ANUARY 25. 1982 THE NAUTILUS 1^«N ()()28-i:i44 Vol. 96 No. 1 A quarterly devoted to malacology and the interests of conchologists Founded 1889 by Henn' A. Pilsbry. Continued by H. Burrington Baker. Editor-in-Chief; R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William J. Clench Curator Emeritus Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, New York 10024 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, Ohio 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, California 90007 Dr. Arthur S. Merrill 103 West 8th Avenue Cudjoe Gardens Summerland Key, Florida 33043 Dr. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Joseph Rosewater Division of Mollusks U.S. National Museum Washington, D.C. 20560 Dr. G. Alan Solem Department of Invertebrates Field Museum of Natural History Chicago, Illinois 60605 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, Ohio 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Charles B. Wurtz 3220 Penn Street Philadelphia, Pennsylvania 19129 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, Florida 32901 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 22.55 Melbourne, Florida 32901 Tlie Nautilus (USPS 374-980) ISSN 0028-1344 OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbcjurne, FL 32901) Mail: Box 2255, Melbourne, FL 32901 Second Qaas Postage paid at Melbourne, Florida and other post offices Subscription Price: $12.00 (see inside back cover) $14.00 (foreign); institutions $18.00 THE NAUTILUS Volume 96, number 1 - January 25, 1982 ISSN 0028-1344 CONTENTS Joseph Rosewater A New Species oiHipjiopu^- (Bivalvia: Tridaciiidae) Francesco B. Trama Occurrence of the Asia^c Clam Corbicula Jluininea in the Raritan River, New Jersey 6 William D. Shepard Rediscovery of a Portion of the Isely Unionid Collections 8 Philippe Bouchet and Francoise Danrigal Napoleon's Egyptian Campaign (1798-1801) and the Savigny Collection of Shells 9 Clement L. Counts, III and Robert S. Prezant Shell Microstructure of Corbicula flmninea (Bivalvia: Corbiculidae) 25 Paul W. Parmalee and Walter E. Klippel A Relic Population oi Obuvaria retussa in the Middle Cumberland River, Tennessee 30 Publications Received 2 Recent Deaths 2 THE NAUTILUS January 25, 1982 Vol. 96(1) PUBLICATIONS RECEIVED Brooks, Stanley Truman (1902-1960), Biobibliosraphy by R. I. .lohnson in Ocr. fnpers MoUuaka. va]. 4, no. 59, pp. 266-268(1981). Clarke, Arthur H. 1981. The Tribe Alasmidontini (Unioni- dae: Anodontinae), Part 1: Pegias, Alasmidon.la, and A rcidenx. Smithsonian Contributions to Zoology, no. 326, pp. iii + 1-101, 32 figs. Ecologic and taxonomic study with illustrations of shell, soft parts, gloehidia, descriptions, distributional maps, synonymies, and simple anatomical test for sex determination. Alusmidontn mliusta n. sp.. extinct mussel from (jharlotte, N(" (not Vnio mlntatif Sowerby, 1840). Habe, T. and Iwao Taki. 1981. A Catalogue of Molluscs of Wakayama Prefecture, the Province of Kii. 1. Bivalvia, Scaphopoda and Cephalopoda. F'ubl. Seto Marine Biol. Lab.. Special Series, vol. 7, no. 1, pp. xx 304, 13 pis. (One in color). Based on Kuroda's Manuscript. Eleven new species of .Japanese Bivalvia are described. Synonymies very complete. 51 cephalopods treated by I. Taki (pp. 234-264). .Johnson, Richard I. 1981. Recent and Fossil Unionacea and Mutelacea (Freshwater Bivalves) of the Caribbean Islands. Occasional Papers on Mollii.ank, long-time shell dealer, editor of "Shell Notes" (1942-51), born in Lantana, Florida, June 10, 1895, died Oct. 11, 1981, in Ferandina Beach, Florida. Sexauer, Howard T. 1981-82 President of the Sanibel Captiva Shell Club, well-known poet, former resident of Montpelier, Vermont, died at age 82 in Florida on Dec. 1, 1981. Zeigler, Rowland Franklin, born .lune 1, 1915, in Greely- ville, SC, formerly chief of obstetrics and gynecology at McLeod Hospital in P'lorence, SC, author (with Porreca) of "Olive Shells of the World" and articles in medical journals, died at age 66, on Sept. 21, 1981, in Durham, NC. His widow, Edna Jo continues his interests in olive shells. I'ln' I'irsI a>vrhfdogiral wtirk liij mi .■\mmcaii jiuhHuhed in A mtn'icit. THOMAS Savs CONCHOLOGY 1819 A beaut iful facsimile copy of the third edition of Say's fiindeinental umrk from Nicholson's British Encyclopedia Philadelphia 1819 This rare work, never before reprotluced in full, con- tains 20 pages and four lithograph plates of Say's new land and freshwater species of mollusks. This is the expanded version of the first (1816) and second (1818) edition. 32 pages, paperbound. The [ireface, by R. Tucker .Abbott, tells the history of this article and gives brief biographies of the author, the editor, the publisher and the printer. Limited edi- tion. $5.00, including postage. F'oreign postage, plea.se add $1.00. Amki;i('an MALACOUHiisTS, Inc. P.O. Box 2255 Mi-llMiurne. FL 32»(I1-I«2S Vol. 96(1) January 25, 1982 THE NAUTILUS A NEW SPECIES OF HIPPOPUS (BIVALVIA: TRIDACNIDAE) Joseph Rosewater Department of Invertebrate Zoology National Museum of Natural History Washington, D.C. 20560 ABSTRACT Hippopus porcellanus n. sp. is described from Sibutu Island, Sulu Archipelago, Philippines, bringing the known members of the genus Hippopus to 3, including H. hippopus (Linne, 1758), Recent, I ndo- Pacific, and H. gimteri Mansfield, 1937, Fossil. Lower Miocene of Florida. For several years shell dealers have received large numbers of a rather distinctive giant clam belonging to the genus Hippopus which they refer to as the "China Clam", and which they consider to be distinct from the "Horse's Hoof or "Bear Paw", Hippopus hippopus (Linne, 1758). It is much thinner and smoother than the usually elaborately sculptured H. hippopus, and lacks most of the characteristic strawberry color of the latter. When I reviewed the classification of Tridacnidae this phenotype was believed to intergrade completely with H. hippopus (Rose- water, 1965, p. 361). Recent examination of nearly two dozen specimens of the "China Clam" and comparison with H. hippopus persuades me that they are separate species. Hippopus porcellanus new species (Figs. 1-4) Description: Shell reaching 216 mm (about 8 1/2 inches) in length, semicircular in outline and globose in shape; usually only moderately in- flated; with valves closed byssal orifice is very narrowly gaping. Valves not excessively heavy, markedly translucent, colored occasionally with weak strawberry blotches arranged concentri- cally or scattered; color of interior porcel- laneous, of exterior grayish white. Surface of valves remarkably clean, except for scattered coralline algae and debris. Primary radial sculp- ture consisting of 13 or 14 low rib-like folds distributed over surface of valve, extending on- to ventral slope where they become obsolete. Secondary radial sculpture consisting of low rib- lets which are nearly obsolete on primary folds but are more prominent in their interstices. Riblets varying somewhat in width, usually lack- ing spines or evidencing only microscopic con- centric imbrications. A few low tubular spines present on primary fold bordering ventral slope and occasionally on ventral portions of other primary folds; folds usually smooth dorsally. Concentric sculpture consisting of microscopic, wavy, imbricate lines of growth. Dorsal margin undulate, with series of 8-9 rounded to squarish, medially projecting, interdigitating processes representing extremities of rib interstices. Hinge line usually longer than half the length of valve. One oblong cardinal tooth in each valve; 2 elongate posterior laterals in right and a single moderately sharp one in the left valve. Liga- ment secondarily prosodetic. Umbos directed postero-medially. Edge of byssal orifice with a series of 8-12 light-yellow, rather poorly developed plicae which remain fairly constant in size or become only slightly larger posteriorly. Ventral slope moderately concave. Hinge plate suffused with orange. Pallial line entire, mod- erately narrow. Muscle scars central, medium sized; the posterior adductor scar round in both valves, the posterior pedal retractor scar is smaller and elongate, the two extending over portions of two fold interstices in each valve. Area within pallial line, excluding muscle scars, dull; pallial line, muscle scars and areas to edge of shell shiny. Prodissoconch unknown. The anatomy and life history of this species are unknown. Following my studies (Rosewater, 1965) several persons have achieved success in studying the spawning and development of Hip- THE NAUTILUS January 25, 1982 Vol. 96(1) Vol. 96(1) January 25, 1982 THE NAUTILUS popus hippopus (Jameson, 1976; Gwyther and Munro, 1981). Measurements (mm): Tridacnidae are indis- criminately inequivalved, and the figures given under "length" and "height" are always the max- imum measurement. "Width" is the greatest distance through opposed valves. Lticality Sihiitu MaMmte (Philippines) Zmtiliiiiniiia hi Id. SnIii Sra No. Specimens 10 5 1 (i Range of Lengths 87-138 75-1.57 216 93-214 Range of Heights 60-105 53-115 167 74-150 Range of Widths 46-78 35-101 141 48-126 Range of Weights, Left Valve (grams) 27-142 11-221 479 38-399 Average Width/Length .60 .60 .66 .58 Types: Holotype ANSP (The Academy of Natural Sciences of Philadelphia) 246600, 157 mm length, 113 mm height; 3 paratypes ANSP 354770, 155 mm length, 115 mm height, 101 mm length, 81 mm height, and 75 mm length, 53 mm height; 1 paratype USNM (National Museum of Natural History, Washington, D.C.) 807720, 133 mm length, 98 mm height. Type locality: Sibutu Island, Tawi Tawi Group, Sulu Archipelago, Philippines (4°46'N; 119°29'E), du Pont- Academy Expedition, 1958. Other Material Examined: ANSP 209699 Zamboanga, Mindanao Island, Philippines, ex. A. B. Bronson, 1956 (10 specimens, see Meas- urements); ANSP 228977, Masbate Island, Philippines, du Pont-Academy Expedition, 1958 (1 specimen, see Measurements); USNM, "Sulu Sea", ex. A. D'Attilio and John Root (6 speci- mens, see Measurements). Distribution: Philippines, mostly known from the Sulu Archipelago; 1 specimen from Masbate Island, central Philippines. Etymology: "porcellanus", an adjectival name referring to the porcelain-like appearance of the shell of this species. Remarks: Shells of this new species are con- sistently smoother, more semicircular in outline, proportionately lighter in weight and thinner than H. hippopus (see figs. 1-4). When viewed apart from the latter they present a rather ro- tund appearance although measurements show they are no more obese, but, in fact, are often less wide. Compared with H. hippopus. H. porcellanus has a very narrow byssal orifice, with the plicae less well developed and lighter in color (fig. 4). The single posterior lateral tooth of the left valve is moderately sharp in H. porcellanus while in H. hippopus it is blunt. The only other large bivalve species with which H. porcellanus is likely to be confused is Tridacna derasa (Roding, 1798). Specimens of T. derasa, of the same length as H. porcellanus (200 mm-f ), are smooth, may be similarly semi- circular in outline and exhibit similar obesity. It is quite likely that living T. derasa has a con- siderably more colorful mantle, that of H. hip- popus, at any rate, being rather sombre olive. The shell of T. derasa is even smoother than that of H. porcellanus, lacking in development of tubular spines and having low primary and secondary folds, although sometimes developing strong, continuous, undulate concentric ridges (Rosewater, 1965, pi. 281, fig. 1). Shells of T. derasa lack the orange and yellow coloration present in Hippopus in the areas of hinge and byssal orifice. Byssal plicae number 6-7, are low and elongate to nearly obsolete in T. derasa while in H. pjorcellanus they are shorter and number from 8-12. The posterior adductor muscle/posterior pedal retractor scar complex is comparatively larger in T. derasa and its umbos tend to be less distinctly convoluted than in H. porcellanus. although size for size the former tends to have a heavier shell. I originally thought that H. porcellanus dif- fered from H. hippopus subspecifically rather than specifically. As subspecies the two would normally have more or less discreet geographic ranges. While it appears that H. porcellanus lives mostly in the southern Philippines, more precisely the southern Sulu Sea, a specimen has been reported from Masbate Id. in the central Philippines. Furthermore, H. hippopus is distributed in the same area besides having a more extensive range in the western Pacific. Since the ranges of the two species seem not to be in any real way mutually exclusive, the sub- specific status seems doubtful, and they are here considered to be separate species within the THE NAUTILUS January 25. 1982 Vol. 96(1) genus HippoptLs. It is quite obvious that the known geographic range of H. porcellanus is considerably more restricted than that of//, hip- popus (see Rosewater, 1965, pi. 272). ACKNOWLEDGMENTS Special thanks are expressed to those who assisted in gathering information and donating specimens: Anthony D'Attilio, John Root, Robert W. Morrison, R. T. Abbott, and Betty and Robert Lipe. I am grateful to R. Robertson, ANSP, for loaning specimens with accurate locality data. R. D. Turner, and K. J. Boss, Museum of Comparative Zoology, Harvard University, and H. A. Rehder, and R. S. Houbrick, USNM, offered helpful criticism. LITERATURE CITED Gwyther, J. and J. L. Munro. 1981. Spawning induction and rearing of larvae of Tridacnid Clams (Bivalvia: Tridacni- dae). AquaeuHurc 24:197-217. Jameson, Stephen C. 1976. Early life history of the Giant Clams Tridarna crocea Lamarck, Tridarna maxima (Roding). and Hippopus hippopus (Linnaeus). Pacific Science 30(3):219-233. Rosewater, Joseph. 1965. The Family Tridacnidae in the Indo-Pacific. Indo-Pacifie Mollusca l(6):347-396. Shell Factory. 1963. Shells of the Winid. Catalog no. 109, 160 pp. Fort Myers. Florida. "China Clam" illustrated on p. 45. OCCURRENCE OF THE ASIATIC CLAM CORBICULA FLUMINEA IN THE RARITAN RIVER, NEW JERSEY^ Francesco B. Trama Department of Biological Sciences Rutgers LTniversity New Brunswick, New Jersey ABSTRACT The mid-Atlantic range of the Asiatic clam, Corbicula fluminea (Muller. 1771^) is extended to the Raritan River in central New Jersey. A successfully breeding population was found in the nontidal region near a water supply intake. These clams have also colonized far upstream into the South Branch of the Raritan but not the North. Branch. No reason for this difference is known. Living specimens were found downstream in the tidal portion of the river which is freshwater but polluted. There was no evidence of reproduction in this region of the river. Since it was first observed in this country in the Columbia River, Washington (Dundee and Dundee, 1958) the Asiatic clam, Corbicula fluminea (Muller, 1774; (alias manilensis Philip- pi) has spread rapidly into many river systems across the United States (Sinclair, 1971). Exten- sion of its range to the mid- Atlantic region was documented by Diaz (1974) in the James River, 'This study was supported iiy the Center for Coastal and En- vironmental Studies. Rutgers University. Assistance of Joy Bergelson in pierforming field collections is acknowledged. Virginia, and by Fuller and Powell (1973) in the Delaware River between Philadelphia, Pennsyl- vania and Trenton, New Jersey. Diaz estimated the year of introduction in the James River to be 1968 and Fuller and Powell concluded that Cor- bicula was present in the Delaware River since at least 1971 or 1970. Crumb (1977) later re- ported it in the Delaware River between Tren- ton and Burlington in September 1971. On March 26, 1981 many empty shells and a few living Corbicula were collected from a tidal (Init freshwater) region of the Raritan River Vol. 96(1) January 25, 1982 THE NAUTILUS near New Brunswick, New Jersey (Sta. 1, Fig. 1). Collections of macrobenthos taken from this same region in September 1980 did not contain any specimens of Corbicula. The largest shell collected in March 1981 was 25 mm in length and was estimated to be between 3 and 4 years of age at the time of death. It was concluded that the year of invasion was not later than 1978 and that the invasion site had to have been upstream from this point of initial discovery. During July and August 1981 the occurrence and range of Corbicula in the Raritan River was studied. Samplmg extended upstream from Sta. 1 (tidal but freshwater) into the North and South Branches of the Raritan River (Fig. 1). At Sta. 1 many empty shells were again found. Many live specimens, however, were also collected from current-swept bottoms covered by gravel and small pebbles The livmg clams fell into two size classes; largest specimen was 25 mm in length and had a grossly distorted shell growth that appeared to commence from a shell length of 20 mm. The smallest specimen was 15 mm in length and appeared nt)rmal. The greatest population density and evidence of reproduction were found at Sta. 2 which is in the vicinity of the Elizabethtown Water Com- pany's intake on the Raritan River which is non- tidal at this point. Largest living clams ranged from 15 to 17 mm in length and a first year class ranged from 2 to 6 mm in length. No Corbicula were found in the Millstone River (Sta. 3) im- mediately above its confluence with the Raritan River. The substrate in this portion of the Millstone River was soft muck and, therefore, was not a preferred habitat for these clams. Very few living Corbicula were taken at Sta- tions 4 and 5, and these ranged from 7 to 10 mm in length. Many empty shells were found at Sta. 4. Even though a suitable type substrate existed at Stations 6, 7 and 8 only one specimen of Cor- bicula (10 mm) was found at Sta. 6. None was found at Stations 7 or 8, nor was there evidence of dead Corbicula shells anywhere in the North Branch of the Raritan River. On the other hand, tliere were significant populations in the South Branch. Specimens ranged from 2.5 to 10 mm in length at Stations 9, 10 and 11. No dead no. 1. Samjile files ,„, the Ranta,, Hirer. Netr Jersey. Samplvngfor Corbicula look place belween July 2:1 and August 2ti. 1981. 8 THE NAUTILUS January 25, 1982 Vol. 96(1) specimens or empty shells were found in those locations. It seems that the Asiatic clam has established itself in the tidal and nontidai, freshwater (even polluted) regions of the Raritan River. The population center (initial colonization?) is in the nontidai waters of the main stem in the vicinity of the intake of the Eiizabethtown Water Com- pany. Furthermore, this foreign species has suc- cessfully colonized the South Branch of the Raritan River but for some reason has not as yet moved very far into the North Branch of the Raritan River. This discovery extends the north- ern range of Corbicula in the mid-Atlantic region. LITERATURE CITED Crumb, S. E. 1977. Macrobenthos in the tidal Delaware River between Trenton and Burlington, New .Jersey. Chesapeake Science 18:253-265. Diaz, R. J. 1974. Asiatic clam, Corbicula manilensis (Philip- pi), in the tidal James River, Virginia. Chesapeake Science 15:118-120. IXindee, D. S. and H. A. Dundee. 1958. Extension of known ranges of four mollusks. The Nautilus 72:51-53. Fuller, S. L. H. and C. E. Powell. 1973. Range extensions (jf Corbicula manilensis (Philippi) in the Atlantic drainage of the United States. The Nautilus 87:59. Sinclair. R. M. 1971. Annotated bibliography on the exotic bivalve Carbicula in North America, 1900-1971. Sterki- anaNo. 43:11-18. REDISCOVERY OF A PORTION OF THE ISELY UNIONID COLLECTIONS William D. Shepard Oklahoma Biological Survey Norman, Oklahoma 73019 During a recent renovation of the Inverte- brate Range at Stovall Museum of Science and History (University of Oklahoma), a collection of unionids was found carefully wrapped in old newspapers and stored away. Part of these were labeled as collected by F. B. Isely in Oklahoma between 1908 and 1912. The remaining unionids were also from Isely's collections. Upon inquiry with several malacologists, it appears that this is the largest extant portion of Isely's collec- tions. These are part of the specimens collected by Isely for his study of the clams of Eastern Oklahoma, which appeared in 1925. After rearrangement of the above specimens and merging of several other collections, a series of unlabeled unionids were found. These differed from all the other specimens, however, in that they had small, circular, numbered, cop- per tags wired to the shells. On further examina- tion it was determined that these were a portion of the specimens used by Isely in his classical migration study (Isely 1914). They were also col- lected in Oklahoma. The rediscovery of these specimens is impor- tant for several reasons. They were used by Ise- ly and therefore are now voucher specimens for liis works. Their presence also allows some in- sight into the taxonomy of his time or at least Isely's interpretation of it. Secondly, the specimens have a historical importance in that some of the streams Isely collected have now been altered so as to deplete the original unionid fauna. Finally, some of these specimens repre- sent the only collections from various streams in Oklahoma. Isely's specimens plus others donated to Stovall Museum form a nucleus for a growing collection of Oklahoma unionids. The Stovall col- lection combined with that located at OU's Biological Station on Lake Texoma (Texas- Oklahoma border) represents perhaps the major collections of Oklahoma unionids. LITERATURE CITED Isely, F. B. 1914. Experimental study of the growth and migration of freshwater mussels. Bureau of Fisheries Document, No. 792. 1925. The freshwater mussel fauna of Eastern Oklahoma. I'rur. OkUi. Acad. Sci. 4:43-118. Vol. 96(1) January 25, 1982 THE NAUTILUS NAPOLEON'S EGYPTIAN CAMPAIGN (1798-1801) AND THE SAVIGNY COLLECTION OF SHELLS Philippe Bouchet and Francoise Danrigal Museum National d'Histoire Naturelle 55, Rue de Buffon 75 Paris (5°), France ABSTRACT During the Napoleonic campaign to Egypt a collection of Red Sea, Mediter- ranean and continental niollut^k^ ivas brought together by Savigny who published upon them in 1817 as folio engravings. These served as type figures for a number of species described by 19th ceyitury malacologists, including Audouin, Ehrenberg, Deshayes, Philippi, Hupe, Landrin, Jonas, Issel, Morlet, Vaillant, P. Fischer, Tapparone-Canefri. von Martens, Weinkauff, Monterosato, Jousseaume, H. Fischer and finally Pallary. A checklist and illustrations of 86 of these original types frmn the Savigyiy collection, now in the Paris Museum, are presented, supplemented by reproductions of the unpublished color vellums of opisthobranchs and cephalopods. Jules Cesar Savigny was born at Provins, France, on April 5, 1777. In 1793, in the middle of the Revolution, the Convention transformed the Royal Gardens into a "Museum d'Histoire naturelle". Savigny was then 16 years old and was sent to Paris to study at the "Ecole de sante" and the Museum. His professors were Lamarck, Cuvier, Daubenton; they noted the young student's assiduity and also took note that he was always wearing the same Nankin costume. Lamarck was then writing the "Nouvelle En- cyclopedie" and assigned Savigny his first work, the drafting of the part on the sorrel plant. This paper showed the wealth of his knowledge and soon he was named professor of botany at the "Ecole Centrale" in Rouen. Cuvier advised him to wear a wig to look older and more serious! But Savigny was never to go to Rouen. Napo- leon Bonaparte was already preparing the Egyptian Campaign. Bonaparte not only wanted to conquer the country but also make it a model of French culture. He added to the expedition troops a "Commission des sciences et des arts", con- sisting of scholars, painters, architects and others. Bonaparte asked his friend Cuvier to name the naturalists who would join the expedi- tion. Savigny and Geoffroy Saint Hilaire were selected for the zoological part. Savigny was then 21 years old; he had been trained as a botanist, but Cuvier's opinion was that "he would become a zoologist when he would decide to". The Expedition After the Battle of Campoformio in 1797, Bonaparte realized that he could not invade England, so he decided to make war on Egypt. In 1798, 38,000 men gathered at Toulon on 335 ships; among them were the members of the "Commission des Sciences et des Arts". On Floreal 30th, year VI, (i.e. May 19th, 1798) Geoffroy Saint Hilaire and Savigny left Toulon. They were to reach Alexandria 41 days later. Savigny was placed in the 4th class of the Com- mission, with a very low salary, but thanks to Geoffroy Saint Hilaire, he was soon admitted to the first class. The Nile was flooding and the expeditionary forces were directed to Cairo which they en- tered on July 23 after the Battle of the Pyra- mids. During this time the Commission settled in Rosetta for a month. Savigny and Geoffroy Saint Hilaire stayed for several weeks in the islands of Lake Menzale; in September they visited Salahied and Damiette. They worked all during the autumn in the delta and left Damiette for Cairo in December. Geoffroy Saint Hilaire was exhausted, but Savigny was in very good condition. A current joke of the time in Cairo was to call scholars, the donkeys, then used as "taxis", because the members of the Commission were always travelling on donkeys. Reprints of this article are available for U.S. $7.00 (postage paid) from American Malacologists. Inc.. P.O. Box 2255, Melbourne. FL. ;32901, U.S., A. 10 THE NAUTILUS January 25, 1982 Vol. 96(1) Between August 1 and 2, Nelson defeated the French Naw in Aboukir and cut all contacts with F" ranee. In Cairo, Bonaparte organized the country under his rule and founded the "Institut d'Egypte". He then prepared the expedition to Syria 13,000 infantrymen and the cavalry left in the beginning of February, 1799. After sev- eral victorious battles, Bonaparte was finally stopped at Saint Jean d'Arce, where 4,000 were killed. On June 14 the army was back in Cairo, Savigny was the only naturalist to follow the troops. In August, Bonaparte turned over the com- mand to Kleber and returned to Paris, where the "Directoire" was weakened. The members of the "Commission" were sent to Suez at the end of 1 799. Savigny was very e.xcited by the wealth and beauty of the Red sea fauna. But Kleber had to face the Turks and the British and the scholars were sent back to Cairo. Kleber was murdered shortly after he won the Battle of Heliopolis; France was then finally defeated at Canope in 1801. The journey of the 45 members of the "Com- mission" back to France was rather unfortu- nate. On April 6, 1801, they left Cairo for Alex- andria. On their arrival they were quarantined by the French general, Menou. They were trans- ferred to a brig, and waited 21 days for permis- sion to leave the bay. The British troops stopped them, took them to Abukir, then back to Alexan- dria. Menou had surrendered on terms stipulat- ing that the collections gathered by the "Com- mission" in Egypt should be given to the British. The scholars rebelled and threatened to destroy the collections. Finally, on September 26, 1801, they were allowed to go back to France with their collections. The Original Plates Savigny returned to Paris in February, 1802. He did a great amount of work on the material he took back with him and published various papers and memoirs between 1801 and 1810. Among them were the study of 1,200 insects and his famous "Histoire naturelle et mytholo- gique de I'Ibis" (1805). In 1817, he published 125 plates for the atlas of the "Expedition d'Egypte", in "grand-aigle" size (71.5 x 52 cm); in 1826 they were reprinted by Panckoucke in reduced size (68.5 x 52 cm). These engraved plates were prepared from the original colored vellums now preserved in this Museum's library. They had beon drawn very accurately, with a wealth of details, by the best artists of the time. Because the "Commission" judged that they were too difficult to reproduce in color, only black and white copies were printed. Savigny suffered progressively failing eye- sight, approaching blindness after 1815, for which reason he never published any text to ac- company the plates. But Cuvier found that this iconography was unique and Audouin was cho- sen to write an explanatory part. Audouin was a professor of entomology at the Museum, and was not prepared for this work. But he was the son-in-law of Brongniart, a member of the "In- stitut", director of the Sevres manufacture, and a friend of the powerful Cuvier. There is considerable difficulty in determining the exact dates of publication of the "Explica- tion sommaire des planches", of which two edi- tions were printed. The first edition bears on the title page "public par les ordres de sa majeste I'Empereur Napoleon Le Grand" and is dated 1809 on the cover page, but was evidently published much later (Laissus, 1973); the Mollusca are treated in tome 1, part 4, and begin with a copy of a letter dated Nov. 1, 1825! The register of the letters and declarations of the "Commission d'Egypte", kept in the Manuscript Department of the Na- tional Library, Paris, shows that the Natural History section belongs to the third issue of reports. On April 13, 1826 the last manuscript by Audouin concerning shells was sent to the "Imprimerie Royale", but we could not deter- mine whether the first edition, third issue, was actually printed before the end of 1826. However it seems wise to assume that Sherborn (1897:287) was right and that this part "may be safely regarded as dated 1826." Almost simultaneously a second edition, "dediee au roi", was being printed by Panckouke. This bears 1827 on the title page but it appears also that this date is a fake. Pallary accepted this date (1827) in a bibliography, but mentioned 1829 in the text (Pallary 1926). In fact the earliest date we can ascertain is July 19, 1828 (Bibliographie de la France, 1828). The text was published while Savigny was still alive. When he became aware of the contents, he became very irritated and sent a letter to the Academy of Sciences. Later on, L. Pfeiffer pro- nounced similarly .severe judgement, and re- proached Audouin with the fact that he had taken all the benefit of Savigny's work. P. Fischer and Pallary had the same opinion. In the checklist, we have given references to both edi- tions of Audouin's text because the first one is apparently very rare in libraries. Vol. 96(1) January 25, 1982 THE NAUTILUS 1 1 Savigny's work, despite its unfinished condi- tion, was to interest naturalists for a long time. The very short explanations by Audouin were later completed, first by Issel (1869), then by Pallary (1926). But many other authors referred to Savigny, among them, Jonas (1846), Pfeiffer (1846), Vaillant (1865), P. Fischer (1865, 1870, 1871), von Martens (1866), Tapparone-Carnefri (1875), Jousseaume (1888). Many of the speci- mens depicted by Savigny have therefore be- come the types of new species, which are enumerated below. The Collection From Egypt, Savigny brought back a rich col- lection of birds, fishes, mammals, insects and shells. Included were also mollusks in alcohol, given by Savigny to Cuvier in 1802; later on these were said to be deposited in the Gallery of Anatomy but we have been unable to trace them. In 1853, the manuscripts, notes and collec- tions of Savigny were offered to the town of Versailles. Curation was entrusted to the "Societe des sciences naturelles et medicales de Seine et Oise". Deshayes once planned to publish a report on the Mollusca collection but never did (Landrin, 1865). In 1864, the Society realized its inability to curate the material properly and asked for help from the Museum. Valenciennes and Gratiolet were contacted, but the Society and the Museum lacked the sum of 2,500 francs to fund the naturalist who intended to work at Versailles. The collection then sank into obli- vion. In 1919, the council of the "Societe des Sciences naturelles" was disbanded because of World War I. The librarian of the Versailles library took this opportunity to transfer the col- lection in a very rough fashion to a cellar in order to gain some space. In 1926, when Pallary wrote his "Explication des Planches", the vellums and collections of Savigny were considered to be lost. In April 1927, he visited his friend the bryozoologist, Canu, in Versailles and realized that some shells in the cellar of the municipal library matched exactly some depicted on Savigny's plates. Be- tween August and September, he carefully studied the shells which were finally to join the national collections in 1930, together with the 5 volumes of vellums. Pallary (1931, 1932, 1934) produced a very detailed history of the expedition and the collec- tions together with a biography of Savigny. He evidently had planned to publish a report on the collection after he had rediscovered it, but only a small part was published (1932); this concerned only the shells not depicted in the plates. We are therefore producing here the latest chapter of this two centuries-old story with the realization of the nomenclatorial importance of this material. Acknowledgements We are grateful to the librarians who helped us during this work and to A. Foubert for Figures 1-51, Mrs. Guillaumin, Centre de Microscopie du C.N.R.S. for Figures 52-84 and P. Lafaite for the colorslides of the vellums. Figures 93-101. We wish to thank Mr. Richard E. Petit of South Myrtle Beach, South Carolina, for making possible the publication of the color plate. Checklist of the names based partly or entirely on Savigny's plates. It has been our intention to simply list the material on which a name is based, when the original description refers to the 1817 plates. Lectotype designation is the affair of the mala- cologist actually engaged in a process of revi- sion of a group and should not be done for the mere sake of it, as is too often the case in such checklists. We are aware of the fact that, strictly, only the very specimen depicted on Savigny's plate should be considered the tyyje of later authors. It is, indeed, possible to recognize the ty^ie when it is a large and distinct shell, but it is not so when dealing with the many microgastropods, of which Savigny appears to have been the earliest collector in the Indo-Pacific. (legyptmcii Ehrenberg, 1831 (Siuxinen) Fig. (51 Savigny 1817: pi. 2, fig. 24 Ehrenberg 18.31: (no page number). Signature E. Name based on an unknown number of animals from Damiette and Savigny's figure. One shell in coll. Savigny. Ehrenberg collection in the Berlin Mu.seum may contain specimens. iwgyptidcd Chenu, \8A5 (Tridacna) Savignv 1817: pi. 10, fig. 1-2 Chenu '1845: 2, pi. 7. fig. 1-2 Name based on Savigny's plate and a number of subfossil shells from the Suez area. .Judging from the illustrations, however. Chenu's shells have nothing to do with Savigny's, and are tyjjical Tridacna mfuimd (Rbding). affinis Issel, 1869 (Chiton) Fig. .') I Savignv 1817: pi. 3, fig. 8 lsseri869: 234 Name based on Savigny's figure and seven specimens from the Gulf of Suez, now in MGD (Genoa, Italy). One specimen in coll. Savigny. See also sariynyi Pilsbry 1892. (irsMomsin Issel, 1S69 (Turbo) Fig. 2.5 Savigny 1817: pi. 5, fig. 28 12 THE NAUTILUS January 25, 1982 Vol. 96(1) Issel 1869: 220 Name based on Savignv's figure and a subfossil shell which could not be traced in MGU. Five shells in coll. Savigny. audouini Jonas, 1846 (Fasciolaria) Fig. 42 Savignv 1817: pi. 4, fig. 17 Jonas i846a: 63 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. audouini Jousseaume, in Lamy, 19lH(Pristis) Fig. 86 Savignv 1817: pi. 8, fig. 11 Lamy 1918: 30 Name based on Savigny's figure and two specimens from Suez, now in MNHN. No Savigny material left. bacillum Issel, 1869 (Cerithium) Fig. 77 Savignv 1817: pi. 4, fig. 28 Issel 1§69: 340 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. bertholleti Issel, 1869 (Rissoina) Fig. 38 Savignv 1817: pi. 4. fig. 2 Issel 1869: 208 Name based on Savigny's figure, named by Audouin 'Rissoa de Bertholett'. Two shells in coll. Savigny. bourguignati Landrin, lS65(Unio) Fig. 4 Savigny 1817: pi. 2, fig. 3 Landri'n 1865: 5, fig. 1-3 Name based on Savigny's figure and specimens collected by Savigny in Damiette (Landrin actually saw the collection). Three syntypes in coll. Savigny. brongnartii Audouin, \S26 (Tricoiia) Fig. 26 Savignv 1817: pi. 5, fig. 23 Audouin 1826: 41; 1828: 181 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. aiillaudi von Martens, 1866 iSpatka) Fig. 1 Savignv 1817: pi. 7, fig. 1 von Martens 1866: 9 Name based on Savigny's figure and specimens in the collec- tions of Caillaud, Mousson, Leiden Museum and British Museum. Two bivalve specimens and one valve in coll. Savigny. calloiia P. Fischer, 1 871 (Gen.a) Fig. 32 Savigny 1817: pi. 5, fig. 10 Fischer 1871: 218 Name based on Savigny's figure and specimens collected at Suez by Gaudry, which could not be traced. Two shells in coll. Savigny; the small one (3.2 mm) is here depicted. The larger one (5.8 mm) is chipped and smaller than the natural size shell depicted fig. 10,5 in Savigny. carinata Pallarv, 1926 (Risella isseli var.) Savigny 1817: pi. 5, fig. 34 Pallary 1926: 84 Name based on Savigny's figure and an unknown number of shells from Suez. See under isseli. ringulata Issel, 1869 (Eulimella) Fig. 73 Savigny 1817: pi. 3, fig. 25 Issel 1869: 182 Name based on Savignv's figure and 3 specimens from Suez, two of which are in MGO (Genoa). One shell in coll. Savignv. The name was changed to TurboniUa isseli by Trvoii <1886:339) because it is supposedly congeneric with fur- bonilla nngutatn Dunkcr 1860. Issel 1869: 177 Name ba.sed on Savigny's figure and a specimen from Suez. There are two shells with tnis locality m the Issel coll. in MGD. One specimen in coll. Savigny. c/j/ppomoru-s Jousseaume. \SSS {Clypeomorus) Fig. 40-41 Savigny 1817: pi. 4, fig. 10 .Jousseaume 1888: 171 Name based on Savigny's figure and an unknown number of specimens from the southern Red Sea. In the Jousseaume coll. (MNHN), there are 33 shells from Massawa and 12 shells from Obock. Two shells in coll. Savigny. eoenobita VaiWant, IS65 (Mytilus) Fig. 12 Savigny 1817: pi. 11, fig. 3 Vaillant 1865: 115, 122 Name based on Savigny's figure and several specimens from Suez, now in MNHN. One specimen in coll. Savigny. mncentrica Audouin, 1826 (Doris) Savigny 1817: Gasteropodes pi. 1, fig. 5 Audouin 1826: 14; 1828: 128 Name based on Savigny's figure. No material left. Fig. 97 Fig. 18 corbieri JoTtus, lSi&(Chama) Savigny 1817: pi. 14, fig. 8 Jonas 1846c: 126 Name based on Savigny's figure. Two complete specimens in coll. Savigny. craticulata Issel, 1SG9 (Odontostomia) Fig. 75 Savigny 1817: pi. 3, fig. 39 Issel 1869: 180 Name based on Savigny's figure and subfossil specimens, which could not be traced in MGD (Genoa). One shell in coll. Savigny. ruvieri Audouin, l&2G(Emargmula) Savigny 1817: pi. 1, fig. 9 Audouin 1826: 27; 1828: 152 Name based on Savigny's figure. No material left. Fig. 89 Fig. 37 dautzenbergi Pallary, 1926 (Donovania) Savigny 1817: pi. 4, fig. 20 Pallary 1926: 71 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. desgenettii "Risso" Audouin, \S2G (Bul.lal Fig. 55 Savignv 1817: pi. 5, fig. 6 Audouin 1826: 39; 1828: 178 Name based on Savigny's figure. Five shells in coll. Savigny. desmarestii Audouin, 1826 (/Sissoaj Savigny 1817: pi. 3, fig. 21 Audouin 1826: 36; 1828: 171 Name based on Savigny's figure. No material left. Fig. 91 Fig. 74 dysmatica Issoi. \St'^>lOdontostomia) Savigny 1817: pi. 3, fig. 36 Fig. 72 doliiformis Pallary, l92(ifPyrgulina) Savigny 1817: pi. 3, fig. 42-43 Pallary 1926: 63 Name based on Savigny's figure and several specimens from Suez, which could not be traced. Five shells in coll. Savigny. dnrbignii Audouin, \S26IRissoa) Fig. 68 Savignv 1817: pi. 3, fig. 22 Audouin 1826: 36; 1828: 171 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. dftrbigvii Audouin, \?<2(>(Snssurella) Fig. 63 Siivignv 1817: pi. 5, fig. ,30 Audouin 1826: 42; 1828: 183 Name based on Savigny's figure. Five shells in coll. Savigny. Vol. 96(1) January 25, 1982 THE NAUTILUS 13 doriae Issel, 1869 (Stomatella) Fig. 48 Savigny 1817: pi. 5. fig. 8 Issel 1869; 228 Name based un Savigiiy's figure and 10 specimens from Suez, which could not tje traced in MGD. Seven shells in coll. Savigny. draparnaudi Audomn, IS26 (Tricolia) Fig. 46 Savigny 1817: pi. 5, fig. 19 Audouin 1826: 41; 1828: 181 Name based on Savigny's figure. Two shells in coll. Savigny. elata Semper, in Issel, 1869(Sfa/io/a^ Fig. 84 Savigny 1817: pi. 3. fig. 15 Issel 1869: 330 Name based on Savigny's figure. Fifteen shells in coll. Savigny. elegans Audouin, 1S2G (Tritonia) Savigny 1817: Gasteropodes pi. 2, fig. 1 Audouin 182?: 15; 1828: 130 Name based on Savigny's figure. No material left. Fig. 94 Fig. 24 eroopotitanus Issel, 1&Q9 (Turbo) Savigny 1817: pi. 5, fig. 27 Issel 1§69: 219 Name based on Savigny's figure and one specimen from Suez which could not be traced in MGD. Six shells in coll. Savigny. erythraea Hupe, 1SS4 (Blanivillia) Fig. 94 Savigny 1817: pi. 8, fig. 6 Hupe 1854: 223 Name based on Savigny's figure. Two valves in coll. Savigny. erythraea Issel, 1869 (Nassa costulata var.) Fig. 44 Savigny 1817: pi. 6, fig. 4 Issel 1869: 126 Name based on Savigny's figure and specimens from Suez which could not be traced in MGD. One shell in coll. Savigny. erythraealsse], 1SQ9( Lticina) Fig. 20 Savigny 1817: pi. 8, fig. 8 Issel 1869: 84, pi. 1, fig. 9 Name based on Savigny's figure and 3 specimens from Suez which could not be traced in MGD. One shell in coll. Savigny. erythrofa Issel, 1869 (Area lactea var.) Savigny 1817: pi. 10, fig. 7 Issel 1869: 89 Name based on Savigny's figure and 6 specimens from Suez. Six paired valves in coll. Savigny. favrei Landrin, 1865 (Helix) Fig. 21 Landrin 1865: 2, fig. 1-3 Name based on a shell from the Savigny coll., not depicted in the atlas. Holotype in coll. Savigny. ferussacii Audouin, 1826 (Scalaria) Fig. 81 Savigny 1817: pi. 3, fig. 13 Audouin 1826: 35; 1828: 169 Name based on Savigny's figure. One shell in coll. Savigny is considered to be the holotype. feuilletii Audouin, 1826 (Neritina) Fig. 28 "Savigny 1817: pi. 5, fig. 11 Audouin 1826: 40; 1828: 179 Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. fourierii Audouin, 1S2& (Bulla) Fig. 54 Savigny 1817: pi. 5, fig. 5 Audouin 1826: 39; 1828: 178 Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. Ireminvillii Audomn, lS2(\(Kist;ua) Fig 67 Savigny 1817: pi. 3, fig. 20 Audouin 1826: 36; 1828: 170 Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. gennesi H. Fischer, 1901 (Claneulus) Fig 49 Savigny 1817: pi. 3, fig. 3 Fischer 1901: 123, pi. 4, fig. 11-12 Name based on Savigny's figure and one specimen from Djibouti, now in MNHN. Five shells in coll. Savigny. gevtiluomiana Issel, lS69(Eulima) Fig. 78 Savigny 1817: pi. 3, fig. 32 Issel 1869: 183 Name based on Savigny's figure and four shells from Suez. There are 6 shells witfi this locality in MGD. Three .speci- mens in coll. Savigny. quenni Audouin. 1826 fTrico/ia^ Savigny 1817: pi. 5, fig. 24 Audouin 1826: 41; 1828: 181 Name based on Savigny's figure. Pallary argued that this is only a color form of T. brongnarti. No material left. girardi Audouin, 182& (Bulla) Fig. 52 Savigny 1817: pi. 5, fig. 3 Audouin 1826: 39; 1828: 178 Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. hemprichi Ehrenberg, 1831 (Helix) Fig. 27 Savigny 1817: pi. 2, fig. 12 Ehrenberg 1831: Helix no. 4 (no page number) Name based on a number of snails collected near Alexandria and on Savigny's figure. 24 shells in coll. Savigny. The Ehrenberg coll. is supposedly in Berlin. hemprichi \&se\, \i>%9 (Trochus) Fig. 22 Savigny 1817: pi. 3, fig. 6 Issel 1869: 329 _ Name based on Savigny's figure. Six shells in coll. Savigny. /wjrn'dMS Orbigny, li,2() (Octopus) Fig. 100 Savigny 1817: Cephalopodes pi. 1, fig. 2 Orbignv 1826: 144 Name based on Savigny's figure. No material left. kumboldti Audouin, \B2() (Anatola) Savigny 1817: pi. 5. fig. 1 Audouin 1826: 39; 1828: 177 Name based on Savigny's figure. No material left. immaculata Audouin, 1826 (Doris) Savigny 1817: Gasteropodes pi. 1, fig. 2 Audouin 182?: 13; 1828: 126 Name based on Savigny's figure. No material left. Fig. 85 Fig. 93 infracostata Issel, \SS9(Riselln) Fig. 64 Savigny 1817: pi. 5, fig. 40 Issel 1869: 195 Name based on Savigny's figure and three specimens from Suez, which could not be traced in MGD. Three shells in coll. Savigny. i'sseli Semper, in Issel, 1869 (Risella) Fig. 65 Savigny 1817: pi. 5, fig. 35 Issel 1869: 194 Name based on Savigny's figure and several specimens from Suez and Zanzibar. There are 2 shells from Zanzibar with a label in Semper's handwriting in MGD. Five shells in coll. Savigny. isseli Nevill & Nevill, 1875 (Marginella) Nevill & Nevill 1875: 95 Fig. 57 14 THE NAUTILUS January 25, 1982 Vol. 96(1) New name for Marginelta pygmaea Issel 1869, non Sowerby 1846. See pygmaea. isseli Trvon, XSSQ (TurhoniUa) New name for Eulimella cingulata Issel 1869. See this name. tstfemicuTO Issel, 1869 ^Cardwmj F'g- 1^ Savignv 1817: pi. 9, fig. 11 Issel 1§69: 74 , , • f Name based on Savigny's figure and several sp?cirneris from Attaka, near Suez, which could not be traced in M(,t). One valve in coll. Savigny. jmnardi Audouin, 1 826 (Scalaria) Fig' '^'^ Savignv 1817: pi. 3, fig. 4 Audouin 1826: 3.5; 1828: 169 . . , u „ ■ ii Name ba.sed on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. hi n th a Audouin , 1 826 (Cypraea) Fig. 33 Savigny 1817: pi. 6, fig. 27 Audouin 182?: 4.5; 1828: 190 . . , u „ ■ u Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. lamarckii Audouin, 1826(PMpa^ Fig. 83 Siivignv 1817: pi. 2, fig. 1 Audoufn 1826: 31; 1828: 161 . , . „ ■ Name ba.sed on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. teroii Landrin, 1 865 (Helix) Fig. 23 Landrin 1865: 4, fig. 1-4 , , ^ . . .. „ Name based on a shell collected by Savigny at the Fyra- mides, not depicted in the atlas. Holotype in coll. Savigny. lessepsianus Vaillant, 1865 iLUhodomus) Fig. 1 1 Savignv 1817: pi. 11, fig. 1 Vaillant 1865: 123 , • . Name based on Savigny's figure and several specimens trom the gulf of Suez, two of which are in MNHN. One specimen in coll. Savigny. Fig. 17 mareoticum Pallarv, 192S (Cardiiim) Savigny 1817: pi. ?), fig. 10 PallarVl926: 109 Name ba.sed on Savigny's figure. One specimen in coll. &ivigny is considered to be the holotype. marmorata Audouin, 1 826 (Doris) Fig. 95 Savignv 1817: Gasteropodes pi. 1, fig. 7 Audoutn 1826: 14-15; 1828: 129 Name based on Savigny's figure. No material left. marmorata Pallary, 1926fGCTta; Fig. 31 Savigny 1817: pi. 5, fig. 9 Pallarv 1926: 76 Nameoased on Savigny's figure. Four shells in coll. Savigny. marfen-si Issel, 1869 (Alaba) Fig. 71 Savignv 1817: pi. 3, fig. 26 Issel lJi69: 206 Name based on Savigny's figure and a subfossil specimen from the Red sea, which could not be traced in MGI). Four shells in coll. Savigny. rtwngii Audouin, 1826rB»/W Fig. 58 &vi'gny 1817: pi. 5, fig. 7 Audouin 1826: 39; 1828: 178 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. oblongun Audouin. \&2i\ I FJ nun ibrnnchus) Fig. 98 Savigny 1817: Gasteropodes pi. 3. fig. 1 Audouin 1826: 20-21; 1828: 140 Name based on Savigny's figure. No material left. olivaefonni'i Issel, 1869 (Tomatina) Fig. 56 Savigny 1817: pi. 6, fig. 25 Issel 1869: 171 , , . Name based on Savignv's figure and three specimens trom Suez, one of which is in MGD. One shell in coll. Savigny. perlntus Issel , 1869 (Triforis) Fig. 79 Savigny 1817: pi. 4, fig. 4 Issel 1869: 152 ,,,.., ■ , Name based on Savigny's figure and subfossil specimens, ot which one is in MGD with the data 'Red sea'. One shell in coll. Savigny. See also savignyanus. pharaonis P. Fischer, 1871 (Area) Fig. 7 Savignv 1817: pi. 10, fig. 9 Fischer 1871: 213 , , • , Name based on Savigny's figure and several specimens trom Suez which could not Be traced. One valve in coll. Savigny. iMraonis P. Fischer, 1870 (Mytilus) Fig. 10 Savigny 1817: pi. 11, fig. 5 Fischer 1870: 169 , ■ r Name based on Savignv's figure and several specimens trom Suez, of which six bivalve specimens and one valve are m MNHN. One specimen in coll. Savigny. philippii Issel, 1869 (Cydostremn) Fig. 66 Savigny 1817: pi. 5, fig. 33 Issel 1869: 189 , . • ^ c Name based on Savignv's figure and 6 specimens trom Suez which could not be traced in MGD. Two shells in coll. Savigny. pulvis Issel, 1869 (Cerithium) ' Fig. 35 Savigny 1817: pi. 4, fig. 5 Isseri869: 1.50 , , Name based on Savigny's figure and one or several specimens from Suez of which one is in MGD in Genoa. One shell in coll. Savigny. miqmaea Issel, 1869 (Marginella) Fig. 57 Savigny 1817: pi. 6, fig. 26 Issel 1869: 150 , c, ■ ■ r , Name based (with question mark) on Savigny s figure anti one shell from Suez, now in MGD. Four shells in coll. Savigny. reticulata Philippi, 1853 (Scissurella) Fig. 62 Savignv 1817: pi. 5, fig. 29 Philippi 18.53: 38, pi. 6, fig. 11 ^,. , , . ,. New name fi)r "Scissurella decussatn Orbigny. .-^ucJouin (1826:42; 1828:183), not Orbigny, 1824. Therefore the name is based on Savignv's figure; Phihppi also refers to specimens from thp Red Sea collected by Hempnch and Ehrenberg. Four shells in coll. Savigny. richardi Audouin, 182& (Cardiuvi) Fig. 14 Savigny 1817: pi. 9, fig. 14 Audouin 1826: 51; 1828: 201 Name based on Savigny's figure. One specimen m coll. Savigny is considered to be the holotype. rissoi Audouin, 1826 (Tricolia) Fig. 47 Savignv 1817: pi. 5, fig. 18 Audouin 1826: 41; 1828: 181 Name based on Savigny's figure. Two shells in coll. Savigny. rissoi Weinkauff, 18S5 (Rissoina) Savignv 1817: pi. 4, fig. 1 Weinkauff 1885: 63. pi. 15d Fig. 76 ,. „„, ^ fig. 13 Name based on the name 'Manzelia de Risso' Audouin (1828: 171) and a shell from Mauritius. Two shells in coll. Savigny. Vol. 96(1) January 25, 1982 THE NAUTILUS 15 roemeriana Issel, IS&d (Verms) Fig. 62 Savigny 1817: pi. 8, fig. 3 Issel 1869: 64 Name based on Savigny's figure and 7 valves from Suez, now in MGD. Seven valves in coll. Savigny. savigniana Audouin. IS26 (Bursatella) Fig. 99 Savigny 1817: Gasteropodes pi. 2, fig. 2 Audouin 1826: 17-18; 1828: 134 Name based on Savigny's figure. No material left. saifignyana Ehrenberg, 1831 (Helix) Savigny 1817: pi. 2, fig. 20 Ehrenberg 1831: Helix no. 9 (no page number) Name based on three specimens collected near Alexandria and Savigny's figure. Pallary however (1926: 47) argues that Ehrenberg's species is different from the one on Savigny's figure, which indeed depicts Zonites algirus (^Linne), represented by 4 shells in coll. Savigny. If Ehrenberg s types proved to be lost (they are supposedly in the Berlin Museum),. the present material appears to be formally avail- able for lectotype designation if it proves necessary for nomenclature stability. savignyanus delle Chiaje, 1828 (Murex) Savigny 1817: pi. 4, fig. 4 delle C"hiaje 1828: 222. pi. 49, fig. 32-34 Name based on Savigny's figure and additional material from southern Italy, presumably lost. One shell in coll. Savigny. See also perlatus. smngnyi Blainville, 1827 (Sepi'a^ Savigny 1817: Cephalopodes pi. 1, fig. 3 Blainville 1827: 285 Name based on Savigny's figure. No material left. Fig. 101 Fig. 45 sarngnyi Deshayes, \S44 (PlarKucis) Savigny 1817: pi. 4, fig. 29 Deshayes 1844d: pi. 109 (with two unnumbered pages of text) Name based on Savigny's figure and an unknown number of specimens from Madagascar, which could not be traced. Three shells in coll. Savigny. savignyi Deshayes, 1844 (Purpura) Fig. 30 Savigny 1817: pi. 6, fig. 1 Deshayes 1844a: 112 Name based on Savigny's figure. A single specimen in coll. Savigny is considered to be the holotype. sarignyi Jor\2LS, 1S46 (Cytherea) Fig. 15 Savigny 1817: pi. 8, tig. 17 Name based on Savigny's figure. Eight bivalve specimens and three valves in coll. Savigny. savignyi P. Fischer, \S€i5 (Cerithium) Fig. 39 Savigny 1817: pi. 4, fig. 8 Fischer 1865: 244 Name based on Savigny's figure and several specimens from Suez which could not be traced. Two shells in coll. Savigny. savignyi Vaillant, 1865 (Diplodonta) Fig. 9 Savigny 1817: pi. 8, fig. 7 Vaillant 1865: 124 Name based on Savigny's figure and several specimens from El Toueneb bank. Red sea, two of which are in MNHN. Two specimens in coll. Savigny. savignyi Issel, 18&9 (Litiopa) Fig. TO Sa\'ignv 1817: pi. 3, fig. 19 Issel 1§69: 197 Name based on Savigny's figure and ten shells from Suez, of which seven are in MGD. Ten shells in coll. Savigny. savignyi Issel, \9i%9 (Marginella) Savigny 1817: pi. 6, fig. 18 Fig. 88 Issel 1869: 115 Name based on Savigny's figure and a number of specimens from Suez, of which two are in MGD. No Savigny material left. sanqnyi P. Fischer, 1871 iPeclunculus) Fig. 3 Savigny 1817: pi. 10, fig. 14 Fischer 1871: 219 Name based on Savigny's figure and one sjiecimen from Suez, which could not be traced. Two shells in coll. Savigny. savignyi Tapparone-Canefri, IHlh (Fasciolana) Fig. 43 Savigny 1817: pi. 4, fig. 14 Tapparone-Canefri 1875: 612 Name based on Savigny's figure. A single shell in coll. Savigny is considered to be the holotype. samgiiyi Morlet, \S7S (RingieuLa) Fig. 59 Savigny 1817: pi. 6, fig. 7" Morlet"l878: 117, pi. 5, fig. 1 Name based on Savigny's figure and one shell from the gulf of Suez, now in MNHlSl. Two shells in coll. Savigny. savignyi Jousseaume, 1888 (Mesodesma) Fig. 5 Savigny 1817: pi. 8, fig. 5 Jousseaume 1888: 206 Name based on Savigny's figure and two left valves from Cameron island. Red sea, apparently lost. One specimen in coll. Savigny. savignyi Pilsbry, 1892 (CaUistochiton keterodon var.) Savngny 1817: pi. 3, fig. 8 Pilsbry 1892: 277, pi. 60, fig. 16 (copied from Savigny) Name based on Savigny's figure. It is therefore an objective synonym of Chiton affinis Issel 1869. which is based on the same figure. savignyi Monterosato, 1899 (Meleagrina) Fig. 2 Savigny 1817: pi. 11. fig. 8-9 Monterosato 1899: 392 Name based on Savigiiy's figure and many specimens from Cyprus and Alexandria, proliably in the Museo Comunale, Roma. Two specimens in coll. Savigny. savignyi Pallary, 191% (Fissurella) Fig. 50 Savigny 1817: pi. 1, fig. 5; Pallary 1926: 34 Name based on Savigny's figure and .several specimens from Suez, which could not be traced. Two shells in coll. Savigny. .•OTr/'gwV? Pallary, l'i2f>(Gastrochaenal Fig. 13 SavignV 1817: "pi. 1, fig. 15; Pallary 1926: 39 Name based on Savigny's figure and one specimen from Suez, which could not be traced. One bivalve specimen and 5 valves, some being fragmentary, in coll. Savigny. savignyi ? aWaxy , 1926 (Lioconcha) Fig. 6 Savigny 1817: pi. 9, fig. 5; PallaiT 1926: 107 Name based on Savigny's figure. One shell in coll. Savigny is considered to be the holotype. savignyi Pallary, 1926 (Nassa) Fig. 90 Savigny 1817: pi. 6, fig. 3; Pallary 1926: 87 Name based on Savigny's figure. No material left. savignyi Pallary, 1926 (Sculus) Fig. 92 Savigny 1817: pi. 1, fig. 10; Pallary 1926: 35 Name based on Savigny's figure. No material left. savigniji "?\\\\vpp\" Krauss, \^i?, (Siphonaria) Savigny, 1817: pi. 1, fig. 1; Pallary 1926:28 Krauss, 1848:61; Reeve, vol. 9, pi. 5, sp. 20. 3 syntjpes in MNHN, seguenziana Issel, 1869 (Rissoina) Fig. 82 Savigny 1817: pi. 4, fig. 3 16 THE NAUTILUS January 25, 1982 Vol. 96(1) Issel 1869: 209 Name based on Savig7iy's fig:ure. A single sl^ell in coll. Savigny is considered to be the holotype. seynperiana Issel, lS&9(Lucina) Fig. 19 Savignv 1817: pi. 8, fig. 12 Issel lS69: 82 Name based on Savigny 's figure and one specimen from Suez, which could not be traced in MCID. A single shell in coll. Savigny. sismondiana Issel, 1869 (Rissoa) Fig. 87 Savigny 1817: pi. 3, fig. ,33 Issel 1^69; 205 Name based on Savigny's figure and three shells from Suez, two of which are now in M(jD. No Savigny material left. sue2irasi'.s' Issel. 1869(A/(iryjnW;o; Fig. 60 Savigny 1817: pi. 6, fig. 17 Issel ito: 115 Name based on Savigny's figure and several shells from Suez, six of which are in MOD. Si.x shells in coll. Savigny. siigillata Jonas, 1846 (CythereaJ Fig. 8 Savigny 1817: pi. 9, fig. 3 Jonas i846a: 64 Name based on Savigny's figure and two additional shells which could not be traced. Two bivalve specimens and one valve in coll. Savigny. tiberiana Issel, 1869 (Cingula) Fig. 69 Savigny 1817: pi. 3, fig. 16 Issel 1^69: 199 ^ ^ Name based on Savigny's figure and several specimens from Suez, of which si.x are in MGD. About three dozen shells in coll. Savigny. tigrina .Audouin, 1826(Dorrs^ Savigny 1817; Gasteropodes pi. 1. fig. 3 Fig. 96 Audouin 1826: 13; 1828: 127 Name based on Savigny's figure. No material left. tninrala (Ferussac m,s) Audouin, "[S'l^lPhysa) Fig. 29 Savigny 1817: pi. 2, fig. 27 Audoum 1826: 34; 1828: 166 Name based on Savigny's figure. Two shells in coll. Savigny. There are also several shells from Syria, collectea by Bruguiere and Olivier, in the Ferussac coll. (MNHN). undata Pallary, 1926 (Risella isseii var.) Savigny 1817: pi. 5. fig. 35.3 Pallary" 1926: 84 ^ NameDased on Savigny's figure and a number of shells from Suez, which could not De traced. See under isseii. unicolor Dautzenberg, in Pallary, 1926 (Cassis turgida var.) Savignv 1817: pi. 6, fig. 6 Pallary 1926: 88 Name based on Savigny's figure and an additional shell in Dautzenberg's private collection, which could not be traced by Abbott (1968: 199). One shell in coll. Savigny. venusta Issel, 1869 (Turbonilla) Fig. 34 Savigny 1817: pi. 3, fig. 34 Issel 1869: 175 Name based on Savigny's figure and three shells from Suez, now in MGD. Nine specimens in coll. Savigny. rillae Issel, 1869 (Cingula) Fig. 80 Savigny 1817: pi. 3, fig. 17 Issel 1869: 198 Name based on Savigny's figure and six shells from Suez, four of which are in MGD. Seven shells in coll. Savigny. villersii Audouin, IS'ZG (Bulla) Fig. 53 Savigny 1817: pi. 5, fig. 4 Audouin 1826: 39; 1828: 178 Name based on Savigny's figure. Two shells in coll. Savigny. FIGS. 1-84. Specimens from the Snvinny Egyptian collec- tion yww in the Musinim National il'Hi^toire Naturelle in Paris. France. 1, Spatha caillaudi van Martens, 68 mm. 2, Melcagrina savi^nyi Monterosato, 58 mm. 3, Pectunculus savigiivi P. Fischer, 18 mm. 4, Unio bourguignati Landrin, 2.i mm. 5, Mesodesma savignyi Jousseaume, 20 mm. 6, Lioconcha savignyi Pallary, ,W mm. 7, Area pharaonis P. Fischer, J,8 mm. 8, Cytherea sugillata Jona^, 3J, mm. 9, Diplodonlii savignyi Vnillant, 28 mm. 10, Mylilus pharaonis P. Fischer, .12 mm. 11, Lithodomus lessepsianus Vaiilant, 10 mm. 12, Mylilus coenobita Vaiilant, 7 mm. 13, Ga.sirochaena savignyi Pallary, 11 mm. 14, Cardium richardi Audouin. 10 mm. 15, Gytherea savignyi Jona.s, 35 mm. 16, Cardium isthmic'um Issel, SJ, mm. 17, Cardium mareoticum Pallary, 2J, mm. 18, Chama corliierei ,)onas, .H7 mm. 19, Lucina semiieriana Issel, 4 mm. 20, Lucina erythruea Issel, 16 mm. 21, Helix favroi Landrin, 12 mm. 22, Trochus liemprielii Issel, 2 mm. 23, Helix leroii Landrin. 7 mm. 24, Turbo cr(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'ptiaca Ehrenberg, 6.5 mm. 62, Scissurella reticulata Philippi, 1.75 mm. 63, Scissurella dorbignii Audomn, 1.75 mm,. 64, Risella infracostata hsel, 1.15 mm. 65, Risella isseii St-niper in Issel, 1.15 mm. 66, Cyclostrenia philipii /sse/, 1.6 mm.. 67, Rissoa freminvillii Audouin, 3.0 mm. 68, Rissoa dorbignii Audouin, 2.5 mm. 69, Cingula tiberiana /sse^, 1.25 mm. 70, Litiopa savignyi Issel, 2.55 mm. 71, Alaba martens"! Issel, 2.3 mm. 72, Odontostomia clysmatica Issel, 2.7 mm. 73, Eulimella ciiijjiilata Issel. 2.6 mm.. 74, Pyrgulina doliiformis Paltary. 2.1 mm. 75, Odontostomia craticulata I.'isel. 2.05 mm. 76, Rissoina rissoi Weinkauff, 3.Jt mm. 77, Cerithium bacillum Issel, 3.75 mm. 78, Eulima gentiluomiana Issel. 2.6 mm. 79, Triforis perlatus Issel, i.7 mm. 80, Cingula villae Issel, 2.0 mm. 81, Scalaria {erassa.ci\ Audouin. 1.95 mm. 82, Rissoina seguenziana Issel. 2.8 mm. 83, Pupa lamarckii Audouin, 2.2. mm. 84, Scaliola elata Semper in Issel, 2.05 mm. 87 ''%\^ 89 ■^i.^ 90 FIGS. 8.5-92. Types fiffun's copied from Savigny (no type m.aterial found in SurK/nii inlltrtion). 85, Anatola humboldti .-1 udoum, 35 mm (?). 86, Pristis audouini Jousseaume, 31 mm.. 87, Rissoa sismondiana Issel, 2.5 mm. 92 88, Marginella savignyi Issel. h mm.. 89, Emarginula cuvieri Audauin, 9.5 mm. 90, Nassa savignyi Pallary, 9 mm. 91, Rissoa desmarestii Audouin, 3 mm.. 92, Scutus savignyi Pallary, 7 mm. <■'■'■' ■" ■ ■ ;. i.-.m, 3.9 mm. FIGS. 96-103. Savigny's unpublished color vellum.s of opisthobranchs and cephalopods in Bibliotheque Centrale, MNHN. Only black and white prints were previously published. >^^--^3 t V -vi?^ 96, Doris immaculata Audouin. i6 mm. *.# %^ ■ r f. -^ %. 97, Tritonia elegans Audouin, 50 mm. 98, Doris marmorata /I Mrfoitiw, 30 mm. 99, Doris tigrina Audouin. 22 mm. 100, Doris concentrka Audouin. 21 mm. ' )-^**&::: ^ ■•i-i^' ,1 ■■ ' ap*"; 102, Bursatella savignyana Audouin. 130 mm. 101, Fleurobranchus obiongus .Audouin, 30 ?«/«, \y^ 103, Octopus horridus Orhigny. mantle length 30 mm. 24 THE NAUTILUS January 25, 1982 Vol. 96(1) FIG. 94. Blainvillia erythraea Hnpe. 58 mm. FIG. 95. Sepia savignyi BlainviUe, mantle length 85 mm. From an unpublished color vellum in the Bibliotheque Cen- trale, MNHN. LITERATURE CITED Abbott. R. T. 1968. The Helmet shells of the World (Cassi- dae). Part 1. Indv-Paafic Mollusca 2(9):7-202, pi. 1-187. Audouin, V. 1826. Explication sommaire des planches de Mollusques de I'Egypte et de la Syrie publiees par J. C. Savigny. Description de I'Egypte ou recueil des observa- tions et des recherches qui ont ete faites en Egypte pend- ant I'expedition de I'armee fran^aise, publie par les ordres de sa majeste I'empereur Napoleon le grand. Histoire Naturelle, Animaux invertebres l(4):7-56. Imprimerie imperiale, Paris. 1828. (same title), 2 erne edition, dediee au roi, 22:117-212. Panckouke, Paris. [1827 on title page is in- accurate]. BlainviUe, H. M. Ducrotay de 1827. Dictionnaire des sciences naturelles, 48. Levrauft, Strasbourg. 572 pp. Chenu, J. C. 1845. Illustrations Conchyliologiques, vol. 2: Bivalves marins. Genus Tridaena. 2 pp., 8 pi. Paris. Chiaje, S. delle 1828. Memorie sulla storia e anatomia degli animali senza vertebre del regno di Napoli, 3:1-232, pi. 31-49. Napoli. Deshayes, G. P. 1844a. Histoire naturelle des animaux sans vertebres, 2eme edition, 10. Baillere, Paris. 638 pp. , 1844b. (no title). Mag. Zool.. Mollusques: pi. 109 (with two unnumbered pages of text). Ehrenberg, C. G. 1831. Animalia Mollusca, in: Hemprich & Ehrenberg: Symbolae Physicae, IV: Animalia Everte- brata. Unnumbered pages. 3 plates. Fischer, P. 1865. Note sur les launes conchyliologiques des deux rivages de I'isthme de Suez. Jour. Conchyt. 13: 240-248. 1870. Sur la faune conchyiiologique marine des baies de Suez et de I'Akabah. Jour. Conchyl. 18:161-179. 1871. Sur la faune conchyiiologique marine de la bale de Suez. Joar. Conchyl. 19:209-226. Fischer, II. 1901. Liste des coquilles recueillies par M. de Gennes a Djibouti et Ali-Sabieh, avec la description de plusieurs formes nouvelles. Jour. Conchyl. 49:96-130, pi. 4. Hup^, H. 1854. Note sur un nouveau genre de Mollu.sque acephale, genre Blainvillie (Blainvillia). Re^>. Mag. Zool., (2)6:219-224. Issel, A. 1869. Malacologia del Mar Rosso. Ed. della Biblio- teca Malacologica, Pisa. 387 pp., 5 pi. Jonas, J. H. 1846a. Beitrag zur Erklarung der in der Des- cription de I'Egypte abgebildeten, nebst Beschreibung einiger anderer in rothen Meere und den angrenzenden Landem lebender Mollusken. Z. Malak. 3:59-64. 1846b. Unbeschriebene Konchylien des rothen Meeres. Z. Malak. 3:65-67. 1846c. Beitrag zur Erklarung der in der Des- cription de I'Egypte abgebildeten . . . Z. Malak. 3:120-127. Jousseaume, F. 1888. Description des Mollusques recueillis par Mr. le Dr. Faurot dans la mer Rouge et le golfe d'Aden. Mem. Soc. Zool. France 1:165-223. Krauss, F. 1848. Die sildafrikanischen Mollusken. 140 pp. Stuttgart. Laissus, Y. 1973. Napoleon et rimprimerie. Description de I'Egypte; bilan scientifique d'une expedition militaire, iyi L'art du Livre a I'lmprimerie Nationale: 190-205. Im- primerie nationale, Paris. 295 pp. Lamy, E. 1918. Les Tellines de la mer Rouge (d'apres les materiaux recueillis par M. le Dr. Jousseaume). Bull. M2(.s. natn. hist. nat. 24:26-33. Landrin, A. 1865. Coquilles nouvelles. Aubert, Versailles. 6 pp., 1 pi. Martens, E. von 1866. Uebersicht der Land- und Siisswasser Mollusken des Nil-Gebietes Ma/.aA-. Bl. 7:1-21. Monterosato, A. di 1899. Coquilles marines de Chypre. Jour. Conchyl. 47:392-401. Morlet, L. 1878. Monographie du genre Ringicula Deshayes et descriptions de quelques especes nouvelles. Jour. Conchyl. 26:113-133, pi. 5. Nevill, G. and H. Nevill 1875. Descriptions of new marine Mollusca from the Indian Ocean. Jour. Asiatic Soc. Bengal 44(2):83-104, pi. 7-8. Orbigny, A. d' 1826. Tableau methodique de la classe des Cephalopodes. Ann. Sci. Nat. 7( separate reprint). Pallarv, P. 1926. Explication des planches de J. C. Savigny. Mm. Inst. Egypte ll:Viii -h 138 pp., pi. 1-18. 1931. Marie Jules-Cesar Savigny. Sa vie et son oeuvre. Premiere partie. Mem. Inst. Egypte 17:1-106. Deuxieme partie (1932). Ibid. 20:1-112. 'Troisieme partie (1934). Ibid. 23:1-202. 1932. Inventaire de la collection malacologique de Savigny. Bull. Mus. natn. hist. nat.. 2nd ser., 4(3): 313-321. ■ 1933. Les sources d'information concernant les savants et artistes de I'Expedition d'Egypte. Bull. Inst. Egypte 15:221-228. Philippi, R. A. 1853. Die Gattungen Delphinula, Scissurella und Globulus. Systematisches Conenylien-Cabinet von Martini und Chemnitz ("second" edition), 2(4):l-57, pi. 1-8. Von Bauer & Raspe, Niirnberg. Pilsbry, H. A. 1892. Manual of Conchology 14: Polyplaco- phofa. 350 pp., 68 pi. Philadelphia. Savigny, J. C. 1817. Description de I'Egypte, ou recueil des observations et des recherches qui ont ete faites en Eg^'pte pendant I'Expedition de I'armee fran^aise, publiee par ordre du gouvernement. Histoire naturelle. Planches, vol. II. Imprimerie royaie, Paris. 1826. (same title), 2 eme edition, dediee au roi. Pjinf rCouf kp P urm Sherborn, C. D. 1897. On the Dates of the Natural History portion of Savigny's 'Description de I'Egypte'. Proc. Zool. Soc. Lond.. 1897:285-288. Ta|iparone-Canefri, C. 1875. Studio monografico sopra i Muricidi del Mar Rosso. .Ann. Mus. Civ. St. nat. Genova 7:564-640, pi. 19. Tr>'on, G. W. 1886. Manual of Conchology 8. Philadelphia. Vaillant, L. 1865. Recherches sur la faune malacologique de la bale de Suez. Jour. Conchyl. 13:97-127, pi. 6. Weinkauff, H. C. 1885. Die Gattungen Ris.'ioina und Rts.wa. Systematisches Conchylien-Cabinet, 2nd ed., l(22):l-205, pi. 1-25. Bauer & Raspe, Nuremberg. Vol. 96(1) January 25, 1982 THE NAUTILUS 25 SHELL MICROSTRUCTURE OF CORBICULA FLUMINEA (BIVALVIA: CORBICULIDAE)i Clement L. Counts, III and Robert S. Prezant College of Marine Studies University of Delaware Lewes, DE 19958 U.S.A. ABSTRACT The shell microstructure ofCorbicula tluminea was examined using scanning electron microscopy. The interface between the periostracum and calcareous shell. and a marginal periostracal loop, are described with a discussion of possible func- tions. Surface morphology of shells with periostracum chemically removed re- waled periodic concentric surface ridges. The complex crossed-lamellar crystalline arrangement of the shell is confirmed. Shell structure of bivalves in the genus Cor- bicula (Miihlfeld, 1811) has been examined by several investigators (Tsujii, I960; Toots and Fox, 1972; Taylor et al., 1973; Mackie, 1978). Tsujii (1960) reported that three shell layers (periostracum, prismatic and pearl [nacreous of other authors]) are present in C. japonica Prime, 1864, collected from brackish waters of the Tose River at Tsu, Japan. Taylor et al. (1973) reported that shells of Cor- bicula flumin-ea (Miiller, 1774), C. occidens (Deshayes, 1854), C. cuneiformis (Sowerby, 1817) and the fossil species C. cordata (Morris, 1854) are composed totally of aragonitic calcium carbonate. Toots and Fox (1972) found that aragonite, as is typical of this mineral through geologic time, had metamorphosed to calcite in fossil shells of Corbicula (Leptesthes) fracta collected in the Upper Cretaceous strata of Wyoming. The outer calcareous shell layers of Corbicula fluminea and C. occidens are arranged in a finely crossed lamellar pattern, the inner layers are complex crossed lamellar and an indistinct myo- stracum occurs at the pallial line (Taylor et al., 1973). The complex crossed lamellar layer of C. fluminea and C. occidens consists of laths ar- ranged in regular columns as in the Limopsacea (Taylor et al., 1969). 'University of Delaware College of Marine Studies Contribu- tion No. 155. Mackie (1978) examined the shell structure and mineralogy of Corbicula fluminea in a com- prehensive ultrastructural study of the fresh- water Sphaeriacea. Corbicula fluminea has the thickest shell of the 23 species of sphaeriaceans studied. Mackie reported that the individual layers of the complex crossed lamellae, which are found in all Sphaeriacea, are inclined in op- jx)site directions. No pallial myostracum, nacre- ous or outer prismatic layer is present. Mackie (1978) postulated that thicker-shelled species of sphaeriaceans have correspondingly thicker periostraca that protect the shell from dissolu- tion in acid waters of organically enriched habitats in which they are found (Ingram et al., 1953; Mackie and Qadri, 1973). No single study has carefully examined the ultrastructure of the interior and exterior sur- face features of the shell of the Asiatic clam. The present paper presents such an analysis of the shell of Corbicula fluminea. using scanning elec- tron microscopy. The outer surface morphology is emphasized. MATERIALS AND METHODS Specimens of Corbicula fluminea were col- lected, through the kindness of Dr. Ralph W. Taylor, from Tygarts Creek, Carter County, Kentucky, in October 1978. The bivalves were first relaxed in 7% magnesium chloride or pro- pylene phenoxytol solutions. Periostracum was tlien removed with a 5% sodium hjTJOchlorite 26 THE NAUTILUS January 25, 1982 Vol. 96(1) solution. Shells were washed in distilled water, dehydrated through 70% ethanol and placed in a hot air oven (73°C) for 48 hours to dry. Shells were then fractured and shell fragments were selected from the posterior and anterior adduc- tor muscle scars, the hinge and ligament, the growing edge, the outer surface near the grow- ing edge, the pallial line, and the purple inner surface. Dried shell fragments were mounted on aluminum SEM pin stubs with double adhesive tape and silver paint and stored in the hot air oven (73°C) until coated with metal. Fractured shells with intact periostracum were dehydrated in a series of increasing con- centrations of ethanol solutions to 100% in which they were stored. Periostracum was also prepared on whole shells by fixation in 3%i gluteraldehyde for 7 days. Shells were fractured and fragments placed in 50% ethanol (5 min.) and 100% ethanol (28 hrs.). All specimens were dried in a Denton DCP-1 critical point drying ap- paratus using carbon dioxide as a transfer agent. Specimens were then mounted on pin stubs. Mounted specimens were coated with a thin layer of gold in a Denton Vacuum Evaporator and examined in a Philips PSEM 501 SEM. RESULTS A well-developed periostracum covers the calcerous shell oi Corhicula jlmrdnea. This layer consists of an extremely smooth external sur- face, is internally homogenous, and averages 10 ym in thickness. It is firmly conjoined with the underlying aragonitic crystals. The periostracum terminates along the ven- tral shell margin in a reflective loop before in- FIG. 1. Pi'notitnicat loop al vetitral shell margin. Uirectiun of growth to the top. Hortzonlal field width = i!55 piw. gressing to the secretory mantle (Fig. 1). Thus, upon emergence from the mantle, the periostra- cum remains disattached from internal calcer- ous shell and extends up to 580 \j.m beyond the shell edge in a loop that often bends back over the outer shell margin. Where the periostracum closely overrides the outer shell surface, the at- tachment is very tight and rarely is this organic layer seen to peel away from the calcareous ex- oskeleton. Ultrastructural examination of fractured shell fragments confirms the complex crossed lamel- lar structure reported by Taylor et al. (1973) and Mackie (1978) (Fig. 2). Bands of crystalline laths are arranged approximately normal to each other. Individual layers or bands have an average thickness of 10 ^im (Fig. 3) while in- dividual crystalline laths are approximately 0.7 \jin in diameter. A cursory examination of the outer surface oi shells from which periostacum has been re- moved with sodium hj'pochlorite reveals annula- tions or major ridges visible to the unaided eye that are separated by fine, concentric striations FIGS. 2-7. 2, Radially fractured shell o/ Corbicula fluminea demo iist rat iri.y complex crossed lamellar structure. Direc- tion of growth lo the right. Horizontal field width = 900 lim. 3, Normally arranged crystalline laths composiyig the com- plex crossed lamellar shell. Direction of growth to the right. Horizontal field width = 8 (jto- 4, Outer shell surface. periostracum removed, showing concentric major annulationis (arrows), unequally spaced, and finer, subannular concen- tric striae helween major annulae. Direction of growth lo the right. Horizontal field width = 5.3 mm. 5, Shell surface, perkmlnwum removed, in fractured specivwn. Two growth zones are found in association with the major annulae (ar- rows). Direction of growth lo the right. Horizontal field width = 2.J, mm. 6, Growth zones demonstrating rugose, perforate zones terminating in a thick band of shell, the periphery of which apparently marks the end of a growth peri.od (A), a smooth hand oj shell that becoines rugose, perforate, and grooved and then smooth, terminating in a small ridge (B) and a wide band of nigi.se. perforate shell (C). Direction of growth to the left. Horizontal field uidth = .'ISO urn.. 7, Rugose, per- forate bond oj oilier shell surface sculpture organized into radial ridges and grooves with surj'ace perforatiuris. Direction of growth In the nght. U'jrlzonhil field width = 82 \im. Vol. 96(1) January 25, 1982 THE NAUTILUS 27 (Fig. 4). Under higher magnification (Fig. 5) three major striations are often found in association with each large concentric annula- tion. The surface morphology of a single major ridge system (Fig. 6) changes with growth, showing different patterns of rugosity that ap- Vol. 96(1) January 25, 1982 THE NAUTILUS 28 pear to be associated with cessation of growth and that alternate with smooth areas at the ap- parent beginning of renewed shell growth. The largest region of rugose shell was found in areas preceeding an apparent growth halt (Fig. 7). This region demonstrated an overall pattern of perforations and grooves that are radially oriented. The grooves were either interrupted or anastomosing and had an average width of 1-2 fjim. Perforations within this zone (Fig. 8) were of various sizes, ranging from the ir- regularly ovoid largest holes measuring 0.75 x 1.00 nm at their greatest dimensions, to the smallest, more circular holes with an average diameter of 0.5 j^m. The largest perforations usually occurred at the bottom of the radial grooves. The small perforations were usually found on walls bordering grooves. No perfora- tions penetrated very deeply into the interior of the shell (Fig. 9). This rugosity is a natural oc- currence on the calcareous shell of Corbicula fiwminea and is not an artifact caused by chemical removal of the periostracum. This is evidenced by the alternating smooth areas. The zone of perforated grooves terminates in a narrow (approximately 20-30 f.im) band of thick, radially arranged microstructures (Fig. 10). This calcareous band is continuous with the FK.S. 8-10. o, hiny.-ijroove sculplnre ikrminstrntiHn varying Kizes of perforations. Direction of yrowth to the right. Horhotit'it fielil u-idlh = li) nm. 9, FrnHitrc section of shell with periostracum removed. Note .luperficinl perforations. Direction of ijrowlh to the right. Horizontal field width = S2 tim. 10. Thick, concentric hand (D) terwinatimj the ridge- groon sralplurc. \,w shell (E) appears to be laid down from beneath this band. Direction of growth to the lejl. Horizontal field ic'dth = I,,-; ii7,i. Vol. 96(1) January 25, 1982 THE NAUTILirS 29 FIG. 11. Thr 5 fjiiii ridge (B) xcpdnilutg smoolh bam! (F) from rugose band (C) of shell. Perforation's of the. rugose band are r}ot organized into ridge-groove patterns at this stage. Tile sntiMith hand has few. scattered perfdrolnois. Dinclidii of groiiili III Hie right. Hnrizniiliil field ivnllli = to jj/h. radial-perforate zones, and has few perfora- tions. The leading edge of this zone apparently represents the end of a growing period. A concentric ridge averaging 5 ytm in width was found 110 ^m from the band of apparent growth cessation (Fig. 6). Between these two points, three transitional surface patterns were found; a band of smooth surfaced shell averag- ing 25 [im in width with diffuse perforations, a 25 fim-wide band of grooved, perforate surface similar to that preceeding the thickened calcium band before the apparent growth pause, and a 60 yim wide band of smooth surfaced shell that terminates in the 5 ^^m-wide ridge. Immediately following the 5 \xm ridge, the shell surface becomes more rugose with perfora- tions averaging 2-4 \xm in diameter (Fig. 11). These perforations are heterogeneous but gradually gain organization as they meld with the ridge-groove pattern found later in the growth period (Figs. 6, 7, and 8) and terminate in the thickened surface sculpture (Fig. 10). DISCUSSION The periostracum tightly fills the irregular contours of the underlying calcareous shell sur- face as shown in Figin-e 10. Presumably the periostracum has fully polymerized prior to the initiation of calcium carbonate deposition on this organic template (Waite and Wilbur, 1976), so that this intimate association is more likely a result of aragonitic nuclei being deposited in rugosities of a preformed periostracal mold. In any event, the tight fit between periostracum and calcareous shell typically leaves an intact outer organic layer over the aragonitic shell and the only evidence of periostracal wear occurs around the umbones. The marginal periostracal looj) is an interest- ing morphological feature whose function is unclear. The periostracal extension might allow free range of motion of the mantle so that upon withdrawal of the mantle the periostracum would easily follow. Thus upon adduction perio- stracum would form an effective seal at the shell edge. The curvature of the periostracal loop, if maintained during mineralization might offer a mold for formation of calcareous concentric shell ridges. Loop size corresponds well with that of major concentric ridge size, lending sup- port to this hypothesis. Extensive periostracal folding along the mantle margin occurs com- monly in many marine bivalves [for example. Area zebni (Waller, 1980)]. "Excess" marginal periostracum is probably a reflection of perio- stracal growth exceeding development of cal- careous shell. This phenomenon still requires careful analysis. Morton (1977) found that populations of Cor- bicula Jluminea in Plover Cover Reservoir, Hong Kong, spawn twice a year. He pointed out that shell growth appeared to stop during these episodes as energy is redirected to gonadal development. The first spawning in the spring of the Plover Cover population was the most in- tensive, followed by a less intensive one in the fall or late summer (Morton, 1977). Although Morton (1977) reported no shell surface changes that correspond to spawning seasons, if metabo- lism is redirected so that only gonads are developed and shell growth ceases, this halt should be reflected in the surface morphology of the shell. Fuziwara (1975, 1978) reported Cor- hirula leana spawns biannually in response to thermal stimuli and while portions of the gonad ai'e mature throughout the year, the overall rate of gametogenesis is unknown. Jones (1981) recently reported variations in shell growth in- crements that correspond to thermal variation 30 THE NAUTILUS January 25, 1982 Vol. 96(1) in the marine environment. The pattern of .sur- face shell sculpture seen in the Tygarts Creek, Kentucky, population may be the result of redirected energetics during gametogenesis and change in surface sculpture seen in Figure 10 represents the terminal period of shell growth before gonadal development or a response of the bivalve to fluctuations in thermal regime. ACKNOWLEDGMENTS We thank Dr. Ralph W. Taylor, Department of Biological Sciences, Marshall University, Huntington, West Virginia, for his assistance in the collection of the bivalves used in this study. We are also indebted to Dr. Melbourne R. Car- riker, College of Marine Studies, University of Delaware, Lewes, for his review of the manu- script and Dr. Gerald L. Mackie, Department of Zoology, University of Guelph, Ontario, for his comments and discussion. Appreciation is also extended to Mr. Walter H. Denny, Department of Mechanical and Aerospace Engineering, Uni- versity of Delaware, Newark, for his technical assistance, Ms. Pam Palinski for her photo- graphic help, and Ms. Linda Leidy for typing the final versions of the manuscript. LITERATURE CITED Fuziwara, T. 197.S. On the reproduction of Carhiculii Icana Prime. Venus 34(l):54-56. 1978. On the ovulation n{ Ciirhiruin lennn Prime. VV?!«.s 37(l):22-28. Ingram, W. M., U. G. Rallinger and A. K. Gaul'in. 19.'):i. Ke- latitmship of Sphaerium nolidum Prime to organic pollu- tants. Ohio J. Sci. 53(4):230-235. Jones, D. S. 1981. Annual growth increments in shell.s of Sjiisula soLidissima record marine temperature varia- bility. Science 211(4778):165-166. Mackie, G. L. 1978. Shell .structure in freshwater Sphae- reacea (Bivalvia: Heterodonta). C'nuidian Jour. ZooL 56(l):l-6. Mackie, G. L. and S. V . Qadri. 1973. Abundance and dis- tribution of Mollusca in an industrialized portion of the Ottawa River near Ottawa - Hull, Canada. Jour. Fish. Res. Bd. Canada. 30:167-172. Morton, B. 1977. The population dynamics of Corbicula fluminea (Bivalvia: Corbiculidae) in Plover Cove Reser- voir, Hong Kong. Jour. ZooL. London, 181(l):21-42. Taylor, .J. D., W. I. Kennedy and A. Hall. 1969. The shell structure and mineralogy of the Bivalvia. Introduction. Nuculacea - Trigonacea. Bull. British Mus. (Nat. Hist.). ZooL Suppl. 3. pp. 1-125. 1973. The shell structure and mineralogy of the Bivalvia. II. Lucinacea - Clavagellacea. Conclusions. Bull. British Mus. (Nat Hist.) ZooL. 22(9):2,55-294. Toots, H. and J. E. Fox. 1972. Inversion of aragonite to calcite in Corbicula (Leptesthes) fracta from upper Creta- ceous strata of Wyoming. Contrib. Geol. 12:11-14. Tsujii, T. 1960. Studies of the mechanism of shell and pearl formation in molluscs. Jour. Fac. Fish.. Prefect. Univ. M(V 5:1-70. Waite, J. H. and K. M. Wilbur. 1976. Phenol oxidase in the periostracum of the marine bivalve Modiolus demissus. Jour. Exper. ZooL. 195:359-367. Waller, T. R. 1980. Scanning electron microscopy of shell and mantle in the order Arcoida (Mollusca: Bivalvia). Smithsonian Cuntr. ZooL 313: 58 pp., 46 figs., 1 table. A RELIC POPULATION OF OBOVARIA RETUSA IN THE MIDDLE CUMBERLAND RIVER, TENNESSEE Paul W. Parmalee and Walter E. Klippel Department of Anthropology University of Tennessee Knoxville, TN 37916 ' ABSTRACT Recovery in 1980-1981 of three specimens o/Obovaria retusa (Lamarck, 1819) from cull piles left by commercial shelters provides additional evidence of an ex- tant relic population of this naiad in the middle Cumberland River, Trousdale County, Tennessee. The fornior dir-triliution of Obovaria retusa (Lamarck, 1819) included the Ohio, Cumber- land, and Tennessee River systems. This mus- sel, called by the common name Golf Stick by Vol. 96(1) January 25, 1982 THE NAUTILUS 31 Boepple and Coker (1912) and known variously as Pink, Ring Pink, Ram's Horn Pink, Pink Pigtoe or Rosebud by commercial shellers, typically inhabits the deep stretches of a river having swift current and a substrate composed of coarse sand and gravel. Obovaria retusa has been extirpated throughout most of its former range. Stansbery (1970) stated that "A popula- tion still living in the impounded lower Ten- nessee [below Pickwick Dam] had apparently not reproduced since impoundment and is ex- pected to die out. The only known breeding population of this once widespread species is a small one in the Green River near Munfordville, Kentucky." Stansbery has since commented (Personal Communication, October 1980) that the Green River population appears to be no longer viable and very possibly has been entirely eradicated. Commercial shellers still occasion- ally take a large old relic individual from stretches of the Tennessee River below Pick- wick Dam, but the animal is extremely rare judging by the limited numbers encountered. Prior to impoundment, 0. return occurred throughout the Cumberland River "although by no means abundant anywhere" (Wilson and Clark, 1914). However, Neel and Allen (1964). in their study of the mussel fauna of the upper Cumberland River basin before impoundment, found it to be fairly common on the big stream bars below the falls. 0. rptui^n was not found during a collection made by Stansbery (1969) after impoundment of the same general area surveyed by Neel and Allen (1964). Tennessee Valley Authority biologists (TVA, 1976), how- ever, reported finding the Ring Pink (numbers and location not given) at a sheller's cook-out area during their mussel survey of the middle Cumberland River in Wilson, Trousdale, and Smith counties (CRM 270.0-305.0). Although a few valves of 0. retusa were recovered from two prehistoric Indian rock shelter middens (Wood- land Period, c. 1000 B.C. - A.D. 1000) in Smith County by Parmalee, Klippel and Bogan (1980), no fresh specimens were taken by brailing or found in sheller's cull and stock piles during their three year middle Cumberland River naiad survey in Smith County. Examination of a series of cull and stock pile belonging to a commercial sheller operating from a location along the Cumberland River (CRM 275.2), c. 6.5 km south of Hartsville, Trousdale County, Tennessee were made No- vember 14, 1980 and January 8 and September 17, 1981. In addition to the various piles of shell, sorted on the basis of species, size, and/or nacre color, several hundred naiads had been dis- carded at the water's edge. It was in this area that the first specimen of 0. retusa was found; it was a very large female (in mm: length, 83; height, 79, breadth, 50) and, by all appearances, old with a heavily eroded umbo and slightly deformed posterior margin (Figure 1). It is very similar in size and apparent age to those speci- mens taken in the Tennessee River below Pick- wick Dam. The second specimen, another fe- male and represented by the right valve, was recovered January 8, 1981 from a large cull pile of "pinks" (£'?h'pfio spp., Cyclonaias tuberculata, Lampsilis orhiculata, Epioblasma sulcata). Measurements (mm) of this valve are as follows: length, 64, height, 60, estimated breadth of paired valves, 40. During re-examination of these abandoned cull piles (no commercial shelling in this area since summer 1980) on September 11, 1981, the / • VU'i. 1. Sf)ecirnens o/ Obovaria retusa taken hy commercial shelters. .>iummer 1980. in the Cumberland River. Trouxdale County, Tennessee. Top shell 8.3 mm in length. 32 THE NAUTILUS January 25, 1982 Vol. 96(1) left valve of another female was found. This specimen was slightly larger than the one repre- sented by a right valve and measured 71 mm in length, 63 in height, and estimated 44 mm in breadth of paired valves. Diligent searches failed to locate the other valves of these two in- dividuals (Figure 1). Admittedly three specimens or even several do not represent a viable population, but they do provide evidence for the continued existence of 0. retusa in the middle Cumberland River. Although the two smallest of the three speci- mens appear to be considerably younger than the large individual, the outer rest-lines are crowded and too obscure to establish even an estimated age with any degree of certainty. Their smaller size and the lesser extent of umbo erosion suggests individuals somewhat younger than the larger female, but it has been found that in the case of certain species (e.g. Epiobbtsma brevidens. Cyprogenia irrorafa) or individuals that impoundment has the effect on naiads of causing the production of extremely heavy and thick but stunted shells (Parmalee. Klippel, and Bogan, 1980). The assemblage of naiad species from this sec- tion of the river (CRM c. 273.0-278.0) where the specimens of 0. retusa were taken exhibits some interesting differences from the one reported by Parmalee. Klippel, and Bogan (1980) at CRM 291.0-296.8. Although many species such as Lampsilis orbiculata appear to occur in about the same frequency in the stretch of river below Hartsville as they do upstream some 16-18 km, others, including Actinonaias ligamentina and Ptychobranchvisfasciolare, are much rarer in oc- currence. In contrast, Quadrula quadruLa ap- pears in greater numbers below Hartsville. It is of interest to note that a relic population of Epioblasma sulcata continues to survive in the Cumberland River below Hartsville; we sal- vaged 38 specimens (all males) of this endan- gered naiad from the cull piles. It is evident from the paucity of specimens taken during the past five years that 0. retusa continues to exist only as a relic population in the middle Cumber- land River and that, as Stansbery (1970) com- mented regarding the population of this naiad in the lower Tennessee River, it can be expected to die out. ACKNOWLEDGMENTS We thank Miles Wright, Frank H. McClung Museum, University of Tennessee, Knoxville, for photographing the specimens in Figure 1 and Betty W. Creech for typing the manuscript. LITERATURE CITED Hiiepple, J. V. and R. E. Cuker. 1912. Mussel resources ol ihe Holston and Clinch rivers of eastern Tennessei-. Bureau of Fisheries Document No. ?65.'3-13. Neel, Joe K. and William R. Allen. 1964. The mussel fauna of the upper Cumberland Basin before its impoundment. Malacologia l(.3):427-4.59. Parmalee, Paul W., Walter E. Klippel and Arthur E. Bogan. 1980. Notes on the prehistoric and preserit status of the naiad fauna of the middle Cumberland River, Smith Coun- ty, Tennessee. The Nautilus 94(3):93-in5. Stansbery, David H. 1969. Changes in the naiad fauna of the Cumberland River at Cumberland Falls in eastern Kentucky. The Ann^r-ican Malacol'igical l.'nion Annual Reports for 1969. pp. 16-17. 1970. Eastern freshwater mollusks (1) the Mis- sissippi and St. Lawrence River systems. In; Proceedings of the Ajnerican Malafological Union Sympostum on Rare and Endangered Mollusks. Malacologia 10(l):9-22. Tennessee Valley Authority. 1976. Mussel fauna of the Cumberland River in Tennessee. Unpub. Report: Div. Environmental Planning and Div. Forestry, Fisheries, and Wildl. Develop. Wil.son, Charles B. and H. Walton Clark. 1914. The mussels of the Cumberland River and its tributaries. Department of Commerce, Bureau '^'^'V^^>^SA^^^^^^^^i^«^«^iM^^^«^i^i^ american malacologists ff-Ti P O BOX 2255 (305) 725-^iO \^ MELBOURNE, FLORIDA 32901 , U.S A PUBLISHERS OF DISTINCTIVE BOOA'.S' ON MOUMSKS Abbott, R, Tucker, editor. THE BEST OF THE NAUTILUS. 1976 288 pp An exciting and nostalgic anthology of stiell collecting in ttie Nlneteentti Century ISBN 0-91 5726-02-X Clotfi $13.95 Aral Vol. 96(2) April 21, 1982 THE NAUTILUS 35 JOSEPH CHARLES BEQUAERT 1886-1982 Dr. Joseph C. Bequaert, Agassiz Professor of Zoology emeritus of the Museum of Compara- tive Zoology, Harvard University, died in Amherst, Massachusetts, at the age of 95, after a lingering illness, on January 12, 1982. Dr. Bequaert was born in the small town of Thourout, near Bruges, Belgium, and as a schoolboy was interested in plants and mollusks. In 1908 he received his Ph.D. in botany from the University of Ghent, and he spent the rest of his long and productive life studying many facets of the natural world. His first entomological field work was in the Belgian Congo where he was Entomologist for the Belgian Sleeping Sickness Commission from 1910 to 1912. He stayed on in Africa after the Commission gained control of the disease by isolating its victims, and between 1913 and 1915 he continued his botanical work, being in charge of botanical explorations for the Belgian Colo- nial Government. During his time in Africa he collected mollusks, many of them new, from the Congo River, throughout the jungles to the mountainous Ugandan border. This material was studied by Dr. Henry A. Pilsbry and formed the basis of his two-volume work on the mol- lusks of the Belgian Congo, the second volume of which was written in collaboration with Dr. Bequaert. In 1916 Joe Bequaert immigrated to the United States, and by 1921 he was a naturalized citizen. His first position in the U.S. was at the American Museum of Natural History as Re- search Associate in Congo Zoology. He left the American Museum in 1922 for Boston, where he was first Instructor (1923-1925) and then Assis- tant Professor in the Department of Tropical Medicine at the Harvard Medical School, 1925-1945. In 1927 he married Frances Brown of Ohio; they had two children, Helen and Frank. After his retirement from Harvard in 1956, he sought warmer climes and went to the Univer- sity of Texas as a Visiting Professor in biology. In 1960 he moved to Tucson, Arizona as he was Visiting Entomologist and Curatorial Assistant at the University of Arizona, where he orga- nized the mollusk collection and continued to collect and study the little known land and fresh- water snails of the area. His last paper was on the "Mollusks of the Arid Southwest" with Walter B. Miller, published in 1973. In 1980 he and his wife Frances moved to Amherst, Massachusetts, to be near their daughter. Dr. Helen Holmes. He died there at the age of 95, on January 12, 1982. Dr. Bequaert's prolific bibliography (over 250 papers, more than 50 of them on mollusks) at- tests to his broad background and interests. His work includes medical research on sleeping sick- ness and schistosomiasis, extensive molluscan studies, particularly of the Achatinidae and Strophochelidae, many works on wasps and other insects, botanical papers, and even a paper on the use of the fauna of putrefaction to determine time of death. He was a member of many scientific societies, president of the American Malacological Union in 1954, and an Honorary Associate in Entomology and Mala- cology at the Museum of Comparative Zoology from the time of his retirement. Dr. Bequaert, affectionately called Uncle Joe by the many young students then at the MCZ, was available to advise and help at all times. He was an in- credible linguist (said he grew up on the streets of Belgium speaking four languages) and was always willing to help translate difficult passages in any of the European languages. Dr. Bequaert's contribution to malacology and to the Mollusk Department of the MCZ will be appreciated by students for many generations. Memorial contributions to support research in land molluscan systematics may be given to either the Friends of the Department of Mol- lusks, Museum of Comparative Zoology, Har- vard University, Cambridge, MA 02138 or the American Malacological Union, 3706 Rice Boulevard, Houston, Texas 77005. William J. Clench, Curator Emeritus Department of Mollusks Museum of Comparative Zoology 36 THE NAUTILUS April 21, 1982 OXYCHILUS DRAPARNALDI IN IOWA Vol. 96(2) Terrence J. Frest and R. Sanders Rhodes II Department of Geology University of Iowa Iowa City, lA 52242 ABSTRACT Feral and greenhouse colonies o/Oxychilus draparnaldi (Beck, 1837) in Iowa City, Johnson Co., Iowa constitute the first Iowa record for an introduced land snail. The species has now been reported as feral in 13 states and in greenhouses in 13 states and the District of Columbia. The feral colony was established in or prior to 1966, occupies approximately 2,U00 sq. ft., and contains several hundred individuals. Differences in land snail faunas in adjacent areas with and without 0. draparnaldi, plus feeding experiments, suggests that 0. draparnaldi reduces local populations of some land snail species. Records of introduced mollusks in Iowa are extremely rare. Barnhard (1978) lists two slugs and one fresh-water snail: as far as we know this paper constitutes the first record for an intro- duced land snail in Iowa. In addition, the Iowa City 0. draparnaldi occurrence is of signifi- cance because it is the only Midwest feral colony and because some information as to date of in- troduction (and hence rate of spread) is available. In the course of a systematic survey of the land snails of Johnson County, Iowa in 1976-78 a single dead specimen of 0. drapanaldi was recovered from a drift pile along a fence in the northwest corner of Hickory Hill Park, a city facility in northeastern Iowa City. Subsequent intensive searches for the species resulted in the discovery of living specimens in July 1978 on the west side of a ravine just southwest of the south terminus of St. Clements Street in Iowa City (SEV4 NWV4 SWV4 SWV4 sec. 2,T. 79 N., R. 6 W., Iowa City West IVz' topographic quadran- gle: Fig. 1). Repeated visits to the site were made in March - August 1979 and 1980 to collect 0. draparnaldi and define the limits of the col- ony. Litter and drift samples were taken from the colony and adjoining areas to assess the ef- fects, if any, of the introduction. A laboratory colony was maintained for one year to check feeding preferences. Also a visit made to one local greenhouse (Pleasant Valley Orchards and rv-^' '■■:'■• OAK LAN P fMfT^Y \ FIG. 1. Portion uflow' topoyrtiiiliic iptadran- =i— Barra Palma Sola ^-1- - Punta Los Munecos r^ iPunla Boca Andreal ^ -—Punta Delgada (Punta Las Literas) ■Jt - Punta Limon } 1 — Punta Villa Rica a' \ 96125 \ FIG. 1. Mapofth: ii-ealnyiGulfofMexiroshov'tngllu'Piintit del Morro-PwnXa r>etg\ernments of Tokyo Prefecture and Miyake- ima for financial and logistic support during his study. The efforts of J. A. Dalton, J. L. )icksved, G. Hodgson, T. Takada, K. Tateyama, 1. Umeda and M. J. Zaiser in field observations nd data collection are greatly appreciated. R. uiiidall is thanked for sharing observations of h-iipeUa predation at Guam. Dr. C. Birkeland itidly offered expert advice and supplied impor- int literature. Drs. R. T. Abbott, H. A. Rehder, Is. B. W. Myers, and Mr. A. D'Attilio gener- usly provided information on the taxonomy of the Thaidinae and C. J. Ferraris, Jr. on Philip- pine corals. Dr. J. Rosewater kindly supplied a needed reference. W. E. Old, Jr. and S. S. Horenstein and M. Werner, respectively, pro- vided bibliographic assistance, photographed the specimens (Figs. 1-3), and tj-ped the manuscript. We are indebted to Dr. R. Robert- son for critically reading the manuscript. We also thank P. Kott and M. Cope. LITERATURE CITED .\rakawa, K. Y. 1958 ("1957"). On the remarkable sexual dimorphism of the radula of Drupella. Venus (Japanese Jour. Malacol.) 19(3-4):206-214. Birkeland, C. (In press). Terrestrial runoff as a cause of out- breaks of Acanthaster planci (Echinodermata: Asteroi- dea). Mar. EcoL, Progress Series. Blainville, H. M. D. de. 1832. Disposition methodique des especes recentes et fossiles des genres Pourpre, Ricinule, Licorne et Concholepas de M. de Lamarck. Nouv. Ann. Mus. Nat. d'Hist.. Paris 1:189-263. B(jrn, I. 1778. Index Rerum Naturalium Musei Caesarei Vindobonetisis. Pt. 1: Testacea. Vienna, xiii + 458 p. Branham, J. M. 1973. The crown of thorns on coral reefs. BioScwnce. 23{4):219-226. Cernohorsky, W. 0. 1969. The Muricidae of Fiji, Pt. 2, Sub- family Thaidinae. The Veliger 11(4):293-315. 1972. Marine Shells of the Pacific. Pacific Pub- lications, Sydney, 411 p. 1978. Tropical Pacific Marine Shells. Pacific Publications, Sydney, 352 p. Coe, W. R. 1956. Fluctuations in populations of littoral marine invertebrates. Jour. Mar. Res. 15:212-232. Cooke. A. H. 1895. The Cambridge Natural History. Mol- luscs, vol. 3. pp. i-ix, 1-459. Macmilian & Co., New York. Demond, J. 1957. Micronesian reef-associated gastropods. Pacific Sci. 11(3):275-341. Emerson. W. K. and W. 0. Cernohorsky. 1973. The genus Drupa in the Indo-Pacific. Indo-Pacific Mollu.tca 3(13):801-864. Endean, R. 1973. Population explosions of Acanthaster planci and associated destruction of hermatypic corals in the Indo-West Pacific region. In: Biology and Geology of Coral Reefs, vol. 2, Biology 1, Jones, 0. A. and R. Endean (eds.). Academic Press. New York, p. 390-438. Fanklioner, P. V. 1970. Notes on feeding and the functional morpholog\' of the gut in the reef dwelling muricid Morula elata (Blainville). Western Soc. Malacologists. Abst. & Proc. for 1969, pp. 15, 16. Glynn, P. W. 1976. Some physical and biological determi- nants of coral community structure in the eastern Pacific. Ecological Monographs 46:431-456. Goreau. T. F., J. C. Larg, E. A. Graham and P. D. Goreau. 1972. Structure and ecologj- of the Saipan reefs in relation to predation by Acanthaster planci (Linnaeus). Bull. Mar. Sci. 22(1):113-152. Ida, H. and J. T. Moyer. 1974. Apogonid fishes of Miyake- 82 THE NAUTILUS April 21, 1982 Vol. 96(2) jima and Ishigaki-jima, Japan, with description of a new species. Japan. Jour. Ichthyol. 21(3):113-128. Melville, R. V. [Sec] 1980. International Commission on Zoological Nomenclature. Opinion 1154, Dnipella Thiele, 1925 (Mollusca, Gastropoda): Designation of a type species by use of the plenary powers. Zooi Norrmi. Bull. 37(2):85-88. Moyer, J. T. 1978. Crown of thorns starfish invades Miyake- jima.' Marine Park Jour. (Japan) 44(1): 17 (in Japanese). Ponder, W. F. 1973. The origin and evolution of the Neo- gastropoda. Malacologia 12(2):295-338. Porter, W. P. 1972. Predation by Acanthaster and its effect on coral species diversity. Amer. Natur. 106(950): 487-492. Randall. J. E. 1972. Chemical pollution in the sea and the crown-of-thorns starfish (Acanthaster planci). Biotropica 4(3): 132- 144. Reese, E. S. 1977. Coevolution of corals and coral feeding fishes of the family Chaetodontidae, p. 267-274. Third International Coral Reef Symposium Proc. vol. 1, Univ. Miami. Miami, Florida. Robertson, R. 1970. Review of the predators and parasites of stony corals, with special reference to symbiotic proso- branch gastropods. Pacific Sn. 24(l):43-54. Shepard, J. W. and J. T. Moyer. 1980. Annotated checklist of the fishes of Miyake-jima, Japan. Pt. 1, Pomacentridae, Chaetodontidae, and Pomacanthidae. Publ. Seto Mar. Biol. Sta. 15(l-4):227-241. Sutcliffe, W. H., Jr. 1972. Some relations of land drainage. nutrients, particulate material, and fish catch in two east- ern Canadian bays. Jour. Fish. Res. Bd. Canada 29(2): 357-362. 1973. Correlations between seasonal river dis- charge and local landings of American lobster (Homarus americanus) and Atlantic halibut (Hippoglossus hippo- glossus) in the Gulf of St. Lawrence. Jour. Fish. Res. Bd. Canada 30(6):856-859. Taylor, J. B. 1975. Planktonic prosobranch veligers of Kaneohe Bay. Ph.D. Diss. Univ. Microfilms Inst., Ann Arbor, Michigan, 331 p. Taylor, J. D. 1976. Habitats, abundance and diets of muri- cacean gastropods at Aldabra Atoll. Linnean Soc. Zool. (London) Jour. 59(2):155-193. 1980. Diets and habitats of shallow water preda- tory gastropods around Tolo Channel, Hong Kong. In: The malacofauna of Hong Kong and southern China, Morton, B. (ed.) Univ. Hong Kong Press, p. 163-180. 1978. Habitats and diet of predatory gastropods at Addu Atoll, Maldives, Exp. Mar. Biol. Jour. 3(1): 83-103. Taylor, J. D., N. J. Morris and C. N. Taylor. 1980. Food specialization and the evolution of predatory prosobranch gastropods. Paleont. (London) 23(2):375-409. Tribble, G. W. and R. Randall. [MS]. Description of coral environment and community structure of the shallow water corals of Miyake-jima, Japan. Wu, S.-K. 1965. Studies of the radulae of Taiwan muricid gastropods. Inst. Zool., Acad. Sinica Bull. 4:95-106. SUCCINEA AVARA SAY FROM THE SOUTHERN GREAT PLAINS OF THE UNITED STATES Dorothea S. Franzen Illinois Wesleyan University Bloomington, Illinois 61701 ABSTRACT Succinea avara Sa?/, 182Jt, was described from shells collected from the North- west Territory. This study is o/S. avara collected at 21 stations in four states of the Southern Great Plains Region of the United States. Shells and anatomical structures are described and illustrated; habitats of a selected representative series from the 21 stations are described. Succinea avara Say, 1824 Sucnnea avara Say, 1824, p. 250; pi. 15, Fig. 6. Succitiea avara Say, Pilsbry, 1948, pp. 837-840; Fig. 442. B, b, C, c, D. Shells of Succineidae present only a few dis- tinctive characteristics; specific determinations depend largely upon anatomical characteristics of the soft parts. It is questionable whether the anatomy of the soft parts as now identified with a species actually corresponds to the anatomy of the soft parts of the holotypes and paratypes which are known only from their shells. It is im- V^ol. 96(2) April 21, 1982 THE NAUTILUS 83 possible to determine the exact location of the type locality when geographically it is desig- nated only as being in a general area, as iorSue- Hnea avara Say, "Inhabits the Northwest Terri- :ory" (Say, 1824, p. 260). The succineas taken Tom a locality today may not be representative )f the fauna existing fifty to a hundred or more /ears ago. Such are the problems concerning ;he identification of S. avara Say. The shells of the holotype and paratj^aes of '^uccinea avara, Academy of Natural Science Philadelphia No. 59542, are small and probably mmature. ^ype description: "S. avara. Shell suboval, pale reddish-yellow, subdisphanous, fragile, covered with an earthy crust; whorls three, minutely wrinkled; body whorl very large; spire small; aperture large, subovate, about two-thirds of the whole length of the shell. Length three-twentieths of an inch. Inhabits the North-west Territory. This small species of Succinea occurs in humid places very frequently under stones and near the water. The shell is always completely in- crusted with a coating of earth. It may at once be distinguished from either the ovalis or of campestris by its very small size. PI. 15, fig. 6." (Say, 1824, p. 260.) Pilsbry (1948, pp. 837-840, Fig. 455 and Fig. 42, A, a, B, b, C, c, D) described what he con- idered to be S. avara. Fig. 442, B, b, are draw- igs of the reproductive system of a specimen rom Warm Spring Canyon, Arizona, collected y Ferriss in 1909; C, c, are of a specimen from lurricane Fault, Arizona. The purpose of this paper is to describe shell nd anatomical features, as well as habitats, of uccinea avara Say of the Southern Great 'lains of the United States. Collections were lade by Artie L. Metcalf and by me at 21 sta- ons located in the western half of Texas, east- rn and southern New Mexico, southern and orthwestern Oklahoma and southwestern Kan- as (Fig. 1). Succineidae are generally regarded as being mphibious- living on wet ground, in woods, long shores, or appearing on the surface allowing a rainfall heavy enough to soak the round. However, we have sometimes found Fid. 1 . Four-state map shomng locations of the 21 collecting sidlionti. them in habitats far removed from wet areas. At the stations where S. avara was obtained, the annual precipitation ranges from about 9 to 44 inches. All stations are subjected to periods of from 3 to 5 months in which the precipitation is less than an inch per month (Yearbook of Agriculture, 1941). S. avara included in this study were taken in early spring, summer, and early winter months during which time the snails were either active or aestivating. They were found in damp places under rocks, logs, boards or dead stems of Yucca sp. The following are the locations of the 21 collecting sites and descriptions of habitats of a selected representa- tive series often of those sites. The field number designations and descriptions of habitats are by Artie L. Metcalf. My field numbers, enclosed in parentheses, serve as a reference to my records. Sites of my collections are indicated by my field numbers and the descriptions of such habitats are mine. Stations 1. Field No. ALM 968 (DSF 402); 30 June, 1969, along the Cimarron River, along New Mexico Rd. 325; 7.5 mi W of its junction with New Mexico Rd. 370, Union County. New Mex- ico. Altitude: 5,250 ft. above mean tide (AMT). Annual precipitation about 17". Habitat: under logs on moist floodplain of the river and also under rocks on adjacent hillside. 2. Field No. ALM 1058 (DSF 412); 30 ,Iuly, 1970; 2.5 mi N of Mt. Dora, Union County, New Mexico. 84 THE NAUTILUS April 21, 1982 Vol. 96(2) 3. Field No. ALM 1060 (DSF 41 1); 31 July, 1970; valley of North Canadian River, 0.1 mi N of bridge over river on New Mexico Rd. 370; about 22 mi N NW of Clayton (S 33, T 29 N, R 52 W), Union County, New Mexico. 4. Field No. ALM 673, DSF .342; 8 May, 1966; .5 mi SE of Rincon; HW^ 1-25 at junction with Engle Road, Dona Ana County, New Mexico. Altitude: 4370 ft. AMT. Annual pre- cipitation about 9.5". The locality is in Jornada del Muerto- a basin between two mountain ranges. Habitat: Succinea avara were aestivating on dead stems of Yucca data Engelm. growing in a "tobosa grass swale" where the predominant plant is tobosa grass, Hilaria mutica (Buckl.) Benth. The snails were found on the side of the dead Yucca stems lying on the ground where there was a trace of moisture. 5. Field No. ALM 948 (DSF 398); 19 November, 1968: valley of Tularosa River and falls, 0.5 mi SW of bridge over Tularosa River, U.S. Hwy 70; 8.5 mi SW of Tularosa (NW '/i, SW 'A, S 32, T 13 S, R 11 E), Otero County. New Mexico. 6. Field No. ALM 946 (DSF 397); 19 November, 1968; Mescalero Indian Reservation, Valley of the Tularosa River, 1.05 mi S of Tularosa Spring (SW 'A.SW 'A. S 7, T 14 S, R 13 E), Otero County, New Mexico. 7. Field No. ALM 998 (DSF 407); 11 October, 1969, and 19 September, 1970; junction of Pehasco and Cox Canyons (SW 'A, S 3, T 17 S, R 13 E), Otero County, New Mexico. .•Mtitude: 7200' AMT. Annual precipitation about 22". Habitat: grassy, riparian meadow where the Penasco River is about 5 ft. wide. The canyon wall is wooded with pre- dominantly Ponderosa pine. 8. Field No. (DSF 492); collected by Artie Metcalf 23 September, 1978; east-central part of Nash Draw Basin (center of S 15, T 22 S, R 30 E), 20 mi E of Carlsbad, Eddy County, New Mexico. 9. Field No. ALM 670; 8 February, 1966, and DSF 339, 7 May, 1966; Franklin Mountains (S 27, T 28 S, R 4 E), about 5 mi NE Canutille, El Paso County, Texas. Altitude: 4900 ft. AMT. Annual precipitation about 9". Habitat: under dead Dasylirion sp. and under moist limestone rocks. A light rain- fall which occurred the first of May resulted in certain areas being relatively moist. 10. Field No. ALM 1396 (DSF 426); 9 August, 1972; 0.6 mi SW of Dalhart on southeast side of U.S. Hwy 54 and bordering Rita Blanca Creek, Hartley County, Texas. 11. Field No. ALM 1283 (DSF 418); 19 June, 1971; 2.3 mi W Channing, south side of Texas Farm Rd. 767, Hartley County, Texas. 12. (DSF 389); collected by Artie Metcalf 23 August, 1968; flood plain of Wolf Creek, 1 mi E of dam on Lake Fryer, Ochiltree County, Texas. 13. (DSF :',90); collected by Artie Metcalf 23 August, 1968; bluffs along north shore of Lake Fryer, Ochiltree County. Texas. Altitude: 2650 ft. AMT. Annual precipitation about 19.31". Habitat: under loose limestone below scarp in Ogallala Formation. 14. (DSF 392); collected by Artie Metcalf 13 September, 1968; bluffs on southeast .-side of Buffalo Lake, 3 mi S of Um- barger, Randall County. Texas. 15. Field No. ALM 1801 (DSF 476); 20 March. 1977; 9 mi ENE or 6.6 mi W of junction of Ranch Rd. 2084 (to Christoval) with Texas Hwy 29. S side of Texas Hwy 29. Schleicher County, Texas. 16. Field No. ALM 1451 (DSF 430); 16 April, 1973; east side of conical hill immediately E of old Fort Lancaster; N of U.S. Hwy 290, Pecos County. Texas. Altitude: 2075 ft. AMT. Annual precipitation about 19". Habitat: under lime- stone rocks on E side of conical hill. The vegetation consists of sparse grass and low xeric shrubs. 17. Field No. ALM 1458 (DSF 428); 18 April, 1973; under limestone rocks opposite Judge Roy Bean Visitor Center, Langtry, Val Verde County, Texas. 18. Field No. DSF 422; 14 July, 1972; Buffalo Bayou, Texas Hwy 6, 1 mi S of junction with HWY I-IO, near Ad- dicks, Harris County, Texas. Altitude: 50 ft. AMT. Annual precipitation about 44". Habitat: Succinea avara found aestivating on a trunk of a pecan tree up to 7' above ground level and on a fence rail 5' above ground level. The ground vegetation comprised nettles and grasses which may have been too dense for the succineas; therefore, they sought more open areas. 19. Field No. DSF 387, 28 July, 1968, and 17 July, 1972; west bank of a bay of Lake Texoma, University of Oklahoma Biology Station, 2 mi E of Willis and Oklahoma State HWT 99, Marshall County, Oklahoma. Altitude: about 500 ft. AMT. Annual precipitation about 38". Habitat: the snails were aestivating under boards and logs lying on the un- ' shaded bank of the lake; temperature at time of collection, 95°F. 20. Field No. ALM 1066 (DSF 414); 1 August, 1970; 6 mi S Turpin; IV2 mi N of Beaver Creek, Beaver County, Okla- homa. Altitude: 2600 ft. above AMT. Annual precipitation I about 18". Habitat: under stones of bluff of Ogallala Forma- tion. The area is a short grass prairie with Yucca glauca Nut tall. 21. Field No. ALM 1559 (DSF 442); 1974; Point of Rocks, an escarpment of Ogallala Formation, 3 mi W of Kansas Hwy 27 on N side of Cimarron River, Morton County, Kan- sas. Annual precipitation about 16.2". Habitat: most of the Succinea avara were under a single large stone lying at base of scarp. Yucca glauca NuJt. and Rhus trilobata Nutt. were common plants. Grama grass and annual ragweed were abundant. The Shell: Shell is conically-ovate, imperfo- rate, amber, somewhat glossy, translucent, usu- ally encrusted with grains of sand or soil. Com- posed of 2V2 to 3% whorls, the shell attains a height of 12.6 mm and a width of 6.6 mm (Table 1). Whorls inflated, sharply incised, increasing rapidly in size from the knoblike nuclear whorl to the large, inflated body whorl. Aperture ovate, ranging from 47.6% to 77.7% of the height of the largest shells obtained at the 21 stations, bounded by a sharp peristome which Vol. 96(2) April 21, 1982 THE NAUTILUS 85 TABLE 1. DtmermoTis of shells o/Succinea avara Say of selected ten oftli£ tumity-one collecting sites. The nwtisurements are of the three largest shells of each locality. Ratios of measurements are listed in the last four columns. No. of Width/ Height of Width of H. Ap / W. Ap / W. Ap station #1 Whorls Height Width Height Aperture Aperture H. Shell W. Shell H. Ap Union Co. , NH 3 1/4 12.6 mm 5.45 mm .432 b . 0 mm 4.05 mm .476 .743 .675 30 June, 1969 3 1/4 10.9 6.3 .577 7.3 4.45 .669 .706 .609 3 1/4 10.0 5.8 .580 8.15 5.1 .815 .879 .625 ^Range (23 shells) 2 3/4 - 3 1/2 6.0- 12.6 3.3 - 6.3 .432 - .689 3.9 - 8 15 2.7 - 5 35 .476 - .815 .706 - .955 .606 - .809 Median 9.10 5.15 .58 6.05 4.05 .684 .818 .683 Station #4 Dona Ana Co. , NH 3 1/4 12.3 mm 6.6 mm .536 7.0 mm 4. 7 mm .569 .712 .671 3 May, 1966 3 1/2 11.3 5.9 .522 6.2 4.2 .548 .711 .677 3 1/4 10.9 5.9 .541 6.2 4.15 .568 .703 .669 Range (29 shells) 3 - 3 1/2 6.65-12.3 3.7 - 6.6 .514 - 664 4.4-7 0 2.5-4 7 .541 - .654 .589- .762 .606 - .739 Median 8.9 5.25 .573 5.44 3.75 .604 .709 .672 station #7 Dtero Co. , NM 3 1/2 9.7 mm 5.2 mm .536 5.65 mm 3. 6 mm .582 .692 .637 11 October, 1969 3 1/2 9.6 5.55 .578 6.2 4.3 .64 5 .774 .693 3 3/4 8.9 4.9 .550 6.1 3.9 .685 .795 .639 ^ange (39 shells) 2 1/2 - 3 3/4 4.15 - 9.7 2.55 - 5.55 .530 - 644 2.9 - 6 2 2.0 - 4 3 .561 - .708 .692 - .876 .571 - .807 'ledian 6.90 3.95 .587 4.55 3.05 .662 .744 .660 Station #9 ;l Paso Co. , TX 3 1/4 12. 1 mm 6. 3 mm .520 6. 75 mm 4.2 mm .551 .666 .622 8 Feb. ; 7 May, 1966 3 1/2 11.7 6.0 .512 6.2 3.9 .529 .650 .629 3 1/4 10.7 5.8 .542 6.35 3.9 .593 .672 .614 ■ iange (58 shells) 2 1/2 - 3 1/2 4.2 - 12.1 2.5-6.3 .404 - 595 2.7 - 6 75 1.95 - 4.2 .528 - .739 .637 - .981 .573 - .756 ledian 8.25 4.35 .537 4.85 3.2 .598 .728 .65 Station #13 ■Jchiltree Co. , TX 3 1/2 11.40 mm 5.6 mm .491 6.85 mm 4. 55 mm .601 .813 .664 23 August, 1968 3 3/4 11.20 6.05 .540 6.45 3.95 .575 .652 .612 3 1/2 11.15 6.1 .530 6.6 4.2 .592 .689 .636 aD, 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) 24 8 - 13 - 1 - 11 - 8 32 8 - 13-1-12-10 49 5 - 12 - 1 - 16 - 6 71 12 - 9-1-11-11 80 13 - 9-1-12-11 88 THE NAUTILUS April 21, 1982 Vol. 96(2) The amber-colored jaw is illustrated in Fig. 4B. Anteriorly the collar has a pointed median fold and posteriorly a rounded indentation. The posterior margin of the basal plate has a pointed median fold flanked on either side by a rounded boss. FIG. 4. A, Representative radula teeth of Succinea avara Say. C, central tooth; 1-L-L, first left lateral; 1-L-M, first left marginal; 2-L-M, second left marginal; 7-L-M, seventh left marginal; 8-L-M, eighth left marginal. B, A jaw o/Succinea avara Say. length of the basal plate; mesocone is flanked on either side by a small ectocone. Laterals have a long, sharply pointed mesocone which, gener- ally, exceeds length of the basal plate; there is a single smaller ectocone; endocone is generally wanting. Differentiation between laterals and marginals is not always sharply defined; fre- quently an endocone is present on the outermost one to three laterals. There may be on either side of a lateral with a split ectocone a tooth with an undivided ectocone. Marginals are char- acterized by a short endocone, a long, sharply pointed mesocone, and an ectocone which is divided into two, three or four cusps. In 5. ovalis and S. vaginacontorta the three small cusps of the ectocone are short and almost equal in size. In S. avara the two medial-most cusps of the ec- tocone are long; the outermost cusps are minute (Fig. 4A). The basal plates of the marginals are as in S. ovalis Say (Franzen, 1959, Fig. 3) and as inS. vaginacontorta Lee {Franzen, 1971, Fig. 3). They are not long and tapering as in the genus Oxyloma (Franzen, 1963, Fig. 1) nor as short and broad as in the genus Catinella (Quick, 1933, Fig. 4); (Franzen, 1979, Fig. 3). ACKNOWLEDGMENTS National Science Foundation Grants-in-Aid No's. NSF G18000 and NSF GB2715 provided laboratory equipment and supported, in part, field studies. The Academy of Natural Sciences of Philadelphia graciously lent shells of the holotype and paratypes of Succinea avara Say. Dr. A. Byron Leonard read the manuscript and offered helpful suggestions. LITERATURE CITED Franzen, Dorothea S. 1959. Anatomy of Succinea ovaltK Say. Proc. Mai. Soc. London 33(5):193-199, Tables MI, Figs. 1-7. 1963. Variations in the Anatomy of the Succi neid Gastropod Oxyloma retusa (Lea). The Nautilus 76(3):82-95, Tables MI, Figs. 1-4. 1971. Anatomy and Geographic Distribution of the Succineid Gastropod Succinea vaginacontorta Lee. The Nautilus 84(4):131-142, Tables MI, Figs. 1-3. 1979. Catinella parallela. a New Succineidae (Puimonata) From Midwestern United States. The Nauti- lus 93(2-3):63-69, Tables 1-2, Figs. 1-3. Pilsbry, Henry A. 1948. Land Mollusca of North America (North of Mexico). Acad. Nat. Sci. Philadelphia Mon. No. 3. Vol. II, Ft. 2: xlvii + 521-1113, 585 figs. Quick, H. E. 1933. The Anatomy of British Succinea. Proc. Mai Soc. London 20(6):295-318, Tables I-V, PI. 23-25, Figs. 1-18. Say, Thomas. 1824. In Appendix to Keating's Narrative Expedition . . . source of St. Peter's River, etc. (North- west Territory) 2:260, PI. 15, fig. 6. Yearbook of Agriculture. 1941. Climate and Man. United States Department of Agriculture, U.S. Government Printing Office, Washington, D.C.: v-xii and 1-1248. INFORMATION FOR SUBSCRIBERS The annual subscription rate for The Nautilus is $12.00 for individuals (foreign $14.00) and $18.00 for institutions (domestic or foreign). Subscrip- tions may begin in January. Send check or money order made out to "American Malacologists" to the Business Manager, P.O. Box 2255, Melbourne, Florida 32901, U.S.A. Back issu£s from volume 72 to date are ob- tainable from the Business Manager. 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FL 32901) Mail: Box 2255, Melbourne, FL 32901 Second Qass Postage paid at Melbourne, Florida and other post offices Subscription Price: $12.00 (see inside back cover) $14.00 (foreign); institutions $18.00 THE NAUTILUS Volume 96, number 3 - July 26, 1982 ISSN 0028-1344 CONTENTS Eileen H. Jokinen Cipangopaludina chinensis (Gastropoda: Viviparidae) in North America, Review and Update 89 Hans Bertsch and Barbara Myers Comments on the Eastern Pacific Trivia ritteri (Gastropoda: Triviidae) 96 Bretton W. Kent An Overlooked Busycon Whelk (Melongenidae) from the Eastern United States 99 M. G. Harasewych and Richard E. Petit Notes on the Morphology ofCancellaria reticulata (Gastropoda: Cancellariidae) 104 William Miller, III and Sheree Kooser The Distribution of Donax variabilis (Bivalvia) at Ship Island, Mississippi: I Some Paleomalacological Aspects 113 J. Gibson-Smith and W. Gibson-Smith Divarilima (Bivalvia: Limidae) and a New Subspecies from the Caribbean 115 J. Gibson-Smith and W. Gibson-Smith The Subfamily Melampinae (Pulmonata: Basommatophora) in Venezuela, with Descriptions of Two New Species 116 J. Gibson-Smith and W. Gibson-Smith An Ancestral Stephopoma (Mollusca: Gastropoda) from the Tertiary „ of Venezuela 120 K. E. Hoagland and Wesley R. Coe Larval Development in Orepidula maculosa (Prosobranchia: Crepidulidae), from Florida 122 Errata 96 COMPENDIUM OF SEASHELLS An Authoritative Guide to over 4,000 Marine Shells of the World in Color by R. Tucker Abbott and S. Peter Dance Two internationally known experts on mollusks have combined their knowledge and writing talents to produce the largest and most complete, all-color, general identification shell book of this century. Dr. Abbott brings his expertise of forty years at the Museum of Comparative Zoology at Harvard College, The Smithsonian Institution and the Academy of Natural Sciences of Philadelphia, while S. Peter Dance contributes his years of experience as a curator at the British Museum of Natural History and the National Museum of Wales. Both authors are world-renowned for their research and many popular books on malacology. Now comes their monumental COMPENDIUM, a pictorial summary of the world's most conspicuous marine species. A LMCJLE FORMAT! Every species is illustrated with its own separate color plate, with all the important data at hand, immediately below it: RESERVE YOUR COPY FOR CHRISTMAS NOW 1. Scientific name, with author and date. October $50.00 (04854-1460), cloth 2. Vernacular English name NATURAL hlSTORY/COLLECTlMQ 4,300color plates 3. Geographical occurrence and depth range Bibliography Index 8Ve x 10^4 400 pages 4. Relative abundance or rarity. \SE^r\: 0-525-93269-0 DUTTOH ^. Habitat. 6. Important synonyms. 4.(l()() Species - 4.()()() cnl<.r phitcs Without this book you would need a shell librar>' costing many hundreds of dollars to identify all these 4.000 species Furthermore, the COMPENDIUM illustrates ^00 species not found in any other popular shell hooks Types lllustnited Hundreds of unique holotypc specimens from Harvard University, the Smithsonian Institution and other American and European museums are shown in full color for the first time, in addition to many beautiful rarities previously hidden away in museum drawers (iuide to tile Litenitiire The COMPENDIUM gives you the leading source-books and indices. Hundreds of monographs are listed under their appropriate genera and families, and a long list of faunal books, useful to local collectors. is arranged by countries. By delvmg into the books in this bibli- ography you can find the answer to almost every molluscan question Every species in the COMPENDIUM is supplied with its author and ihc date it was described, thus enabling researchers lo track each one down. Full (^i\er:iiie The great popular families are fully covered, including double views of 181 species of cowries. 534 cones. 152 volutes. 140 scallops. 75 nutmegs (Cancellariidae) and all the Strombidae, Cassidae. Pleuroiomariidae, Neruidae. etc EXCLUSIVE OFFER ! A handsome bookplate has been designed for the maiden pubHcation of the COMPENDIUM. Your numbered copy with your name, will be personally autographed by both authors. Make check or money orcjer payable to; American Malacologists , Inc - P O Box 225 5 Melbourne, FL 32901. USA Yes , please mail mc an exclusively numbered copy of the COMPENDIUM OF SEA SHELLS , autographed (as I indicate below ) by R. Tucker Abbott and S . Peter Dance. lencbse $5 0.00, plus $1 .5 0 for postage (foreign ad- dress add $2 .5 0; Florida residents, please add 5 % sales tax). No discounts for clubs or multiple orders. Vol. 96(3) July 16, 1982 THE NAUTILUS 89 CIPANGOPALUDINA CHINENSIS (GASTROPODA: VIVIPARIDAE) IN NORTH AMERICA, REVIEW AND UPDATE Eileen H. Jokinen Ecology Section, U-42 The Univeristy of Connecticut Storrs, CT 06268 ABSTRACT The Asian freshwater viviparid snail. Cipangopaludina chinensis (Gray), was introdiwed into North America in the 1890's. The species has spread across the United States and southern Canada and is well-established in the northeastern U.S. Connecticut populations are limited to lakes and ponds of medium-hard to hard waters of calcium levels in excess of 5 ppm. The shell growth ofC. chinensis is allometric with the young snails having a lower shell height to shell width ratio than the adults. The growth patterns and radular cusp number of the typical chinensis and the japonicus morph differ from, each other. A literature review notes the five-year life span, diatom diet, the importance of quality food sources in regulating population parameters, and the role ofC. chinensis as a possible host for Asian helminths.- The Asian freshwater prosobranch snail Cipangopaludina chinensis (Gray, 1834) (Synonym; Viviparus chinensis malleatus (Reeve, 1863)) was first introduced into North America in the 1890's (Abbott, 1950). Since then, malacologists have been documenting the spread of populations of C. chinensis. This species has become well-established in parts of North America and it is time to review the known North American biogeography, ecology and natural history of C. chinensis. This paper will also add information on the snail's en- vironmental preferences, radular structure, and shell growth pattern. Dundee (1974) and Clarke (1978) have syno- nymized C. japonicws (von Martens, 1861), with C. chinensis. Clench & Fuller (1965) believed that the two were different species with in- dividuals of C. japonicus having more elongate shells (higher height/width ratios) with smaller apical angles than C. chinensis. Morphological intergrades exist (Clench & Fuller, 1965), indi- cating either hybrid populations of two species, or different morphological types within one species. Apparently Clarke (1978) has decided upon the latter explanation but extensive com- parative morphological and immunological studies have not been published. Therefore, this paper will differentiate, when possible, between topical C. chinensis and C. japonicus by in- dicating "japonicus" forms as "japonicus" morphs, thereby cautiously following a middle ground that may easily be assimilated into a final resolution of the problem. C. chinetisis was first brought to North America in the early 1890's by sailors returning from Yokohama, Japan. The sailors sold the snails to merchandize dealers in San Francisco's Chinatown (Wood, 1892a, 1892b; Stearns, 1901). Although Wood called these specimens "japoni- ca", the snails were actually C. chinensis, the first introduced japonicus not being recorded until 1911 (Hannibal, 1911; see discussion in Clench & Fuller, 1965). A second west coast in- troduction was reported by Rev. G. W. Taylor who found "japonica" being sold in a Chinese market in Victoria, British Columbia (Anony- mous, 1894). It is not known whether these snails were of the japo7iic2is or the chinensis tj'pe (Clench & Fuller, 1965), and there has been no report of established Asian viviparid popula- tions in British Columbia. Additional west coast I 90 THE NAUTILUS July 16, 1982 Vol. 96(3) colonies have been reported from California and Washington (Clench & Fuller, 1965; Hanna, 1966; Branson, 1977). Individuals of C. chinensis were first found on the east coast by W. Clench who collected speci- mens in Massachusetts from the Muddy River, a stream dividing Boston from Brookline. The snails may have been introduced when goldfish were added to the stream for mosquito larvae control (Johnson, 1915). Additional east coast colonies have been reported from Quebec, Maine, Vermont, New Hampshire, Massachu- setts, Rhode Island, New York, New Jersey, Pennsylvania, Delaware, Maryland, North Carolina, and Florida (Johnson, 1916, 1918; Richards & Adams, 1929; Schmeck, 1942; Jacobson & Emerson, 1961; Clench & Fuller, 1965; Stanczykowska et aL, 1971; Perron & Probert, 1973; Bucci, 1974; Dundee, 1974; Clarke, 1978). Additional populations have recently been found in Connecticut (see below for details). C. chinensis populations have also spread mid- continentally in Ohio, Michigan, Wisconsin, Indiana, Minnesota, Arizona, Colorado, Texas, Iowa, Oklahoma, and Utah (Teskey, 1954; Bran- son, 1959; Clench & Fuller, 1965; Huehner & Etges, 1971, 1972; Barnhart, 1978). Figure 1 is a map of all the recorded references to North American populations of C. chinensis north of Mexico (excluding Hawaii). This species is very well established on the northeast coast where it should now be considered as a permanent part of the freshwater molluscan fauna. The species is also becoming well entrenched in the St. Lawrence-Great Lakes drainage. Is is also com- mon on the west coast between San Francisco and Seattle. Additional colonies are scattered across the continent. The paucity of mid- continental colonies may reflect a lack of infor- mation but it should be noted that some reports of recent continental malacological surveys mention no colonies of C. chinensis (Harman & Berg, 1971; Clarke, 1973). Connecticut Records Over the past several summers I surveyed 215 aquatic sites in southern New England, especial- ly Connecticut, for gastropods. Populations of typical C. chinensis have been found in seven FIG. 1. Thu \iirth American 1982 distributiov q/'Cipangopaludina chinensis (Gray). See text for references. Vol. 96(3) July 16, 1982 THE NAUTILUS 91 lakes and ponds in Connecticut and one lake in \'ermont (Lake Fairlee, Thetford Center, pre- viously reported in Clench & Fuller, 1965). A population of the japonicus-morph lives in one lake in Connecticut (Black Pond). Table 1 sum- marizes the physical/chemical data on the Con- necticut sites. Methods of water chemical analy- sis have been described elsewhere (Jokinen, 1978). Populations of C. chinensis were generally found in larger lakes or ponds with sandy to muddy sand substrates. The snails were either crawling on the substrate or on dock pilings. Even though several investigators have noted C. chinensis living in slower waters of streams (Schmeck, 1942; Stanczykowska et ai. 1971) all Connecticut sites were lentic. Due to the fact that most of the bedrock in Connecticut is of the highly insoluble schist- gneiss type, the majority of the surface waters (75% of those sampled) are very soft (calcium values below 5ppm). Yet, five of the eight C. chinensis populations were found in harder waters (Ca*-' > lOppm) and none lived in waters with Ca < 7ppm. If one statistically compares expected to actual percent occurrence of C. chinensis in soft (Ca < 5ppm) waters (56% ex- pected occurrence, 0% actual) to medium-hard to hard (Ca > 5ppm) waters (44% expected, 100% actual), using an arcsin transformation (Sokal & Rohlf, 1969), there is a statistically significant difference at the t = .02 level. This indicates that C. chinensis is more likely to be found in waters of higher calcium values than Table 1. Physical/chemical data for the habitats of Clpangopaludina chinensis in Connecticut. Lake or pond Area CO. Cond. Ca Mg Type ha S pH mgC/1 ymhos Color ppm ppm Black Pond, Middlefield, Middlesex Co. Bunnell's Pond, Bridgeport, Fairfield Co. East Twin Lake, Salisbury, Litchfield Co. Goodwin Park Pond, Wethers- NR field, Hartford Co. Holbrook Pond, Hebron, Tolland Co. North Farms Reservoir, Wallingford, New Haven Co. Tyler Pond, Goshen, Litchfield Co. Lake Wononskopomus , Salisbury , Litchfield Co. NR 30.6 11 7.3 7.6 194 .024 11.4 4.5 A 16.6 10 7.1 3.5 207 .025 9.9 3.3 N 227.6 15 7.4 33.3 286 .005 35.0 11.6 1 7.0 14.2 280 .222 19.6 4.2 NR NR 1.8 29.3 8 6.4 4.5 74 .059 7.1 2.0 25.3 73.7 7.5 8.6 158 .041 14.0 3.5 6.9 7.2 112 .030 7.8 4.1 NR 142.7 10 7.5 22.8 217 .010 26.6 8.3 Lake Type: N = natural, NR - natural, water level raised, A = artificial impoundment S = total number of snail species in lake -I- -I- Na K ppm ppm 8.2 0.5 22.0 2.9 3.9 2.1 11.9 2.0 4.8 0.3 5.5 1.8 3.1 0.5 4.5 1.6 I 92 THE NAUTILUS July 16, 1982 Vol. 96(3) lower, and that it may have a physiological need for dissolved calcium levels above 5ppm. Boycott (1936) termed mollusks which could not exist in soft water as "calciphilic". Greater sam- ple sizes and physiological tolerance testing are needed for greater resolution of required envi- ronmental calcium values. Natural history and Ecology Data on the life history pattern of C. chinensis has been published by Stanczykowska et al. (1971, 1972). They estimated the maximum life span for female snails as five years, for males, usually three years, sometimes four. All individ- uals grew throughout life, with the female's reaching larger sizes because of their longer life span. The females may begin to produce em- bryos by the end of their first year but the four and five year classes produced the most off- spring. Each female produced a minimum of 169 embryos during a life time (an estimation based on embryo counts published by Stanczykowska et aL, 1971). Crabb (1929) found up to 102 em- bryos in one female. The embryos develop in the uterus, and during spring and summer 100% of the females are carrying embryos. The young begin to appear in the population by June and continue to be born through October. The snails begin a migration into deeper water in October, the females appearing to migrate first (Stanc- zykowska et al., 1971). Quality of food affects the population para- meters of C chinertsis. Stanczykowska et al. (1972) and Plinski et al, (1978) compared snail populations from three different habitats near Montreal, the Ottawa River, a canal, and a lake. The snails fed on the bottom material which was composed of inorganics, organics and algae. The percent composition of substrate materials was identical to the percent composition of the gut contents, indicating that the snails were not selectively ingesting any specific bottom mate- rial. Most of the algae ingested were epiphytic or benthic, indicating that feeding was primarily radular and not by filtration. The ingested algae were primarily diatoms, especially species of Fragilaria, followed by greens, blue-greens, and flagellates. The percent composition of algal forms differed between the three habitats. The Ottawa River snails were larger, heavier, had higher carbon and nitrogen values, and a greater population density than snails from the other two habitats. Sediment and gut analysis revealed that the Ottawa River sediments had not only a higher percentage of algae but a higher diatom content. The lake snails made some attempt to compensate for poorer quality and quantity by consuming greater quanitities of substrate. Increase in ingestion rate with a decreased food quality has also been noted for pulmonate snails (Calow, 1975). No studies have been done to date on why diatoms appear to be more nutritious to C. chirieyisis than other algae. The entire area of freshwater gastropod feeding and nutrition is poorly explored. Allometric Shell Growth Shell growth in C. chinensis is allometric, that is, the proportions of the shell width to the shell height change throughout development. Conse- quently, the shell looks different in juvenile and adult individuals. The subject of allometric growth has been explored by Gould (1966 & 1971), White & Gould (196.5), and Jokinen (in review). Allometric growth may be analyzed by using linear regression and the resultant curves graphed. Figure 2 illustrates the allometric growth pattern of "typical" C. chinensis and compares it to the growth pattern of the japoni- CTis-morph. Shell measurements were made to the nearest 0.1 mm with vernier calipers and in- cluded embryonic shells. The raw data are avail- able from the author. The data were analyzed using the computer program SAS (Statistical Analysis System) (Barr et al., 1979) and both linear and curvilinear regressions (SAS-General Linear Models and Non-linear Models) were run on shell height vs. width and log shell height vs. log width to determine which model best fit the data. Analysis proved that shell growth in both groups is best described by a linear log shell height vs. log width curve. Log shell height was used as the independent variable. The ratio of the shell width to the shell height (W/H) decreases as the shell increases in size, the shell becoming relatively more elongate as it Vol. 96(3) July 16, 1982 THE NAUTILUS 93 18- 1 6- X I- Q 14 12- _l —I 1^10 X o 6- C. CHINENSIS » JAPONICUS- MORPH EMBRYO .4 .6 .8 10 12 14 SHELL HEIGHT 16 OQi 0 FIG. 2. Allometric shell growth o/C. chinensis and the ]a.pomcus-morph. Sketches illustrate the shape of the shells at various points on the curves. The scale bars represent 5 mm. The regression equation C. chinensis is logW = 0.81, (±0.00U S. E.jlogH + 0.17. The regression equation for the ]a.'pomais,-morph is logW = 0.78 (±0.01 S. E.jlogH + 0.23. The standard error values do not overlap, indicating that the two growth patterns are significantly different. The regression coeffi- cients (slopes) are less than one. meaning that the shell width increases in size at a slower rate than the shell height. The shells, therefore, elongate as they grow. If the slopes equaled unity, both height and width would grow at an equal rate and the shell shape would not change with increase in size. grows. The regression equation for the relation- ship between height and width for "typical" C. chiyiettms is logW = 0.84 logH + 0.17, r-square = .998. A value of less than unity for the regres- sion coefficient (slope), in this case 0.84, indi- cates negative allometry, the shell width not in- creasing as fast as shell height. The regression equation for the japonicus-morph, the popula- tion from Black Pond, is logW = 0.78 logH -i- 0.23, r-square = .997. The lower slope value, 0.78, indicates that the shell width of the japo- nicM^-morph increases even less rapidly than C. chinensis shell width. This is reflected in the relatively more elongate shells of the japonicws- morph. Young individuals of C. chinensis not only have a different shaped shell from the older snails but demonstrate marked periostracal spiral hirsuteness. When embryonic shells are placed in water or alcohol and examined under a dissecting microscope, they show a pattern of three spiral rows of stiff periostracal hairs. The hairs are distinctly hooked and the longest hairs are 0.35 mm in length. The hooks face outward from the shell. Some of the larger shells show rows of pits in the same position as the em- bryonic hairs (see Fig. 2). Since the hairs become obliterated as the snails grow post- partum, their function, if one exists, is most likely intra-uterine. The hairs might serve to hook the embryonic shells together to stabilize their positions in the uterus, or they might func- tion during birth. 94 THE NAUTILUS July 16, 1982 Vol. 96(3) Radular Structure Radulae of C. chinensis individuals were ex- amined for basic structure (Fig. 3). The teeth were separated from each other and mounted in Turtox's CMC-9AF mounting medium which contains acid fuchsin for staining chitinous material. The stain picked-up certain structural details of the central tooth where the radular material is thickened. C. chinensis has the typical taenioglossid tooth number of 7 per row. The central, first and second laterals all have a large central cusp bounded on either side by four smaller cusps. The marginal teeth do not have an enlarged cen- tral cusp but have seven small cusps (this number appears to be somewhat variable). Radular structure is similar to that of Viviparus georgianus Lea (as illustrated in Clench, 1962) except for the lack of an enlarged central cusp on the marginals of C. chinensis. An examina- tion of the radular structure of one individual of the Black Pond population {japonicus-mov\>\\) revealed five small cusps on either side of the central cusp on the first lateral tooth and nine cusps on the marginal tooth. It is not known if these differences reflect individual variation (if the two morphs are of the same species) or reflect interspecific variation. A much more ex- tensive examination of interpopulation variation is needed before real conclusions can be drawn. Parasites Information is sparse concerning the role of C chinensis as the final or intermediate host of hel- FIG. 3. Thf mthilar teeth o/C. chinensis. C = central tooth. L-I = first laivnl. LIl = I'eeond lateral. M = marginal (greatly erdargi'li. minth parasites. Penner (1942) found Massachu- setts populations to be negative for schistosome parasites. The snail may serve as a final host to the normally clam-inhabiting trematode A spido- gaster conchicola von Baer. The snails are in- fected by eating embryonated eggs passed in the feces of other snails (Michelson, 1970; Huehner & Etges, 1971, 1972, 1977). The snail also serves as intermediate host to a number of echino- stome trematodes (U.S. Dept. Agriculture, 1946), including Echinostoma cinetorchis Ando & Ozaki, a species which has been reported from humans in Japan, Taiwan and Java (Schmidt & Roberts, 1977). Angiostrotigylus cantonensis (Chen), a nematode which infects vertebrate nervous systems, may use C. chinensis as an in- termediate host (Chang et al., 1968). This parasite has been reported from Hawaii and Costa Rica (Schmidt & Roberts, 1977). Summary The introduced Asian snail, C. chinensis. does well in cool-temperate to warm-temperate cli- mates in permanent ponds, lakes and the slow parts of rivers with mud, silt or sand substrates. Available data indicate that the snail requires a minimum of 5ppm calcium in its habitat and is excluded from very soft waters. This species should be considered a well-established part of the North American fauna, especially in the northeastern U.S. Some basic studies on the natural history and ecology of this species have been undertaken in Canada but more studies should be done to de- termine what effect populations of C. chinensis are having on the native fauna. More studies need to be undertaken on the parasites carried by C chinensis in North America to determine its potential as a second- ary host to both native and introduced Asian parasites. Shell growth is allometric, the snails tending to become relatively more elongate with age. The embryonic periostracal hairs are stiff and hooked and may serve to stabilize the young snails within the uterus. The shell growth pat- tern as well as details of the radula structure dif- fer between the "typical" C. chinensis and the japonicus-morph. Whether or not these should Vol. 96(3) July 16, 1982 THE NAUTILUS 95 be considered to be intraspecific individual or in- terspecific differences awaits more extensive analysis. ACKNOWLEDGMENTS Data collection was supported by a research contract from the Department of Environmen- tal Protection, State of Connecticut. My thanks to L. Penner for loan of shells from the Muddy River. LITERATURE CITED Abbott, R. T. 19.50. Snail invaders. Ndtiirnl History 59:80-8.5. Barnhart, M. C. 1978. Three intmiluced gastropods in Iowa. The Nautilus 92:106-107. Barr, A. J., J. H. Goodnight, J. P. Sail, W. H. Blair, C. M. Chilko. 1979. SAS Users Guide. 1979 ed., SAS Institute, Inc., Raleigh, N.C., 495 p. Branson, B. A. 1977. The Chinese apple snail, Cipatigopnlu- dina chinensis. on Orcas Island. Washington. The Nauti- lus 91:76-77. Bucci, D. A. 1974. Vivipnrus malleatus in Montreal, Canada. The Nautilus 88:55. Calow, P. 1975. The feeding strategies of two freshwater gastropods, Ancylus fluviatilis Mull, and Planorhis con- tortus Linn. (Pulmonata), in terms of ingestion rates and absorption efficiencies. Oecologia (Berl.) 20:33-49. Chang. P. -K.. J. H. Cross, Jr. & S.S.S. Chen. 1968. Aquatic snails as intermediate hosts tor Angiostrongylus cantonen- sis on Taiwan. Jour. Parasitol. 54:182-183. Clarke, A. H. 1973. The freshwater molluscs of the Canadian Interior Basin. Malacologia 13:1-509. Clarke, A. H. 1978. The Asian apple snail, Cipangopaludina chinensis (Viviparidae) in (.)neida Lake, New York. The Nautilus 92:134. Clench, W. J. 1962. A catalogue of the Viviparidae of North America with notes on the distribution of Vivipariis georgianus Lea. Occas. Papers on Mollu.tks. Mus. Comp. Zool., Harvard Univ. 2(27):261-287. Clench, W. J. & S. L. H. Fuller. 1965. The genus Viviparus (Viviparidae) in North America. Occas. Papers on Mol- lusks. Mus. Comp. Zool. Harvard Univ. 2(32):385-412. Crabb, E. D. 1929. Egg laying and birth of young in three species of Viviparidae. The Nautilus 42:125-129. Dundee, D. S. 1974. Catalog of introduced mollusks of East- ern North America (North of Me.xico). Sterkiana 55:1-36. Gould, S. J. 1966. Allometry and size in ontogeny and phy- logeny. Bio. Rev. 41:587-640. Gould. S. J. 1971. Geometric similarity in allometric growth: a contribution to the problem of scaling in the evolution of size. American Naturalist 105:113-136. Hanna, G. D. 1966. Introduced mollusks of western North America. Occ. Papers Calif. Acad. Sci. 48:1-108. Harman, W. N. & C. 0. Berg. 1971. The freshwater snails of central New York. Neiv York State Agric. Exp. Sta. Search: Entomology 1(4): 1-68. Huehner, M. K. & F. S. Etges. 1971. A new gastropod host (or Aspidogaxter conchicola. Jour. Parasitol. 57:1255. Huehner, M. K. & F. S. Etges. 1972. Experimental trans- mission of A.yjidogaster conrhicola von Baer, 1827. Jour. Parasitol. 58:109. Huehner, M. K. & F. S. Etges. 1977. The life cycle and devel- opment ofAspidngaster conchicola in the snails, Viviparus malleatus and Goniobasis livescens. Jour. Parasitol. 63:669-674. .lohnson, C. W, 1915. Viviparus malleatus Reeve in Massa- chusetts. The Nautilus 19:35. Johnson, C. W. 1916. Viviparus malleatus Reeve. The Nautilus 30:48. .lohnson, C. W. 1918. Viviparus malleatus and contectoides in Massachusetts. The Nautilus 31:107-108. Jokinen, E. 1978. Habitats of two freshwater limpets {Fer- ri.<:sia: Ancylidae) from New England. The Nautilus 92:156-160. Jokinen, E. H. (In review). Allometric growth of planorbid snails. Michelson, E. H. 1970. Aspidogaster conchicola from fresh- water gastropods in the United States. Jour. Parasitol. 56:709-712. Penner. L. R. 1942. Studies on dermatitis-producing shisto- somes in eastern Massachusetts, with emphasis on the status oi Scihistosojnatium pathloc.opticum Tanabe, 1923. Jour. Parasitol. 28:103-116. Perron, F. and T. Probert. 1973. Viviparus malleatus in New Hampshire. The Nautilus 87:90. Plinski, M.. W. Lawacz, A. Stanczykowska & E. Magnin. 1978. Etude quantitative et qualitative de la nourriture des Viviparus ?NaWea/Ms (Reeve) (Gastropoda, Prosobran- chia) dans deux lacs de la region de Montreal. Canadian Jour Zool. 56:272-279. Schmeck, E. H. 1942. Viviparus malleatus in Niagara River. The Nautilus 55:102-103. Schmidt, G. D. and L. S. Roberts. 1977. Foundations of Parasitology. C. V. Mosby Co., St. Louis., 604 pp. Sokal, R. R. and F. J. Rohlf. 1969. Biometry. W. H. Freeman & Co.. San Francisco, 776 pp. Stanczykowska, A., E. Magnin and A. Dumouchel. 1971. Etude de trois populations de Viviparus malleatus (Reeve) (Gastropoda. Prosobranchia) de la region de Montreal. I. Croissance, fecondite, biomasse et production annuelle. Canadian Jour. Zool. 49:1431-1441. Stanczykowska, A. M., M. Plinski and E. Magnin. 1972. Etude de trois populations de Viviparus malleatus (Reeve) (Gastropoda, Prosobranchia) de la region de Montreal. II. Etude qualitative et quantitative de la nourriture. Cana- dian Jour Zool. 50:1617-1624. Stearns, R. E. C. 1901. Japanese Vivipara in California. Thv Nautilus 15:91. Teskey, M. C. 1954. The mollusks of Brown County. Wiscon- sin. The Nautilus 68:24-28. U.S. Department of Agriculture. 1946. Index-Catalogue of Medical and Veterinary Zoology. Part 7. Hosts. White. J. F. and S. J. Gould. 1965. Interpretation of the coef- ficient in the allometric equation. American Naturalist 99:5-18. 96 THE NAUTILUS July 16, 1982 Vol. 96(3) FOR SALE A matched pair of blond wood shell cabinets, each 42" wide, 25" deep, and 40" high, with a 3'/z" deep display case on top, covered by 'A" plate glass. Each cabinet contains 24 drawers .3" deep. Price for pair $9.SO.OO F.O.B. Mobile. One shell dredge, 12" wide, 7" high (with sloped lip at mouth), 18" long; complete with break-aloose steel rod bridle and 1" galvanized steel lead chain, also equipped with trailing float. Steel bar frame hot-dipped galvanized after fabrication, covered with 'A" galvanized hardware cloth. Total weight 20 lbs. Price $125.00 F.O.B. Mobile. Contact H. I. Johnstone, 2209 River Forest Rd.. Mobile, Ala. 3(5fi05. I'hone (205) 471-1219. ERRATA Nautilus, vol. 94 (1) J. K. Buttner and R. C. Heidinger, "Seasonal Variations - Corbi- cula - Illinois Fish Pond." Page 9, Table 2, bottom line, for 10"" read 10-\ Page 10, left column near bottom, for P>0.01, read P<0.01. The authors express their regrets. COMMENTS ON THE EASTERN PACIFIC TRIVIA RITTERI (GASTROPODA: TRIVIIDAE) Hans Bertsch Instituto de Investigaciones Oceanologicas Universidad Autonoma de Baja California Ensenada, B.C., Mexico Barbara Myers and Department of Marine Invertebrates, Natural History Museum P.O. Box 1390 San Diego, California 92112 Although Cate (1979) recently surveyed all known species of Triviidae the information he presented on Trivia ritteri Raymond, 1903, was only a condensation of the original description. Not only were no new data presented, but one of the original collecting sites of T. ritteri (Cortez Bank, west of San Diego) was omitted. The find- ing of two specimens of T. ritteri from deep water off San Diego, California, occasioned a search for additional information. References, Synonymy and Published Records: Triiw ritteri Raymond, 1903: 85-86; Catalina Island (60 fms); Monterey; Cortez Bank (54 fms); Catalina Island near Avalon (40 fms); off San Pedro (50 fms); Lowe, 1904: 18-20; C:i(alina Island; Dall, 1921: 140; Oldroyd, 1927: 238-239; Grant & Gale, 1931; 44, 754; Upper Pliocene, Los AiiKfl's; Keen, 1937: 47; Smith & Emer.son, 1955: 99, 101; Ficny, 1956: 151, 154; Anacapa Passage, ,34°00' 38-20" N; ll;»'30' 35-1.3" W (26-27 fms); Anacapa Pas- sage, 34°00' IC" N; W.i'^-zr :j5-05" W (29-31.5 fms); 2 1/2 miles N. of W. end of Anacapa Island, 34°03' 05-45" N; 119°26' 02" to 25' 28" W (46-58 fms); Abbott, 1974: 149. fig. 1637 Puiiula ritteri (Raymond). Burch, 1945: 27-29; 42; off Redondo Beach (25 fms), White's Landing, Catalina Island (40 fms); Pleistocene of Timm's Point, San Pedro. Decoriatrivin ritteri (Raymond). Cate, 1979: 11, 98; figs. 94 and 94a. Geographic and. Bathyynetric Distributions: Trivia ritteri has been recorded from the upper Pliocene and the Pleistocene of Los Angeles, and in the Recent from Monterey to San Pedro, Catalina Island, Anacapa Passage and Island, and Cortez Bank. Our knowledge of the distribution of Trivia ritteri has been based on just a few specimens. In the collections of California Academy of Sciences (CAS), Los Angeles County Natural History Museum (LACM), and San Diego Nat- ural History Museum (SDNHM) are numerous Vol. 96(3) July 16, 1982 THE NAUTILUS 97 lots of Trivia ritteri. An analysis of the accom- panying data of those specimens gives a better understanding of the distribution of T. ritteri (Table 1). Specimens in the LACM collection ex- tend the range of T. ritteri southward more than 400 miles (over 650 km) to midway down the Baja California peninsula (27° N). Trivia ritteri is endemic to the California Province (sensu Valentine, 1966), with a few northern records to Monterey (36° N). Trivia ritteri is usually found deeper than 50 m (Figure 1). More than two-thirds of the 113 specimens analyzed for bathymetric occurrence were found between 51-90 meters deep. Trivia ritteri has a predominantly insular distribution: the majority of specimens have been collected from the offshore islands of southern California and northern Baja California. Mainland collect- ing records are either from several miles off- shore or from deep trenches (such as La Jolla) which rapidly drop off close to the shoreline. TABLE 1. Recent records o/ Trivia ritteri with their depth ranges. Location Specimens Depth Range 1 1 1 1 1 1 M 1 1 1 1 1 1 1 1 1 1 1 ; i-?o 3|-^o A 5 1-60 r< /N 71- 80 / V \ N, ' y^ 9 I-IOO \ ( M I-I20 helk is described from the eastern United States. This new species is compared with B. carica (Gmelin, 1 791) and B. contrarium (Conrad, 18U0) and its subgeneric affini- ties are discussed. A large, sinistral Busycon is occasionally col- lected offshore along the Atlantic coast of the United States between southern New Jersey and northern Virginia. This whelk frequently has a pure white shell and has been referred to as either a sinistral form of B. carica (Gmelin, 1791) or as B. contrarium (Conrad, 1840) (Abbott, 1974, p. 222). Here I show that this whelk is distinct from sympatric B. carica and B. contrarium, and describe this new species. 100 THE NAUTILUS July 16, 1982 Vol. 96(3) FIGS. 1-.5. Busycon laeostomum tl sp. 1, 258 mm (Pope coUeclion); 2, 213 mm (USNM no. 679720), 3, 222 mm (with soft tissues and operculum; Harasewych collection), 4, 221, mm (USNM no. 806850). and 5, 221 mm (siphonal canal broken; USNM wo. 806850). Vol. 96(3) July 16, 1982 THE NAUTILUS 101 Busycon laeostomum, new species Figs. 1-5, 7, 10 Shell Description - Shell sinistral, solid, sub- pyriform, inflated; shoulder rounded, tubercu- late; spire low, subconical; spiral cords on spire weak or absent; aperture broad, elliptical, longer than siphonal canal; anal ridge on parietal wall narrow, weak; siphonal canal open, somewhat flaring; columella broad, strongly bowed; spiral cords on body whorl restricted to dorsum of siphonal canal, weak; shell exterior usually chalky, white or pale pinkish-orange, oc- casionally with indistinct slightly darker suffu- sions; periostracum minutely ciliated; aperture white or light pinkish-orange; apertural lirae ab- sent or rarely weak. Description of Animal - Head and foot black; operculum corneus, elliptical, lingulate, nucleus apical; radula with 3 teeth per row, 5-6 cusps on central teeth. Material Examined - Holotype - Length = 245 mm (U.S. National Museum no. 806849); trapped in 10-15 m of water off Stone Harbor, New Jersey, January 1979. Paratypes - Length = 184 mm (USNM no. 678850); dredged 8-10 km off Avalon, New Jersey, July 1967. Length = 213, 176 and 138 mm (USNM no. 679720); dredged off Ocean City, Maryland, 1969. Length = 224 and 221 mm (USNM no. 806850) and 222 and 220 mm (M. G. Harasewych collec- tion); trapped off Stone Harbor, New Jersey in 10-15 m of water, January 1979. Length = 220 mm (USNM no. 806851); off Avalon, New Jersey, 1968. Length = 258 mm (Theresa R. Pope collection) and 83 mm (author's collection); collected as beach shells at Tom's Cove, Assateague Island, Virginia, April 1977. Length = 163 mm (Geerat J. Vermeij collection); off Ocean City, Maryland, 1977. Type Locality - Stone Harbor, New Jersey; in 10-15 m of water. Range - Southern New Jersey to northern Virginia; offshore. Etymology - laios (Gk.) = left, stoma (Gk.) = mouth. Remarks - B. laeostomum is sinistral and has a ciliated periostracum hke B. contrarium. but otherwise more closelv resembles B. carica in general shell shape (Figs. 6-11, Table 1). B. con- trarium is more slender and has a more angular shoulder than B. laeostomum, although tuber- cles along the shoulder make the latter differ- ence somewhat difficult to observe in photo- graphs (e.g. - Figs. 4 and 10). B. contrarium and B. laeostomum also differ with respect to the development of these tubercles, the nature of the external spiral cords and apertural lirae, the shape of the columella, and the color of the shell. ■WliileS. laeostomum has only rounded tubercles along the shoulder, in B. contrarium these tubercles are frequently elongated into promi- nent spines. B. contrarium also has well- developed spiral cords and apertural lirae which become obsolete in later whorls. In B. laeosto- mum these cords and lirae are very weakly developed or more commonly completely ab- sent, even in the early whorls (e.g. -compare Figs. 9 and 10). The columella of 5. contrarium. is slender and relatively straight compared to broad, strongly bowed columella of B. laeo- stomum. Finally, the dark brown axial streaks that usually occur on the early whorls of B. con- trarium (Fig. 9) do not occur in B. laeostomum. In B. laeostomum 62% of the types (8 of 13) had a pure white shell; the remaining shells were a TABLE 1. Comparison of northern Busycon contrarium and B. carica with B. laeostomum, with respect to spire angle, the ratio of siphonal canal length to aperture length, aperture shape ( = aperture uidth divided by aperture length) and canal shape (= canal uridth halfway along canal divided by canal length). The values listed are means with ranges in- dicated in parentheses. Shells with badly broken lips or siphonal canals were not measured. Differences were tested for signif>.cance with a one-tailed Mann-Whitney U test and the level of significance is indicated by asterisks (i.e.-' in- dicates p <0.05. ** indicates p <0.01. and **' indicates p 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 Janowsky's MAL DE MER ENTERPRISES 946 Ralph Avenue Brooklyn, New York 11236 USA (212) i85-S550 122 THE NAUTILUS July 16, 1982 Vol. 96(3) LARVAL DEVELOPMENT IN CREPIDULA MACULOSA (PROSOBRANCHIA: CREPIDULIDAE) FROM FLORIDA' K. E. Hoagland and Lehigh University Bethlehem, PA and 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 (posthumous) '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." LITERATURE CITED Caswell, H. 1981. The evolution of "mixed" life histories in marine invertebrates and elsewhere. Amcr. Naturalist 117(4):529-536. 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. ENTFORMATION FOR SUBSCRIBERS The annual subscription rate for The Nantilus is $12.00 for individuals (foreign $14.00) and $18.00 for institutions (domestic or foreign). Subscrip- tions may begin in January. Send check or money order made out to "American Malacologists" to the Business Manager, P.O. Box 2255, Melbourne, Florida 32901, U.S.A. Back issues from volume 72 to date are ob- tainable from the Business Manager. Volumes 1 through 71 (if available) may be obtained in reprint or original form from Kraus Reprint Co., Route 100. Millwood, New York 10546. Advertising rates may be obtained from the Business Manager or Editor. 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WANTED -OLD SHELL BOOKS Will pay good prices for libraries, second- 305-725-2260) or write: R Tucker Abbott, hand books and reprints on mollusks, shells American Malacologists. Inc.. P. 0. Box 2255 and conchology. Back numbers of Ue Nauti- Melbourne, FL 32901. Free appraisals. //^s. vols. 40-71 wanted, $1.50 each. Phone (1- americanmalacologists, mc. PUBLISHERS OF DISTINCTIVE BOOKS ON MOLLUSKS THE NAUTILUS (Quarterly) MONOGRAPHS OF MARINE MOLLUSCA STANDARD CATALOG OF SHELLS INDEXES TO THE NAUTILUS ((ieographiral. vols 1-90; Sdentific Names, vols 61-90) REGISTER OF AMERICAN MALACOLOGISTS :T0BER 29, 1»82 THE > :?■•.>•■> •-'•'- m^^' nft NAUTILUS ISSN 0028-1344 Vol. 96 No. 4 A quarterly devoted to malacology and the interests of conchologists Founded 1889 by Henrj' A. Pilsbry. Continued by H. Burrington Baker. Editor-in-Chief: R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William J. Clench Curator Emeritus Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, New York 10024 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, Ohio 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, California 90007 Dr. Arthur S. Merrill 103 West 8th Avenue Cudjoe Gardens Summerland Key, Florida 33043 Dr. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Joseph Rosewater Division of MoUusks U.S. National Museum Washington, D.C. 20560 Dr. G. Alan Solem Department of Invertebrates Field Museum of Natural History Chicago, Illinois 60605 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, Ohio 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Charles B. Wurtz 3220 Penn Street Philadelphia, Pennsylvania 19129 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, Florida 32901 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 2255 Melbourne, Florida 32901 Tlie Nautilus (USPS 374-980) ISSN 0028-1344 OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbourne, FL 32901) Mail: Box 2255, Melbourne, FL 32901 Second O.ass 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 4 THE NAUTILUS October 29, 1982 ISSN 0028-1344 CONTENTS 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 .125 .127 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 .129 ,130 C. John Finlay and Danker L. N. Vink New Records of Cymatiidae (Gastropoda) in the Western Atlantic . ,132 Robert F. McMahon The Occurrence and Spread of the Introduced Asiatic Freshwater Clam, Corbicula fluminea (Miiller), in North America: 1924-1982 .134 John W. Ropes Hermaphroditism, Sexuality and Sex Ratio in the Surf Clam, Spisula solidissima, and the Soft-shell Clam, Mya arenaria . . . ,141 Billy G. Isom and Robert G. Hudson In Vitro Culture of Parasitic Freshwater Mussell Glochidia. 147 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. ,152 ,156 ,151 SUBSCRIPTION NOTICE Now due for 1983, Nautilus volume 97, individuals $13.00 (foreign $15.00); institutions (and agencies) $18.00. Please send your check now to AMERICAN MALACOLOGISTS, INC., P.O. Box 2255, Melbourne. FL 32901. It will save us time and costs if you respond before December 1. Early subscribers may obtain a copy of Say's 1819 Conchology reprint (originally $5.00) at 50% off. Add $2.50 to your payment, postage free. 124 THE NAUTILUS October 29, 1982 Vol. 96(4) Freshwater Snails of Africa and their Medical Importance by David S. Brown hor the first time: A comprehensive account of freshwater snails in Africa and neighbouring islands. An exhaustive guide to species of medical and veterinary importance. An invaluable reference work for malacologists, epidemiologists, parasitologists, freshwater biologists and biogeographers. Published by TAYLOR & FRANCIS LTD, LONDON. Distributed in the Americas by AMERICAN MALACOLOGISTS, INC. FLORIDA. 450 pp. 230 X 152inm (9" 0 85066 145 5 Cloth 153 Figs. ■6") Regular price $55.00 (plus postage) NOW $27.50 ITie knowledge resulting from intensive study in recent years is brought together for the fu'st time in this book. Dr. Brown presents a systematic synopsis of nearly 400 species, most of them illustrated, together with chapters on host/parasite relations, snaxl control, ecology, distribution and biogeography, accompanied by many maps, photographs of shells and drawings. (plus postage) $2. op NEW: SUPPLEMENT 2 Wagner and ABBt)Trs STANDARD CATALOG OF SHELLS Complete species and generic catalogs of six major families, a revision of 1200 World Size Records, and an Index to genera and subgenera. With seven New guide tabs. Fits into your pres- ent binder. 116 pages, iljus. $13.00, plus $1.00 postage (foreign postage $1.50). Families covered: Coralliophilidae (Latiaxis. Magilua, etc.); Triviidae; Ovulidae; Nassariidae (by W. 0. ("ernohorsky, about 2,600 names); Glycymerididae (650 names); Spondylidae (250 names). Order now from AM K RIG AN MALACOLOGISTS, INC., P.O. Box 2255, Melbourne, FL 32901, USA. We accept VISA or MASTER CARD payments. Give your name, card number, and the month and year of expiration. Vol. 96(4) October 29, 1982 THE NAUTILUS 125 A SINISTRAL SPECIMEN OF SCAPHELLA FROM THE WACCAMAW FORMATION (EARLY PLEISTOCENE), SOUTH CAROLINA Jules R. DuBar' and Susan S. DuBar Geoscience Department Morehead State University Morehead, Kentucky ABSTRACT 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. 126 THE NAUTILUS October 29, 1982 Vol. 96(4) ^ SCAPHELLA LOCALI '6 ' r-iT 10 Mile ii-33 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 specimens. 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 Petit. Bivalvia 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) Gastropoda 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) Anthozoa Septastraea crassa (Holmes) P>hinoidea Arbiiciii wacca/naw Cooke I'lypeaster crassus Kier Encope michelini imperforata Kier Rhyricholampus sp. cf. R. ayresi Kier LITERATURE CITED 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 THE NAUTILUS 127 NOTES ON THE FRESHWATER MUSSELS (UNI()NH)AE) OF THE UPPER MONONGAHELA RIVER BASIN, WEST VIRGINIA Michael A. Zeto West Virginia Department of Natural Resources Division of Water Resources 350 North Vance Drive Beckley, WV 25801 ABSTRACT 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 counties. 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 Fairmont. 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. 128 THE NAUTILUS 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). Methods 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. Results 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, 1980. Species Site No. 1 2 3 1 Anodonta q. grand is (Say. 18291 X X Strophitus u. undulatus (Say,18J7) X X X Sinpsonaias ambiqua (Sav. 18251 X Lasmiqona costata (Raf..l820) X X X X Tritoqonia verrucosa (Raf..l82n) X X X /Imblera t^ plicaU (Say. 1317) X Fuscofiaia flava (naf.. 18201 X X PleurobcTO clava (lam.. 1819) X Pleurobema sintoxia (Raf..I320) X Elliptio dilauta (l!af.,lS20) X X X Ptychobranchus fascioUrls (Raf..l820) X X Obovarla subrotunda (Raf..l020) X X X X X Lampsilis radiata luteola (Lam.. 1819) X Lampsllls ventricosa (Carne5.1B23) X X X EpIoblasiM triQuetra (Raf.. 18201 X X Corblcula fluminca (Muller.1774) X 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 specimens. ACKNOWLEDGMENTS 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. LITERATURE CITED 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 THE NAUTILUS 129 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 .VaH(i7«s92(4):153-1.5.5. (.)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(3.s. First International Corhicula Si/mposium. pp. (i9-80. Texas Christian I'niversity Research Foundation. f\)rt Worth. Texas. Sinclair, R. M. 1974. Effects of an introduced clam (Cor- hicula) on water quality in the Tennessee River Valley. In. Proceedings of the Second International Waste Conference. pp. 43-50. Tennessee Department of Public Health. Tennessee Stream Pollution Control Board, Na.shville, Tennessee. Sinclair, R. M. and B. G. Isom 1961. A preliminary report on the introduced Asiatic clam Corhicula in Tennessee. Ten- nessee Stream Pollution Control Board, Tennessee De- partment of Public Health. Nashville, Tennessee, 31 pp. 1963. Further studies on the introduced Asiatic clam (Corhicula) in Tennessee. Tennessee Stream Pollu- tion Control Board, Tennessee Department of Public Health, Nashville, Tennessee, 76 pp. Smith, A. L., R. A. Muia, J. P. Farkas and D. (). Bas.sett 1979. Clams-a growing threat to inplant water sy.stems. Plant Engineering 1979: 16.5-167. Taylor. D. W. 1981. Freshwater mollusks of California: A distributional checklist. Calif. Fisti Game 67:140-163. Taylor, R. W., and R. C. Hughart 1981. The freshwater naiads of Elk River, West Virginia with a comparison of earlier collections. The Nautilus 95:21-2."). Thomas, N. A., and K. M. MacKenthum 1964. Asiatic clam infestation at Charleston. West Virginia. The Nautilus 78:28-29. Thonierson, J. E., and D. G. Myer 19711. Corlncula manilen- sis: Range extension upstream in the Mississippi River. Stcrk-iana 37:29. Thompson, C. M., and R. E. Sparks 1977. Improbability of dispersal of adult Asiatic clams, Corhicula inanilensis. via the intestinal tract of migratory water fowl. Amer. Midi. Natur. 98:219-223. 1978. Comparative nutritional value of a native fingernail clam and the introduced Asiatic clam. Jour. Wildl. Manage. 42:391-396. Trama, F. B. 1982. Occurrence of the Asiatic clam Corhicula fluminea in the Raritan River. The Nautilus 96:6-8. HERMAPHRODITISM, SEXUALITY AND SEX RATIO IN THE SURF CLAM, SPISULA SOLIDISSIMA, AND THE SOFT-SHELL CLAM, MYA ARENARIA John W. Ropes National Marine F'isheries Service Northeast Fisheries Center Woods Hole Laboratory Woods Hole, Massachusetts 02543 ABSTRACT 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 142 THE NAUTILUS 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- mented. 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. Methods The preparation of gonadal tissues for micro- scopic examination was by standard histological methods outlined in Ropes and Stickney (1965). Results 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. DISCUSSION 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 changes. 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) A October 29, 1982 THE NAUTILUS 143 . 100 ^m c D cf am ]0 ^m ' * to tj.f" FIG 1 SecUon. of the gonad, ofhennaph nMl0.05) 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. Location No. clams examined No. hermaphro- dites Reference 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. >1,000 3 Coe S Turner, 1938 1,063 0 Shaw, 1965 > 700 0 Pfitzenmeyer, 1965 1,400 0 Ropes 5 Stickney, 1965 36 1 Shaw, 1970 1,311 10 Otto, 1973 1,785 1 Porter, 1974 2,480 0 Brousseau, 1978 3l. 96(4) October 29, 1982 THE NAUTILUS 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 68. 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 Number of Area Year samples males females Total .X" Long Island 1965Sp 4 17 22 39 0.6410 1965Fa 5 27 18 45 1.8000 1968 7 38 35 75 0.1253 1969 1 4 6 10 0.4000 Total 17 86 81 167 0. 1498 Pt. Pleasant, NJ 1962 8 74 98 172 5.3488 1963 20 244 246 490 0.0082 1964 20 240 244 484 0.0330 1965 14 170 198- 368 2. 1304 1965Sp 4 18 18 36 0.0000 1965Fa 5 28 21 152^/ 1.0000 1968 14 58 73 1.7121 1969 3 S -) -1 10.3571 Total 88 S3- 920 1-59 3.9187 Cape May-Wildwood, NJ 1965Sp 6 18 30 48 3.0000 1968 8 45 36 ^ 81 1.0000 1969 4 26 14 40 3.6000 Total 18 89 80 169 0.4792 Delmarva Peninsula 1965Sp 5 25 23 4^/ 0.0202 5 Virginia 1965Fa 13 46 48 94 0.4256 1968 6 27 33 60 0.6000 1969 1 6 3 9 1.0000 Total 25 104 107 212 0.0471 Grand Total 148 1116 1183 2507 2.2510 1/ Includes a hermaphrodite in the total. 146 THE NAUTILUS 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. ACKNOWLEDGMENTS 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 comments. LITERATURE CITED 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. 76:1.5,V166. 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.' U < 3 O 300 1 T 200 100 0. cyanea 0. ornatus crescent octopus 1200 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. 154 THE NAUTILUS 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- face. 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 THE NAUTILUS 155 DISCUSSION 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. ACKNOWLEDGMENTS 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 author. LITERATURE CITED 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. 1966-67:77-83. Boyle. P. R. 1980. Home occupancy by male Octopus cul- garis in a large seawater tank. Animal Behavior '28:1123-1126. 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. 2:337-343. 156 THE NAUTILUS 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. REDISCOVERY OF THE MARGINELLID GASTROPOD PERSICULA TESSELLATA (LAMARCK, 1822) ON THE PACIFIC COAST OF PANAMA Barry Roth California Academy of Sciences San Francisco, California 94118 William K. Emerson and American Museum of Natural History New York, New York 10U24 ABSTRACT 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 Philadelphia 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. 158 THE NAUTILUS 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. DISCUSSION 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 THE NAUTILUS 159 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 holotype. 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 160 THE NAUTILUS 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 History). 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 THE NAUTILUS 161 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. ACKNOWLEDGMENTS 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. LITERATURE CITED 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. 107(1):1-158. 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., 1848]. 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. Notices 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. 162 THE NAUTILUS October 29, 1982 Vol. 96(4) COMPENDIUM OF SEASHELLS An Authoritative Guide to over 4,000 Marine Shells of the World in Color by R. Tucker Abbott and S. 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