what is known of the 
original type-material, from its description and 
from other sources." Coan and Roth (1966, p. 
283) actually commented that the Chemnitz 
figures differed in pattern and proportions from 
any "P. porcellana" that they had examined at 
the time. We suggest that the neotype designa- 
tion is invalid. 

Coan and Roth (1966) first regarded Persicula 
porcellana as an eastern Pacific species and il- 
lustrated (1966, figs. 16-17) a specimen from 
Isla Coiba, Panama, under that name. Later, the 
same authors (Roth and Coan, 1968) distin- 
guished P. accola as an eastern Pacific species 
(including the specimen they had figured in 
1966), and allocated P. porcellana to the west- 
ern Atlantic, stating that they had examined 
"from the Caribbean, specimens that look like 
Lamarck's type [i.e., of P. tessellata] and still 
others which closely resemble the original 
Chemnitz figures" (Roth and Coan, 1968, p. 63). 
Such specimens include MCZ 265501 (no locali- 
ty; four P. porcellana and four P. a-ccola), USNM 
90175 ("Venezuela," ex Wesleyan University col- 
lection; two, probably P. accola), and USNM 
413838 ("Brazil," ex Ford collection; one P. ac- 
cola). It seems most probable that, with the ex- 
ception of the four true P. porcellana in MCZ 
265501, these all represent misallocations of 
eastern Pacific shells. 

Vol. 96(4) 

October 29, 1982 


In summary, Persicula tessellata (Lamarck, 
1822) and Persicula accola Roth and Coan, 1968, 
ire similar eastern Pacific species occurring in 
;he same part of western Panama. Persicula 
oorcellana (Gmelin, 1791) is the valid name for a 
Caribbean species with more rows of finer spots; 
Marginella obesa Redfield, 1846, is synonymous. 

We are grateful to Mr. James Ernest of 
Balboa, Republic of Panama, for calling our at- 
:ention to his findings of Panamanian Persicula 
md generously providing us with specimens. 
Drs. Pablo E. Penchaszadeh, Instituto de Tec- 
nologia y Ciencias Marinas, Universidad Simon 
Bolivar, Venezuela, and Eliezer de Carvalho 
Rios, Museu Oceanografico de FURG, Rio 
jrande, Brazil, generously supplied information 
ind/or specimens. Dr. Robert Robertson 
ANSP) kindly arranged for the loan of speci- 
mens. Mr. William E. Old, Jr. (AMNH) gave his 
idvice and provided technical assistance. Mrs. 
\my Hkimi (AMNH) typed the manuscript. 
"^Jina Root and Karl F. Koopman (both AMNH) 
jave bibliographic assistance. 


3osc, L. A. G. 1801. Histoire Naturelle des Coqiiilles . . . 

Paris, vol. 5, 255 p. 
Ilhemnitz, J. H. 1788. Neues systematisches Conchylien- 

Cabin^t. 10. Niirnberg, xxiv + 376 p. 
Z;oan, E. V. and B. Roth. 1966. The west American Mar- 

ginellidae. The Veliger 8(4):276-299. 
Dodge, H. 1955. A historical review of the mollusks of 

Linnaeus. Part; 3, the genera Bulla and Valuta of the 

class Gastropoda. Bull. Amer. Mus. Nat. Hist. 


Fairc-hild, H. L. 1887. A history of the New York Academy 
of Sciences, formerly the Lyceum of Natural History. 
New York, published by the author, xii + 190 p. 

Gmelin, J. F. 1791. Syslema naturae per regna tria naturae 
. . . , ed. 13, Leipzig, 1(6):3021-3910. 

.lousseaume, F. P. 1875. Coquilles de la famille des mar- 
ginelles flpc. Mag. ZiioL. ser. 3, 3:164-271, 429-435. 

Keen, A. M. 1958. Sea shells of tropical west America; 
marine mollusks from Lower California to Colombia. 
Stanford, Calif, xi + 624 p. 

1971. Sea shells of tropical west America; 

marine mollusks from Baja California to Peru, ed. 2. 
Stanford, Calif, xiv + 1064 p. 

Kiener, L. C. 1834-1841. Species general et iconographie 
des coquilles vivantes. Genre marginelle. Paris, 44 p. 

Lamarck, J. B. P. A. de M. de. 1822. Histoire naturelle 
des aiiimtmx sans vertebres, 7. Paris, 711 p. 

Redfield, J. H. 1846. Description of some new species of 
shells. Ann. Lye. Nat. Hist. New York. 4(5):163-168 
[Feb., 1846]. 

1848. Descriptions of new species of Bulla and 

Marginella. with notes upon G. B. Sowerby, Jr's. mono- 
graph of the latter genus. Ibid., 4(12):491-495 [Sept., 

Reeve, L. A. 1864-1865. Conchologia iconica; or illustra- 
tions of molluscous aniynals. 15. Monograph of the genus 
Marginella. London, 27 pis. with text. 

Rios, E. C. 1970. Coastal Brazilian seashelU. Rio Grande, 
255 p. 

1975. Brazilian marine mollusks iconography. 

Rio Grande, 331 p. 

Roth, B. and E. V. Coan. 1968. Further observations on the 
west American Marginellidae with the descriptions of two 
new species. The Veliger ll(l):62-69. 

Smith, M. 1950. New Mexican and Panamic shells. The 
Nautilu.9. 64(2):60, 61. 

Sowerby, G. B. IL, 1846. Monograph of the genus Mar- 
ginella. Thesaurus Conchyliorum. or figures and descrip- 
tions of shells. l(7):373-406, [27 Nov., 1846]. 

Wood, W. 1828. Supplement of the Index testaceologicus; 
or a catalogue of shells, British and Foreign. London, 
vi + 59 p. 


A grant of $400 will be awarded by the Ameri- 
can Malacological Union to a graduate student 
'or the purpose of conducting field studies of 
and or freshwater mollusks. Grant proposals 
should include: (1) A brief description of the pro- 
oosed project, (2) an estimated budget of antici- 
Dated expenditures, and (3) a statement from 
;he student's supervising professor that the pro- 
ect is an appropriate part of the student's re- 
search. Proposals must be submitted in quadru- 

plicate by January 1, 1983 to Dr. Alan J. Kohn, 
President A.M.U., Dept. Zoology, University of 
Washington, Seattle, Washington 98195. The 
announcement of the award will be made by 
April 1, 1983. 

This award has been made possible by the 
many friends of Dr. Joseph C. Bequaert who 
died on January 19, 1982 at the age of 95. Dr. 
Bequaert, known as Uncle Joe to his students 
and friends, spent a lifetime in the field studying 
land and freshwater mollusks, and this award is 
a tribute to his memory. 


October 29, 1982 

Vol. 96(4) 


An Authoritative Guide to over 4,000 Marine Shells of the World in Color 

by R. Tucker Abbott and S. Peter Dance 

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talents to produce the largest and most complete, all-color, general identification shell book 
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( luidc til tlic Litcnitiirc 
The COMPENDIUM gives you the leading source-books and indices 
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eo 36 


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