americanmalacologists, inc. PUBLISHERS OF DISTINCTIVE BOOKS ON MOLLUSKS THE NAUTILUS (Quarterly) MONOGRAPHS OF MARINE MOLLUSCA STANDARD CATALOG OF SHELLS INDEXES TO THE NAUTILUS {Geographical, vols 1-90; Scientific Names, vols 61-90) REGISTER OF AMERICAN MALACOLOGISTS JANUARY 31, 1986 THE NAUTILUS FEB 5 1986 I Woods Hole, Mass. ISSN 0028-1344 Vol. 100 No. 1 A quarterly devoted to malacology and the interests of conchologists Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker. Editor-in-Chief: R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Mr. Samuel L. B. Fuller 1053 Mapleton Avenue Suffield, CT 06078 Dr. M. G. Harasewych 363 Crescendo Way Silver Spring, MD 20901 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, OH 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill c/o Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, FL 33149 Dr. G. Alan Solem Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, FL 33149 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, FL 32902-2255 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 2255 Melbourne, FL 32902-2255 Second Class Postage paid at Melbourne, Florida and other post offices The Nautilus (USPS 374-980) ISSN 0028-1344 A quarterly magazine devoted to malacology. Copyright ^1986 by American Malacologists, Inc. OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbourne, FL 32901) Mail: Box 2255, Melbourne, FL 32902-2255 POSTMASTER: Send address changes to above. Subscription Price: $15.00 (see inside back cover) $17.00 (foreign); institutions $20.00 THE NAUTILUS Volume 100, number 1 — January 31, 1986 ISSN 0028-1344 CONTENTS Dedication of the 100th Volume 1 Birth and Growth of The Nautilus 2 Tom Pullev and the Train 8 Joseph Rosewater (1928-1985) A Tribute and Bibliography by Harald A. Rehder . Andrew C. Miller, Barry S. Payne and Terry Siemsen Description of the Habitat of the Endangered Mussel Plethobasus cooperianus 14 Andrew C. Miller, Barry S. Payne and David W. Aldridge Characterization of a Bivalve Community in the Tangipahoa River, Mississippi 18 Richard A. Petit Notes on Species of Brocchinia (Gastropoda: Cancellariidae) 23 William K. Emerson On the Type Species oi Metula H. & A. Adams, 1853: Buccinum. clathratum A. Adams and Reeve, 1850 (Gastropoda: Buccinidae) 27 Eugene Coan Some Additional Taxonomic Units that First Appear in Publications by J. G. Cooper 30 Dee S. Dundee Notes on the Habits and Anatomy of the Introduced Land Snails, Rumina and Lamellaxis (Subulinidae) 32 Award News 37 Notes; Deaths 38 STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULA- TION (Required by) Act of October 23, 1962: Section 4396, Title 39. United States Code, and postal regulation 132-622. 1. Title of publication THE NAUTILUS. 2. Date of filing: September 14, 1985 3. Frequency of Issue: Quarterly (4 per year). 4. Location of known office of publication: 2208 South Colonial Dr., Melbourne, FL 32901. 5. Location of Headquarters of General Business Offices of the Publishers: 2208 South Colonial Dr., Melbourne, FL 32901 , 6. Names and addresses of publisher, editor, and manag- ing editor: Publisher, American Molocologists, Inc., P.O. Box 2255, Melbourne, FL 32901, Editor, R. Tucker Abbott, P.O. Box 2255, Melbourne, FL 32902, Business Manager, Mrs. Cecelia W Abbott, P.O, Box 2255, Melbourne, FL 32902-2255, 7. Owner: American Molocologists, Inc., P.O. Box 2255, Melbourne, FL 32902-2255. 8. Known bondholders, mortgages, and other security holders owning or holding 1 percent or more of total amount of bonds, morfoges or other securities: none. 9, Extend and Nature of Circulation: Average Single 12 Mos. Issue 850 850 A. Total No. copies Printed (Net Press) B. Paid Circulation 1. Soles through dealers and carriers, street vendors and counter sales 2. Moil subscriptions C. Total Paid Circulation D. Free Distribution (including samples) by moil corner or other means E. Total Distribution (Sum of C&D) F. Office use, left-over, unaccounted and back start subscription copies G. Total (Sum of ESF)-should equal net press run shown in A. I certify that the statements mode by me above are correct and complete. (signed) R. Tucker Abbott, Editor none none 735 730 735 730 18 18 753 748 91 102 850 850 THE BEST OF THE An Exciting and Nostalgic Glimpse Into Conchologys ftist Heii' IS a IreiTSLirv of iinsl.Tlf^n and ofton humorous writings takfii from tho first 40 voars of America's oldest mollusk |ourn,il, covering the Lite pioneering f^'riod of 1886 to 1927 The noted m.iKicoiogist and editor, R Tucker Abholl has carefully gathered a wide-ranging series of over 100 articles that depict tiir lore and ex- citement of sli.'il hunlmt; in the United States American M«lacologlst5 P.O. Bo. 2255 288 pp , rinthbound $13,95. Melbourne, FL 32902-2255 Collectible Shells of Southeastern U.S., Bahamas & Caribbean by R, Tucker Abbott. Ph d A 'Take it to ttie Beach' Field Guide WATERPROOF - TEARPROOF 105 beautiful color photos of living animals and their shells 64 pages ol color 300 species il- lustrated. How to clean shells. Where to find them. Includes fossils, pond and tree snails, as well as sealife Collectible Shells stresses conservation, but also has helpful hints about collecting and cleaning shells. The book introduces the tourist and beginner to famous Florida fossils and the unique world ol tree and pond mollusks Printed on a washable, tearproof plastic paper. Drop it in the ocean, use it in the ram, or let your wet shells drip all over it. Keep it on your boat or take It to the shore A popular new seller retailing (or $8.95 Postage a id state lax are included as a big savings American Malacologists, Inc. Publishers of Distinctive Books on Mollusks P.O. Box 2255, Melbourne, FL 32902-2255 W« accept VISA Of MASTERCARD orders Dymai' Piea";e give dale o' e« pifation »n(j youf ctfd number Fo'eigr* customers may send mte'na lionai postal money order ot checK on New Yofh bank o' U S cash by 'egisiered mail Vol. 100(1) January .31, 1986 THE NAUTILUS The Nautilus ITS 100th ANNIVERSARY and Volume 100 Dedicated to all the mollusk enthusiasts, both private concholog-ists and pro- fessional malacologists, who founded and carried on for the last one hundred years America's oldest journal devoted exclusively to the study of mollusks. To its 500 contributors and 15 consulting editors are added many thanks and appreciation. DEDICATED TO THE FORMER EDITORS^ William D. Averell (1853-1928) Founder of the Conchologists' Exchange, 1886 Henry A. Pilsbry (1862-1957) Founder, editor of The Nautilus, 1888-1957 Charles W. Johnson (1863-1932) Business manager, 1890-1932 Horace Burrington Baker (1889-1971) Business manager, co-editor 1932-1957 Editor, 1958-1970 Charles B. Wurtz (1916-1982) Co-editor, 1958-1974; consulting editor, 1974-1982 Bernadine B. Baker (1906-living) Business manager, 1958-1979 Morris K. Jacobson (1906-1980) Consulting editor, 1972-1980 William J. Clench (1897-1984) Consulting editor, 1972-1984 Joseph Rosewater (1928-1985) Consulting editor, 1972-1985 'The present staff is R. Tucker Abbott, co-editor, 1958-1969, editor since 1970; Cecelia W. Abbott, business manager, since 1979; M. G. (Jerry) Harasewych, associate editor, beginning with this volume. Mrs. H. B. Baker, former business manager, lives in Haverford, Pennsylvania. 2 THE NAUTILUS January 31, 198f; BIRTH AND GROWTH OF THE NAUTILUS Vol. 100(1) It was the year 1886. The country was recov- ering from the depression of 1884. President Grover Cleveland, in his first term, was about to dedicate the newly erected Statute of Liberty in New York harbor. There were still 12 states to be added to the Union, and it took a two-cent, brown postage stamp with Washington's like- ness to send a letter across the country. In Philadelphia the only means of transportation were horses, bicycles, the steam locomotive and boats on the Delaware River. The Academy of Natural Sciences of Philadelphia had not built its large new annex and still used gas lights in its halls and study rooms. In England, they were celebrating Queen Victoria's Diamond Jubilee, and the Spanish American War was still 12 years away. In 1886, in some respects, it was the best of times for American conchology. William H. Dall of the U.S. Geological Survey in Washington, D.C. had already produced over 50 major works on mollusks, and Harvard University had just published the first part of his famous "Blake Report" on deepsea mollusks. George W. Tryon, Jr., the Philadelphia author of "Structural and Systematic Conchology," had just completed the tenth volume of his monumental "Manual of Conchology." Verrill and Bush in New England were at the height of their reports on the dredg- ings of the Steamers FUh Hnivk and the AlbatToss. William G. Binney, living in New Jersey, just across the river from Philadelphia, had just revised his classic "Manual of North American Land Shells." The Conchological Sec- tion under the wing of Philadelphia's Academy was nourishing with such active members as S. M / y y ' ^ 7 / // y o-vSr ^-^ ' -'^/^ ■J 11 ^Hy//X Tho end (if a letter written in Dec. 18i»0 to the editors of the newly launc-hiMl XnuiWux hy the Calil'ornia-liased shell collector and newspaper editor, Kdward Warren Ro(ier, admon- ishing them to have it "come out on time." Vol. 100(1) January 31, 1986 THE NAUTILUS 3 $1.00 per Year. ($1.12 to Foreign Countries.) lOcts. a copy. THE NAUTILUS A MONTHLY DEVOTED TO THE INTERESTS OF CONCHOLOGISTS. EDITOR: H. A. PiLSBRT, Conservator Conchological Section, Academy of Natural Sciences, Philadelphia. ASSOCIATE EDITOR : C. W. Johnson, Acting Curator Wagner Institute of Science. Vol. VI. FEBRUARY, 1893. No. 10. CONTENTS: PAGE. Additional Shells from the Coast of Southern Brazil. Wm. H. Dall 109 Description OF A NEW FORM OF CYPR.iiA. John Ford 112 Vol. 84 JULY, 1970 No. 1 THF NAUTILUS THE PILSBRY QUARTERLY DEVOTED TO THE INTERESTS OF CONCHOLOGISTS EDITORS AND PUBLISHERS R. Tucker Abbott, du Pont Chair of Malacology Delaware Museum of Natural History. Greenville. Del. 19807 Horace Burrington Baker, 11 Chelten Road, Havertown, I'a. (Emeritus Professor of Zoology, University of Pennsylvania) Charles B. Wurtz, Biology Department La Salle College, Philadelphia, Pa. 19141 CONTENTS The occurence of Cymatiidae and Cypraeidae in North Carolina. By Hugh J. Porter 1 Eggs and attachment sites for egg capsules of Valvata lewisi. By B. Z. Lang and N. O. Dronen, Jr 9 THE NAUTILUS January 81, 1986 Vol. 100(1) Vol. I. No. 1. This is the first issue of "The Conchologist's Exchange " As encouragpnient is received it will assume the form of a printed sheet with columns for " Exch.-inges in Mollusca," "New Localities," "Answers to Correspondents," ike. This, our first number, has been sent to 500 Conchologists. Subscription price, 25 cents per annum, post paid. Exchanges of 20 words, 10 cents; for each additional 10 words the charge will be 5 cents. The Conchologist's Exchange will be issued semi-monthly, and will endeavour to become a cheap and useful medium for the exchange of those most beauti- itful productions of nature — "The Mollusks." EXCHANGES FOR MOLLUSCA ONLY. CVPR.^A erosa, L. lynx, L. CERITHIUM. maculosum, Kien. eburncum Brug. CVCLOS rOMA sulcatum. Lam. clegans Mull, LVMN.-EA lebraTryon bTROMBINA ^icanahfcra Sby. Fi-^urclla volcano. Rve. Col- umbella fulgunns Lam. Prof. D. S. SHELDON. Davenport, fa SUCCINEApuiris L. HLLIX arbuslorum L. " ncmomlii L " ericelorum. Mull. Toiundato, MuU. " Upicida L ccllaria Mull Pupa muscorum, L. Cionella subcylindriea E LtHNERT, Washington, D C. CONIOBASIS simplex, Suy. cariiiifcra. Lam. bella, Con. p«rangulata. Con, sordiJa L«a symiTielrica, Hald ;bcnum. Lea MeUntbo subsolida Anth. Unioriibigjnosus Lea pustulosus Lea. gracilis, Barnes w. a; marsh. Aledo. Illinois HELIX albolabris. Say. alter- nata. Say. clauaa.Say. elevata, bay. fallax, Say. hirsuta, Say. inflecta, Say. solilaria, Say. monodon, Rackett. Sayii. Binncy. Pupa armifera. Say, corlicana. Say. Fallax, Say Unio cicgans Lea, lachrymosus. Lea, parvus, Barnes EDWARD A. ENOS, Connersvillc, Indiana. NASSAfossataGld. Purpura saxicola Val. Amycla gausapata Gas. Adula falcata Gld. Acmaea spectrum Esch scabra. Nuti, pelta, Esch. Hipponyx cranioidcs. Carp. G. W. PUIFRBAL'GH, Greenfield, Indiana. American and Foreign Unionidac for exchange. Send for list. No responsibility will be ai- umed for the siaQ(Jing of the bovc parties. Address, WILLIAM 0. AVERELL, Proprietor, CHESTNUT HILL, PHILADELPHIA. f ^e Conr^ologists' ^H^ange. A Monthly Publloatlon desltrned for Ood .holOitlerta and Sclentlsta arenerally. "Wm, D Averell. Editor and Publlaher. Vol. II- MARCH and APRIL, 1888. No. 9. oojsra?E2sra?s. Ill,, Wm A. H, Wnght, Tbc .Sl.ell-Hcwing MuUuscb of Kliode Island, II, K, C»ipenier, brief Noics on the LAod and Frcsh-Wner Shells of Mercer Co. Note* on the Umonidaeof Florida, Dr, S. H. Wnght and Berlii Lyogyrus. Gill, and oiher American shells, H. A Pibbry, MarganuiJA Hildrcthiaaa. Lea. U. Shimek, ... Notes on .^mencao Shells, Rev. Wm. M. Beauchamp, A Noted Scienust Dead (Geo, W. Tryon, Jr ) From Phila. Public Ledger, Bell Taps. Editorial Note*, ,,..... The CoDchologiSt in Bermuda, J, Matthew JoDe«. A Collecung Trip to On»et Bay, Maw,. F. C. Baker, . Exchangei, ...... Suodard Work* oa Conchology 109 110 "5 116 "7 118 119 119 120 CHESTNUT HILL, Philadelphia, I'a. Slnylo copies, C cents; 50 oenia a year In the United States and Canada; Foreign Countries, 62 cents; Countrtee not In Universal Postal Union, 7B cents. Vol. lUU(l) January 31, 1986 THE NAUTILUS Raymond Roberts (Cypraeidae), John Ford (Olividae), Dr. Benjamin Sharp (Scaphopoda) and Angelo Heilprin (Florida fossils). But in the late 1880's, in other respects, it was the worst of times for American conchology. It was like the changing of the guard. Aging Isaac Lea died in 1886 at the age of 95; Lucy Say, widow of America's first conchologist, died on Nov. 15, 1887, and Andrew Garrett, the con- chologist of the South Seas, died the same month. Then, like a clap of thunder, George Tryon, at the age of 49, succumbed to a heart attack on February 5, 1888, soon after a walk in a severe winter storm. It was a time when there was no journal in America devoted solely to mol- lusks. Tryons American Journal of Conchology had succumbed in the financial panic of 1873. In Europe, in 1886, there were eight respecta- ble scientific journals devoted solely to mollusks -two each in England, Germany and France and one each in Belgium and Italy (Jutting and Altena, 1958). For their outlets, American malacologists depended on the general publica- tions of the Smithsonian Institution, the Califor- nia Academy of Sciences, Harvard's Museum of Comparative Zoology, and the Academy of Natural Sciences of Philadelphia. The need for a new American serial publica- tion was evident. There were several hundred active conchologists in United States and Canada, many of them already publishing privately or in obscure natural history magazines. Among them were Henry Hemphill, Temple Prime, R. E. C. Stearns, T.H. Aldrich, W. W. Calkins, Charles T. Simpson and Victor Sterki, to mention but a few. The Nautilus had a very inauspicious begin- ning. William D. Averell, a 33-year-old shell dealer and close friend of Tryon's living in near- by Chestnut Hill, began the predecessor of The NautiliLs as the Conchologists' Exchange in July 1886. Volume 1, number 1 was merely a post- card sent out to 500 people announcing that the annual subscription price for this monthly 4-page publication would be all of 25 cents. It was to contain conchological news, exchange notices and answers to correspondents. By February 1887, it had grown to 8 pages and the price rose to 35 cents. The final number, vol. 2, no. 9, issued in April 1888, two months after Tryon's death, had 16 pages. Already it had served as a publication medium for such well- known mollusk workers as F. C. Baker, Josiah Keep of California, C. F. Ancey of Algiers, W. H. Dall, T. D. A. Cockerell and a Harry A. Pilsbry (later to call himself, Henry). But a quirk of circumstances in Philadelphia changed the history of this journal. William B. Marshall, a Jessup Student at the Academy and the assistant to Tryon, left in 1887 to be the Assistant Zoologist at the New York State Museum in Albany, and later to serve under Dall and Bartsch in Washington for 40 years. Just at this time a young newspaper reporter by the name of Pilsbry from Davenport, Iowa, and a contributor to the Conchologists' Exchange, visited the Academy on his way to seek new employment in New York. Tryon was favorably impressed with the 24-year-old amateur con- chologist who already had considerable know- ledge about mollusks, was an excellent drafts- man and had a natural bent for research and publishing. Upon Marshall's sudden departure for Albany, Tryon immediately invited Pilsbry to return from New York and be his new assis- tant. Pilsbry arrived in November 1887, and Tryon died three months later. Pilsbry was elected as conservator of the Con- chological Section in 1888, and he immediately plunged into the task of finishing the marine series of the Manual of Conchology. a publication that supplied much of his salary. There was a reawakening of interest in the Conchologists' Exchange, and in early 1889 a "Prospectus" was sent out announcing its continuation as The Na^dihis with Pilsbry as editor and Averell as business manager. Volume 3, no. 1 began on May 5, 1889. According to Pilsbry's colleagues (H. B. Baker, 1958) the one year's association with Averell "was none too happy" and, at the end of volume 4, no. 1, Averell's share was bought out by another Philadelphian, at the Wagner Free Institute of Science: Charles W. Johnson. This ideal partnership lasted for the next 42 years until Johnson's death in Boston in 1932. Johnson was a very affable and knowledgable gentleman whose forte was entomology and paleontology, as well as malacology. He was particularly successful in inspiring young naturalists, and when he moved to Boston in 1903 to assume the curatorship of the Boston THE NAUTILUS January 31, 1986 Vol. lOU(l) Society of Natural History, William J. Clench as a youngster fell under his spell. Years later, from 1926 to 1932, Clench was the Curator of Mollusks at nearby Harvard, and often assisted Johnson in the business affairs of The Nautilus. It was Johnson's understanding that Clench would succeed him as business manager, but upon the former's death in 1932, Pilsbry (then 69) thought that both editors of The Nautilus should be in the same city; so H. Burrington Balcer, one of America's leading land snail ex- perts and Professor of Zoology at the University of Pennsylvania, began his association as junior editor and business manager in October 1932 with volume 46. Meanwhile, Clench in 1941 launched the marine journal, Johnsonia, naming it after his old mentor. The Nautilus continued to flourish and grow. Its history is well-told in the Pilsbry memorial number of The Nautilus by Dr. Baker (1958, vol. 71, no. 3, pp. 112-115). Pilsbry died on October 26, 1957 at the age of 94, and the editorship of the journal continued under Baker's capable hands until his death on March 11, 1971, at the age of 82. In April 1958, two junior editors joined The Nautilus, one being the land snail expert and aquatic biologist, Charles B. Wurtz (1916-1982) who had received his Ph.D. under Dr. Baker at the University of Pennsylvania. At the age of 34 I had left the Smithsonian Institution in 1954 to become the first occupant of the newly-formed Pilsbry Chair of Malacology at the Academy. I became the other junior editor and often assisted Pilsbry in the preparation of the jour- nal, sometimes entirely assembling and editing it during Pilsbry 's winter sojourns in Lantana, Florida. Also, at this time. Dr. Baker's wife, Bernadine Barker Baker, known to her family and friends as "Bunny," took over the business and subscription management, and continued in that capacity with great success for the next twenty years. In 1969 I helped found and organize the Delaware Museum of Natural History in Green- ville, and became the curator of their Depart- fi^osfEcyds,- "We propose lo Issue a sjxieen paQ,e iliustraied acTavQ monrnly journal ol Concholosy to be trailed THE NAUTILUS. It will lake ihe place cf ihe Concholoaist's Ex- change formerly published oy Wm. D Averell, and will be the successor of that paper. Subscribers lo the Exchange will be crediied on the books ol the Nautilus wlih such amounts as may be due thern upon their subGCrlptlons. The Nautilus will be under the editorial man- agement of Mr. Henry A. Pilsbry, Conservator of the Conchologlcal Section at the Academy ot Natural Sci- ences, and the successor ot the late Mr. Tryon in the publication of The Manual of Conchology. The new Journal is designed to afford a medium of communication between siudents and others interesi- ed in conchology [ to publish original articles contrib- uted by amencan and foreign authors^ to record new discoveries) and lo teach beglnnerG and young studenis of science. America needs a Journal ol this kind, and if we can obtain sufficient encouragement In the shape ol subscriptions we will commenca the publication of The Nautilus at once. We enclose yju a blank for your subscription. If you think well of the prjject please fill it up at once and return it to Mouni Airy, PriUadalphic. Pi The Nautilus. JANUARY, 1896. TO C0NCH0L0QIST8. We regret the necessity which compels ua to start the new year with an editorial of this nature; but it is nothing new to hear that the Nautilus must struggle for existence. This struggle has con- tinued since the Jura, until now we have but a few species, three or four in the Indo-Pacific and one in the United States. Do vou in- tend that the only recent Nautill's In North America shall be- come extinct ? We are willing lo supply the care, but not the en- tire environment. It needs feeding once a nionth in order to add another septum. You are asked to assist in this important function once a year ; and when you see a slip of paper which reads, " Inclosed please find $1,00," it mean.s that it hasconie your turn to " chip in." We hope that you will no longer neglect these little reminders. They mean that your subscription is due. We cannot wait until the end of the year— U'<; mt(a( have \i in advance. Wishing you all a Happy New Year. H. A. P. A C. W. J. Vol. 100(1) January 31, 1986 THE NAUTILUS ment of Mollusks. I moved The Ndutilns. now owned by Mrs. Baker, to Delaware. In 1979, upon the recommendation of Bunny, my wife, Cecelia White Abbott, took over the duties of business manager. There was need for an improvement in the format of the journal which had remained un- changed for several decades. In May 1972 (volume 86) the page size was increased to 8 x IOV2 inches, a two-column format established on a glossy paper, and a board of 13 consulting editors formed. By now, 85 years after its incep- tion, the annual subscription rate had risen from 25 cents to $7.00 for individuals. Today it stands at $15.00. With the demise of the Department of Mollusks at the Delaware Museum in 1979 as a functioning, scientific entity, we moved The Nautilus, now under the jurisdiction of my American Malacologists, Inc., to Melbourne, Florida, where it has been published for the last six years. We have now added an associate editor, Dr. M. G. (Jerry) Harasewych, a newly appointed associate curator in mollusks at the U.S. National Museum in Washington, D.C. For over twenty years The Nautilus served as the official organ of the American Malacological Union at no cost to their organization. Proceed- ings of the meetings were published by them from 1932 to 1953 in the same format as that of The Nautilus. Like all growing organizations, the AMU eventually required its own periodical and now issues the extensive American Mala- cological Bulletin {vo\. 1, no. 1, July 1983) under the capable editorship of Dr. Robert S. Prezant of the University of Southern Mississippi. The life of an editor is not easy, especially in the case of scientific journals that constantly need financial help. The editors of the privately owned Nautilus were no exception, and they had to make repeated appeals for subscribers to "pay up." (Pilsbry and Johnson, 1893). In my 28 years of experience as an editor I had to face financial and production jiroblems, and I some- times found manuscripts poorly written, badly organized, without nomenclatorial knowledge and often lacking references to the very same subject. It is work getting some of them into shape, and sometimes there is little thanks. There is an old adage that "the next time you see an understanding and beloved editor, you will see him laid out horizonally and surrounded by flowers." But there are rewards -a sense of accomplish- ment and the joy of creating- a sense of helping others to participate in "our glorious science of conchology" as Averell put it so often in his editorials of a hundred years ago. A few years ago I gathered together and published a collec- tion of interesting articles and reproductions of ancient advertisements (Abbott, 1975). In The Best of The Nautilus one can sense the spirit of comradship among our early contributors. The Nautilus is for both the lieginning scientist, the enthusiastic amateur with new discoveries, as well as for the accomplished malacologist on the "cutting edge of the science" and immersed in "the state of the art." It is still a joint and cooperative effort. Let's hope that The Nautilus goes on for another 100 years! -R. Tucker Abbott, Melbourne, Florida LITERATURE CITED Abbott, R. Tucker. 1958. From the Pilsbry Chair of Malacology. The Nautilus 71(3):100-103. (editor). 1975. The Best ofTheNnutilus. 280 pp., American Malacologists, Inc., Greenville, Delaware. 1979. Bernadine Barker Baker -A Tribute upon her Retirement. The Nnutiliis 93(2-3):ii. 1983. Charles B. Wurtz-An Obituary. The Nautilus 97(1 ):43. portrait. f331 RENEWAL OF SUBSCRIPTION Enclosed find Two Dollars for one year's subscription to THE NAUTILUS, 234 Berkeley Street, Boston, Mass. Name, Address, Notice.— The rules now governing second-class matter do not permit publications to be mailed to subscribers in arrears. 8 THE NAUTILUS January 31, 1986 \'(il. 10(1(1) Abbott, R. Tuckor ami Chas. B. Wurtz. 1971. Horace Burrington Baker, 1889-1971. (An ol)ituary). The Nautilus 85(l):l-4. 3 portraits. Avereli, William D. (1899). Pospectus for Tlie Nautilus. 1 page, privately printed. Mt. Airy, Philadelphia, Penna. Baker, H. Burringti)n. 1958. Henry Augustus Pilsbry, 1862-1957-An Obituary. The Nautilu.'< 71(3):73-83, 3 portraits. 1958a. The Pilsbry Nautilus. The Nautilu.s 71(30):112-115. Gray, Arthur F. 1933. Charles WillLson Johnson, 18()3-1932. (An obituary). The Nautilus 46(4): 129- 136, portrait. .Jutting, W. S. S. Van B. and C. 0. van Regteren Altena. 1958. A list of Printed Malacological Journals. Basteria 22:10-15, 1 graph. Pilsbry, H. A. and C. W. Johnson. 1893. To Conchologists (a piea for support). The Nautilus 7(3):25-26. Wurtz, Charles B. 1958. Dr. Pilsbry and Fresh-water Mollusca. The Nautilus 71(3):84-87. TOM PULLEY AND THE TRAIN Thomas Edward Pulley, born October 15, 1916, Director of the Houston Museum of Natural Sciences since 1957, and a malacological student of the late William J. Clench of Harvard, passed away on Nove^riber 19, 1985, at the age of 69. In preparation for the 100th anniversary volume of The Nautilus, which in part is dedicated to Dr. Clench, I asked Tom to write a personal anecdote that typified his mentor. If my readers will recall, in my "Farewell to Bill Clench" (The Nautilus 9S(2):55-58), Bill drew a huge arrow in the sandy beach for his students that pointed to a waiting train. Tom Pulley has now caught up with the other students, and here is Tom's personal account of The Train written in July 198U. He asked that this last picture of Bill Clench be included- (RT A). Everyone who knew Bill was aware of his capacity for enthusiasm over a new shell. We who worked with him at the MCZ often wit- nessed the thrill of anticipation with which he opened each new package of shells and spread out the contents on the wide brown table. It was almost like watching the bright-eyed wonder of a small child at his first real Christmas. But Bill had the same happy way of appreciat- ing most of the other simple pleasures of life. He and Julia once invited me and my wife and 5-year-old son to a Sunday afternoon picnic; Bill would grill the hamburgers. When we arrived at his chosen site I was a bit surprised to see that we were on a nice grassy spot, but it was on the railroad-right-of-way. There were houses nearby, but they were mostly concealed by trees. I had known that Bill liked trains, and I soon learned that we were here to wave at the afternoon train when it passed by. We lighted the charcoal and opened some beer and cokes. When the coals were ready. Bill grilled the hamburgers and they were delicious. As we were finishing I saw that Bill key)t glancing at his watch. Soon we heard the faint whistle of William J. Clench, age 8G, in California. Vol. 100(1) JanuarvSl, 1986 THE NAUTILUS 9 one of the last steam trains, and Bill quickly jumped to his feet. He strained to view the big engine when it first came into sight around a distant bend, and long before it was near us he raised both hands in the air and began to wave. As the train approached he began jumping in the air and waving more excitedly. The engineer saw him and began tooting his whistle in a long series of short bursts. The excitement was catching, and by the time the engine had reached us my son was as excited as Bill, and I was amazed to find that I, too, was jumping and waving like a 5-year old. The smiles that were exchanged between Bill and the engineer as the train roared by led me to believe that this must have happened many times before. I understood Bill's joy that afternoon as the train passed by, because I had taken part in it. I later realized that Bill experienced this same kind of happy pleasure in many of the less dramatic events of daily life. To Bill, the thrill of seeing a new shell, meeting an old friend or making a new one, finishing another number of Johnsonia, or even telling one of his old bad jokes was comparable to his pleasure that day by the railroad tracks. - Tom Pulley, Julv 1984 JOSEPH ROSEWATER (1928-1985) A TRIBUTE AND BIBLIOGRAPHY Harald A. Rehder Smithsonian Institution Washington, D.C. 20560 On March 20, 1985, malacologists the world over lost an eminent and treasured colleague, and those close to him a dear friend. His un- timely death left a void in the ranks of malacolo- gists that will be hard to fill, and his associates at the Smithsonian still find it difficult to believe that we will not see him at any moment leaving his room or coming around the corner in the halls or in the range of the Division of Mollusks. Joe was born on September 18, 1928 in Clare- mont. New Hampshire, in the Connecticut River Valley, the only child of Joseph and Alice Tipping Rosewater. His mother had come to Claremont from the Rosewater home in Queens, Long Island, New York, to be with her parents at this time. Joe's father was an accountant with the City of New York, and his mother had been a Navy nurse in World War I. When Joe was eight years old his father died. Joe graduated in June 1946 from John Adams High School in South Ozone Park, Long Island, two months after the death of his mother. Joe's mother, before her death, had appointed her brother Charles to be Joe's legal guardian, and therefore Joe moved to his uncle's home in Claremont, New Hampshire. In the fall of 1946 Joe entered the University of New Hampshire at Durham, graduating in 1950. As a boy he had had an interest in animals, and had kept an aquarium in his room. His special interest in mollusks began apparently in FIG. 1. .Jo.suph RusuwaliT ui a I'ui-fiit iiitVirnial puse. 10 THE NAUTILUS January 31, 1986 Vol. 100(1) his junior year at the University when his zoology professors, Emery F. Swan and George M. Moore, persuaded him to undertake a mollus- can project for a master's degree. This is evidenced by the fact that Joe states that his first field work started in 1949 with collecting mollusks in New England. Joe began working on his Master's degree in the fall of 1950 but his plans were interrupted by the arrival of a draft notice in February 1951. That month he married Mary Carlson of Gilman- ton, New Hampshire, and in May he was formal- ly drafted and went to Fort Meade, Maryland, for basic training. In the fall he was sent to Fort Sam Houston, Texas, to receive further training as a medical technician in radiology. In early March 1952 Joe was sent overseas to Germany where until April 1953 he was a medical techni- cian in radiology in the 97th General Hospital in Frankfurt-am-Main. After his return from Germany and after leav- ing the service, Joe worked as an x-ray tech- nician at the Sacred Heart Hospital in Manchester, New Hampshire, and as an instruc- tor in biology at Mount St. Mary's College in nearby Hooksett until May 1955. That summer he enrolled at the University of New Hamp- shire, attending the summer school and then the academic year 1955-56, receiving a Master's degree in June 1956. In the fall of 1956 he entered Harvard Univer- sity as a graduate student to work under Dr. William J. Clench in malacology. Here, in the company of Richard I. Johnson, Richard W. Foster, Robert Robertson, Arthur H. Clarke and Arthur S. Merrill, he spent four happy and fruitful years under the genial and caring tutelage of Bill Clench and Ruth Turner. In the summer of 1957 Joe came to Washing- ton and spent three months in the Division of Mollusks of the U. S. National Museum as a Summer Intern, working on the family Pleuro- ceridae in connection with his doctoral studies. In the course of his studies he reorganized the North American members of the family Thiaridae in the museum collection. The following summer Joe accompanied Bill Clench on a field trip through Kentucky, Ten- nessee and Georgia, collecting and studying the freshwater faunas of the area. During his years at Harvard Joe was a Teach- ing Fellow in Biology, and in his last year he held a position as Curatorial Assistant in the Department of Mollusks. In the fall of 1959, Joe's final year at Harvard, Bill Clench and Joe were beginning to plan for Joe's immediate future after he received his degree in the coming June, and Bill wrote to me regarding a possible opening at the National Museum. For a year or more we had on file a job application form that Joe had filled out for a position in the Division, awaiting the time when we would get approval to hire an additional staff member. At this time the opportunity opened up to hire a malacologist under a grant that the National Museum had received from the Office of Naval Research and Atomic Energy Commission (ONR-AEC) to work on the marine faunas of the Pacific, especially that of the Marshall Islands- Bikini and Enewetak. This position was offered to Joe and he started working on January 1, 1960. In the year and nine months that he was on this contract he helped identify and arrange many lots of Indo- Pacific mollusks, and spent several days a week reorganizing and expanding the collections of Indo-Pacific marine mollusks, bringing the classification up to date and adding thousands of lots to the collection. In August 1961 Joe made his first visit to the Pacific attending the Tenth Pacific Science Congress in Hawaii. On October 2, 1961 Joe joined the staff as Associate Curator, and for the next twenty- three and a half years was a valued and impor- tant member of the scientific staff of the National Museum of Natural History. With the addition of Joe, the staff of the Divi- sion of Mollusks was once more at full strength, and this fact seemed to infuse new energj' into the divisional activities, stimulated in part no doubt by Joe's enthusiasm in his new position. Recorded in the annual report for that year was a notable increase in the number of specimens distributed in exchange, lent for study and iden- tified for correspondents. Although Joe's doctoral dissertation dealt with a freshwater mollusk, marine mollusks were his first love, as he told me in a letter he wrote when I offered him the opportunity of coming to the Smithsonian on the ONR-AEC grant. He had just finished preparing for the cataloguer the last of the marine collection of Vol. 100(1) January 31, 1986 THE NAUTILUS 11 the old Boston Society of Natural History that had been turned over to the Museum of Com- parative Zoology', and was in the midst of incor- porating Tucker Abbott's Philippine marine shells into the collection. When he came to work in the Division under the grant, sorting and classifying the Indo-Pacific moUusks was, therefore, a familiar task. After he and Ruth Turner had finished the western Atlantic Pin- nidae they had planned to monograph the Indo- Pacific species of the family. Now Ruth sug- gested that Joe do the paper alone, and this he did. He finished the project and it was published before the end of his contract. Following this he began studying the family Tridacnidae and the Indo-Pacific Littorinidae, both of which were eventually published in 1965 and in 1970/72. He continued his interest in these three families and published several papers. He was working on a study of the Eastern Pacific members of the Littorinidae at the time of his death, and also had underway a study of the bivalve family, Periplomatidae. During his years in the Division of Mollusks Joe went on several expeditions and field trips. In February and March 1963 he spent six weeks at the Enewetak Marine Biological Laboratory of the Atomic Energy Commission, and that winter he spent three months in the western Indian Ocean on the research vessel Te Vega as part of the International Indian Ocean Expedi- tion. In August and September 1966 Joe was in Australia spending most of the time carrying on field work in Western Australia with Barry C. Wilson, and in May and June 1970 he was a member of the Mariel King Memorial Moluccas Expedition on the R/V Pele. again together with Barry Wilson. In April 1974 he spent two weeks in Tunisia at the Mediterranean Marine Sorting Center, and in July 1976 he was a member of a party from the Smithsonian that spent ten days making a survey of the littoral fauna of Ascen- sion Island. Later he made several trips to Panama, and to the Pacific coast, including Alaska, in connection with his study on the Lit- torinidae of that fauna. His last trips away from Washington were to the Harbor Branch Labora- tory at Fort Pierce, Florida where for several years he carried on field studies on Periploma and the Littorinidae. Joe was a member of the American Malacolo- gical Union from 1957 on and was its President in 1969. He was also a member and past presi- dent of the Biological Society of Washington, the Society of Systematic Zoology of which he was treasurer 1963-1966, and the Paleontolo- gical Research Institute, Ithaca, New York. He was a member of the National Capital Shell Club and its president in 1965. He served as a Con- sulting Editor of The Nautilus from 1972 to 1984, and contributed 17 articles to that journal. In the Museum he represented the Depart- ment of Invertebrate Zoology on several com- mittees-the Professional Advancement Evalu- ation Committee, the Council of the Senate of Scientists, and the Advisory Committee on the Naturalist Center. On the Departmental Collec- tions Advisory Committee he represented the Division of Mollusks. Joe is survived by his wife Mary Carlson Rosewater and three children, Katherine Louise Rosewater (Waitt) of Sandown, New Hamp- shire, Gail Ann Rosewater of Rockville, Mary- land, and Carl Joseph Rosewater of Kensington, Maryland, and a granddaughter, Megan Rosewater- Waitt. Bibliography of Joseph Rosewater 1957. A comparative study of development in the Peri- winkles. Abstracts of student research projects. Woods Hole Embryology Counie. 1957:14-15, 1 table. 1958a. The Family Pinnidae in the Western Atlantic. Johnsonia 3(38):285-326, 23 plates (with Ruth Turner). 1958b. Book Review: Olsson, A., et al. Pliocene Mollusks of Southern Florida, etc. Johnsonia 3(38):328. 1959a. Mollusks of the Salt River, Kentucky. The Nautilus 73(2):57-63. 1959b. Intertidal stranding of Clione limacina in Massa- chusetts. TheNautUux 73{2):76-n. 1959c. Calvin Goodrich, a bibliography and catalogue of his species. Occasional Papers on MoUusks. Department of Mollusks. Museum of Comparative Zoology. Hanmrd University. 2(24): 189-208, 1 photograph, 1 plate. 1959d. A Pleurocerid index to past collecting in the Ohio and Tennessee River Basins with some thoughts for the future. Annual Report American Malacological Union for 1958, Bull. 25:23-24. 1960a. Egg mass and gross embryology' of Pleurocera canalicutata (Say, 1821). Annual Report .\merican Mala- cological Union for 19.59, Bull. 26:10-12. 1960b. Proposed further use of the Plenary Powers in the case of the Generic name Pleurocera Rafinesque, 1818 (Class Gastropoda). Z.M.(S)83. Bulletin of Zoological NoTnenclature 17(6-8):171-172. 1961a. Preliminary observations on the nervous systems of 12 THE NAUTILUS January 31, 1986 Vol. 100(1) some Pleuroceridae. Annunt Report American Mahi- cological Union for 1961, Bull. 28:10-11. 1961b. Comments on the proposal to place the Generic name Gari Schumacher, 1817, on the Official List, Unemended, Z.N.(S)1461. Bulletin of Zoological Nomenclature 18(5):303. 1961c. The Family Pinnidae in the Indo-Pacific. Indo- Pacific Mollwica l(4):17.'5-226, pis. 135-171, figs., maps. 1963a. Resistance to dessication in dormancy by Tectariux muricatuit. The Nautilus 76(2):111. 1963b. An effective anesthetic for giant clams and other mollusks. Turtux News (Chicago) 41(12):300-302, 2 figs. 1963c. Problems of species analogues in world Littorinidae. Annual Report Anifrican Malacolociical Union for 1963, Bull. 30:.5-6. 196.5a. The Family Tridacnidae in the Indo Pacific. Indo- Pncific Mollusra l(6):347-396, pis. 263-293. 19651). Studying living Tridacnidae in the Marshall Islands. Annual Report American Malacological Union for 1965, Bull. 32:18-19. 1966a. Reinstatement of Melarhaphe Menke, 1828. The Nautilus 80(2):37-38. 1966b. The Giant Clams. Australian Natural History 1.5(8): 250-256. 5 figs. 1967a. Indo- West Pacific Littorinidae. Aurnial Report American Malacological Union for 1966, Bull. 33:27. 1967b. Book Review: Halstead, B. W. 1965. Poisonous and Venomous Marine Animals of the World, vol. 1, Inverte- brates. Science 156(3776):795 (with M. L. .Jones, R. B. Manning, D. L. Pawson, and K. Ruetzler). 1968a. The Zoological Taxa of William Healey Dall. Bulletin U. S. National Museum 287:1-427 (with K. J. Buss and F. A. Ruhoff). 1968b. Itinerary of the Voyage of H. M. S. Blossom. The Veliger 10(4):'350-352. 1968c. Book review: Hyman, L. H. 1967. The Invertebrates, vol. VI. Mollusca 1. Science 160(3823):62-63. 1968d. Book review: An English-Classical Dictionary for the use of Taxonomists. Systematic Zoology 17(3):334. 1968e. Notes on Periplomatidae (Pelecypoda: Anomalodes- mata) with a geographical checklist. Annual Report American Malacological Union for 1968, Bull. 35:37-39. 1969a. (iross anatomy and classification of the commensal gastropod, CaledonieUa montrouzieri Souverbie, 1869. The Veliger ll(4):345-350, text figs. 1-4, pi. 55. 1969b. George Mitchell Moore. 1906-1968. [Obituary]. The Nautilus 9,ZC[):\\\. 1969c. Comment on a review by ,J. D. Rising of R. E. Black- welder, Taxonomy, a Text and Reference Book. Systema- tic Zoology 18(3):361-362. 1969d. Malacological collections-development and manage- ment. In: Natural History Collections, Past- Present - Future, Symposium of the Biological Society of Washing- ton. Proceedings of the Biological Society of Washington 82:663-670. 1970a. The Family Littorinidae in the Indo-Pacific. Part I. Subfamily Littorininae. Indo-Pacific Mollusca 2(11): 417-506, 64 plates. 1970b. Monoplacophora in the South Atlantic Ocean. Science 167(3924):1485-1486, fig. 1970c. Another record of insect dispersal of an Ancylid snail. TheNautihis 83(4):144-145. I97()d. The role of the professional in Malacologj-. Pitts- hurgh Shell Club Bulletin 5:12-13. 1970e. Book Review of: .Johnstone, Kathleen Yerger. Col- lecting Seashells. Grosset and Dunlap. Illus., 198 pp. Atlantic Naturalist 25(4):187-188. 1970f. Discussion of: R. T. Abbott, Eastern Marine Mol- lusks. In: American Malacological Union Symposium on Rare and Endangered Mollusks. Malacologia 10(1):49. 1972a. The Family Littorinidae in the Indo-Pacific. Part II. The Subfamilies Tectariinae and Echinininae. Indo-Pacific Mollusca 2(12):507-534, 21 plates. 1972b. Book Review: Wilson, Barry R. and Keith Gillett. 1971. Australian Shells. National Capital Shell Club Newsletter February 1972, 2 pp. 1972c. Book Review: Brost, F. B. and R. D. Coale. 1971. A guide to Shell Collecting in the Kwajalein .-Xtoll. National Capital Shell Club Newsletter February 1972, 2 pp. 1972d. The Amphi-Atlantic distribution of Liltorina melea- gris. The Nautilus 86(2-4):67-69, plate, map. table (with G. .J. Vermeij). 1972e. Teratological Littorina scahra angulifera. The Nautilus 86(2-4):70-71, plate. 1972f. Life spans of Mollusca. In: Altman, P. L. and D. S. Dittmer. 1972. Biology Data Book, edition 2, vol. 1, pp. 234-235. Federation of American Societies for Experi- mental Biologj', Bethesda, Maryland. 1973a. Book Review: Wilson, B. R. and Keith Gillett. 1971. Australian Shells. The Quarterly Rerieiv of Biology 48(l):42-43. 1973b. Book Review: Graham, Alastair. 1971. British Proso- branch and other Operculate Gastropod Molluscs. The Quarterly Review of Biology 48(1):43. 1973c. A Source of Authors and Dates for Family Names of Gastropods. The Veliger 16(2):243. 1973d. Replacement pages for: The Family Littorinidae in the Indo-Pacific, Part II, the Subfamilies Tectariinae and Echinininae (Indo-Pacific Mollusca 2(12):509-516). Indo-Pacific Mollusca 3(14):63-70. 1973e. More on Penis Shedding Among L('/ion)ja. Neir York- Shell Club Notes, no. 196, November 1973, p. 7. 1974a. Studies on Ascension Island Marine Mollusks. Bulk- tin American Malacological Union for 1973, pp. 30-32. 1974b. Phylogeny of Littorinidae. Littorinid Tidings. Occasional Newsletter of Littorinidae Research Grotip, no. 1, pp. 10-11. 1975a. An Annotated List of the Marine Mollusks of Ascen- sion Island, South Atlantic Ocean. Smithsonian Contri- butions to Zoology 189:1-41, figs. 1-24, 3 tables. 1975b. The Marine Commensal Gastropod, CaledonieUa mxmtrouzieri (Prosobranchia: Hipponicacea) in Thailand. The Nautilus $%iyM. 1975c. Mollusks of Gatun Locks, Panama Canal. Bulletin American Malacological Union for 1974, pp. 42-43. 1975d. Book Review: R. Scase and E. Storey. 1975. The World of Shells. National Capital Shell Chih Neu'sletter. September, 1975. p. 10. 1976a. William Healey Dall, The Legacy He Left for Mala- cology. Bulletin American Malacological Union for 1975, pp. 4-6. 1976b. The NMNH-STRI Survey of Panama 1971-1975. Bulletin American Malacological Union for 1975, Vol. 100(1) January 31, 1986 THE NAUTILUS 13 pp. 48-50. 1976c. Letter U> the Editor. AiistnilniH Slifll ,\eirx No. Hi, p. 2. 1976(1. Book Review: Ruth Fair. Shell Collectors' Cuide. National Capital Shtil Cliili Ni-iivlettrr. Septemlier 1976, pp. 11-12. 1976e. Application to use Plenary Power to designate the type of Pleurocera. Bulletin of Zmilogiral Nnnirnrlatiirr 33(2):10.->-n3. 1977. Bicentennial Booty. National Capital Shrll CluliNcivs- letter. February 1971, p. 8. 1978a. Zoogeography of West African Littorinidae. Bulletin Ameriran Malacological Union for 1977. pp. .31-34. 1978b. A Case of Double Primary Homonymy in East Pacific Littorinidae. The Nautilus 92(3):123-12.5. 1978c. A Malacological Ecpedition to the Moluccas, National Geiyraphir Society Research Reports nf 1969 Projects. pp. .515-523. (With Barry R. Wilson). 1978d. Book Review; Kirtisinghe. P. 1978. Sea Shells of Sri Lanka. National Capital Shell Club Newsletter. September 1978, p. 11. 1979a. A Reconnaissance of West American Littorinidae. Bulletin American Malacological Union for 1978, p, 55. 1977b. Rectifications in the Nomenclature of some Indo- Pacific Littorinidae. Proceedings Biological Society of Wa.-ihington 92(4):773-782. (with W. F. Ponder). 1979c. Further Comments on the Proposed Designation of a Type Species for Pleurocera Rafinesque, 1818. Z.N.(S)83. Bulletin Zoological Nommclature 36(3):139-146. (with J. P. E. Morrison, A. H. Clarke, G. M. Davis, and C. B. Stein). 1980a. Predator Boreholes in Periploma margaritaceuni with a brief survey of other Periplomatidae (Bivalvia: Anomalodesmata). The VV/7>r 22(3):248-251. 1980b. A Close look at Littorina Radulae. Bulletin Ameri- can Malacological Union for 1979, pp. 5-8, 8 figs. 1980c. Subspecies of the Gastropod Littorina .•icahra. The Nautihis 94(4):158-162, 12 figs, 1980d, Book Review: Melvin, A. G. and Melvin, L. S. 1980. 1000 World Sea Shells, Rare to Common with Values. National Capital Shell Club Newsletter. September, p. 4. 1981a. The Family Littorinidae in Tropical West Africa. Atlantule Reports 13:7-48, 6 plates, 2 tables. 1981b. A New Aid to Taxonomic Research on Mollusks. Bulletin American Malacological Union for 1980, p, 67. 1981c. Changes in Shell Morphology of Post Larval Tri- dacna gigas Linne (Bivalvia: Heterodonta). Bulletin American Malacological Union for 1980, pp. 45-48, figs. 1981(1. Malacological .'lourney into Cuba, 1980, The Nautilus 95(4):1.59-162, 1981e. Rectifications in the Nomenclature of some Indo- Pacific Littorinidae -II. Proceedings Biological Society of Washington 94(4):1233-1236. 1982a. A new species of Hippopus (Bivalvia: Tridacnidae). The Nautilus 96(l):3-6, 4 figs. 1982b. A new species of the Genus Echininus (Mollusca: Littorinidae: Echinininae) with a review of the Subfamily. Proceedings Biological Society of Washington 95(1): 67-80, 6 figs, table. 1982c. Book Review: R. K. Dell, Sea Shore Life. Natiaual Capital Shell Newsletter. No. 70, Autumn, p. 15. 1983a. Review of Hawaiian Pinnidae and Revalidation of Pinna exquisita Dall, Bartsch, and Rehder, 1938 (Bivalvia: Mytiloida). Pacific Science 36(4):453-45S. 1983b. Review of: .lames Graham Cooper- Pioneer Western Naturalist, by Eugene Coan. The Nautilus 97(l):44-45. 1983e. Another Bivalve-/l/)/(?-o(^j;(7 Association with Com- ments on Adaptive Significance of Oddly Shaped Lepto- nacea. American Malacological BuUetiii 1:90-91. 1984a. A bibliography and list of the taxa of Mollusca introduced by .Joseph P. E. Morrison (December 17, 1906- December 2, 1983). The Nautilus 98(l):l-9. 1984b. A new species of Leptonacean Bivalve from off Northwestern Peru (Heterodonta: Veneroida: Lasaeidae). The Veliger 27(l):81-89. 6 figs. 1 table. 1984c. Burrowing activities of Periploma margaritaceum (Lamarck, 1801) (Bivalvia: Anomalodesmata: Periploma- tidae). American Malacological Bulletin 2:35-40, 1 plate, 1 table. 1984d. Bermuda marine Mollusk type specimens transferred to the Smithsonian. The Nautilus 98(4):151-153. 1985. Epizoan Communities on Marine Turtles 1. Bivalve and Gastropod Mollusks. Marine Ecology 6(2):127-140, 4 figs., 1 table (with .1. Frazier et al.). 14 THE NAUTILUS lanuary 31, 1986 Vol. 100(1) DESCRIPTION OF THE HABITAT OF THE ENDANGERED MUSSEL PLETHOBASUS COOPERIANUS Andrew C. Miller Barry S. Payne U.S. Army Engineer Waterways Experiment Station P.O. Box 631 Vicksburg, MS 39180 and Terry Siemsen Navigation Planning Support Center P.O. Box 59 Louisville, KY 40201 ABSTRACT A total of 26 species ofunionids, in addition to the Asian Clam. Corbicula, were collected at a mussel bed on the Ohio River near Ohnsted, Illinois. The bed was dominated by Corbicula (1475/m^, 128.2 g/m^ of Tissue Dry Mass, TDM) and Fusconaia ebena (66.0/m^ and 39.7lm^ of TDM) although eight other unionids. Amblema plicata, Elliptio dilatatus, Leptodea fragilis, Ligumia recta, Mega- lonaias gigantea, Pleurobema cordatum, Potamilus alatus, and Truncilla donaci- formis were also judged comynon. In addition, three live specimens of the Fed- erally endangered Orange-footed Pimpleback Mussel, Plethobasus cooperianus, were identified. All appeared healthy and were adult-sized, ranging in total shell length from 68 to 7U mm. While there are no records of recent recruitment for P. cooperianus, it appears that this mussel bed, which contains a diverse cammunity of bivalves in addition to a dense population o/' Corbicula, provides good habitat for this particular species. Three live specimens of the Endangered Plethobasus cooperianus (Lea 1834), the Orange- footed Pimpleback Mussel, were found in September 1983 during a survey of a gravel bar in the Ohio River near Olmsted, Illinois. In the summer of 1982, Williams and Schuster (1982) collected mussels at this site and found a single live P. cooperianus. However, other workers brailed this gravel bar and found no live endan- gered species (Neff & Pearson 1980, Williams 1969). The purpose of this paper is to document the existence of this species, and to provide in- formation on habitat characteristics and com- munity structure where it was collected. The historical range of P. cooperianus in- cluded the Ohio River from western Pennsyl- vania to southern Indiana; the Wabash River below Mt. Carmel, Illinois; the Cumberland River from Cumberland County, Kentucky, to the vicinity of Nashville, Tennessee; the lower Clincli River in Anderson County, Tennessee; and the Tennessee River from near Knoxville, Tennessee, to Kentucky Lake, Benton County, Tennessee. It has also been recorded from the Caney Fork, Holston, and French Rivers in Tennessee and from the Green and Rough Rivers in Kentucky. At present, P. cooperianus is restricted to the Ohio and lower Wabash Rivers where it is uncommon and to the lower Tennessee River in Alabama and western Tennessee where it is also uncommon. The shell of P. cooperianus is up to 87 mm long, 75 mm high, and 45 mm wide; it is heavy, subcircular, and tuberculate. The periostracum is chestnut colored and the nacre is either white or faintly pink and iridescent posteriorly. Superficially this species resembles Q. pustulosa, but the latter species is usually smaller and has fewer and relatively larger tubercles which tend to be laterally rather than radially extended. In addition, Q. pustulosa exhibits white nacre and young specimens have prominent green rays, whereas in P. cooperia- nus the rays are obscure and narrow. The most definitive characteristic in P. cooperianus is the bright orange viscera, which can be seen in live specimens by gently prying the valves apart (Clarke and Fuller 1983). Wilson and Clark (1914) reported collecting two gravid females in June from the Cumber- Vol. 100(1) January 31, 1986 THE NAUTILUS 15 land River; evidently it is a summer breeder although the host fish is unknown. Although ecological data are virtually non-existent, this is a large river species which has been collected from sand and gravel substrate in shoals and riffles. Methods and Study Area On 26-29 September 1983 molluscs were col- lected from a reach of the Ohio River below Lock and Dam 53 near Olmsted, Illinois. The study area included the upper portion of a mussel bed delimited by Williams (1969) and Williams and Schuster (1982). During the four- day study bivalves were collected by hand along the shore, with the use of a 5-ft brail bar, and by an experienced shell diver using underwater breathing apparatus. As part of this work the diver completed three 30-min qualitative searches, and collected six 0.25- m^ quadrat samples from each of four sites on the mussel bed. This project was undertaken for the U. S. Army Engineer District, Louisville, to provide information for a Feasibility Report and Envi- ronmental Impact Statement for the Lower Ohio River Navigation Project. The gravel bar where mussels were collected was about 3 miles long and followed the Illinois shoreline. Substrate consisted of densely packed coarse sand and gravel. Particle sizes varied from less than 1.0 cm to more than 10.0 cm with the greatest fraction of total weight in the 1- to 3- cm range. Water depths where P. cooperia- nus were taken ranged from approximately 3 to 6 m deep. The study site was Ohio River Mile 966.6 to 967.2, approximately 4 miles downriver of Lock and Dam 53; navigation and barge fleet- ing were taking place in the immediate area. No recent evidence of sedimentation, physical abra- sion, or propeller wash from navigation activi- ties, were noted at this portion of the gravel bar. Results Using qualitative and quantitative techniques, 26 species of unionids, in addition to the Asian Clam, Corhicula, were collected at or adjacent to the gravel bar (Table 1). Although nine species were judged common, the most abun- dant bivalves were Corhicula, Amblema plicata, Fusconaia ebena, Lampsilis teres. Megalonaias gigantea, and Potomilus alatios. TABLE 1. Bivalves ccillected at a gravel bar near Olmsted, Illinois. 26-29 September 1983. All taxa were collected alive. Scientific Name Family Unionidae Actlnonaias carinata Amblema plicata Cyclonaias tuberculata Ellipsaria llneolata Elliptio crassidens Elliptio diiatatus Fusconaia ebena Lampsilis ovata Lampsilis teres Lasmigona complanata Leptodea f ragilis Ligumia recta Megalonaias gigantea Obliquaria ref lexa Qbovaria olivaria Flethobasus cooperianus Plethobasus cyphyus Pleurobema cordatum Potamilis alatus Quadrula metanevra Quadrula nodulata Quadrula pustulosa Quadrula quadrula TritoRonia verrucosa Truncilla donacif ormis Truncilla truncata Family Corbiculidae Corbicula manilensis Presence* UC C nc FC FC UC c UC c UC c c c FC UC UC UC c c FC FC FC FC UC C DC * Presence, for this study: UC = Uncommon, < 6 collected FC = Fairly common, 6-20 collected C = Common, > 20 collected A total of 14 species of unionids were collected in 24 0.25-m^ quadrat samples (Table 2). Total unionid density ranged from 28 to 124/m^ (X = 60). Numerically, F. ebena (X = 46/m^) and Trun- cilla donaciformis (X=12/m^) dominated the unionids. Two species, Q. metanevra and Q. nodulata, judged fairly common in the study area, were found in only 1 of the 24 quantitative samples. The gravel bar was numerically domi- nated by Corbicula, with numbers ranging from 556 to 2628/m' (X=1475). Average tissue dry mass (TDM) for Corbicula (128.2 g/m') was ap- proximately three times that for total unionids (X = 39.7). Because of their large size and high 16 THE NAUTILUS January 31, 1986 Vol. 100(1) TABLE 2. Bivalves collected in 24 0.25-ni' quadrat samples at a gravel bar near Olmsted, Illinois, 28-29 September 1983. Species Occurrence* 24 No/m^ 1475 TDM/m^ Corbiaula manilensis 128.2 Total unlonlds Ih 66.0 39.7 Fusconaia ebena 24 45.5 27.4 Truncilla donaaiformis 24 11.5 0.4 Leptodea fvagilis 11 2.5 1.3 Obliquaria reflexa 9 1.8 0.8 Quadrula pustulosa 8 1.5 1.4 Amblema plioata 6 1.0 3.1 Ellipsaria lineolata 7 1.3 1.5 Tritogonia verrucosa 3 0.5 0.4 Lcmpsilis ventrioosa 6 1.2 0.03 Quadrula quadrula 5 0.8 1.3 Quadrula metanevra 1 0.2 0.1 Quadrula nodulata 1 0.2 0.1 Megalonaias gigantea 2 0.3 1.3 Aatinonais aarinata 1 0.2 0.3 * Occurrence - Number of quadrats with live organisms (total quadrats collected = 24) . numbers, F. ebena represented the majority, about 70% (27.4 g/m^ of the total unionid biomass. Although numerically dominant, T. donaciformis. because of_ its small size, represented only about 1% (X = 0.4 g/m^) of the unionid biomass. Based upon these quantitative collections, evidence of recent recruitment (presence of clams less than 12 months old judg- ing from their small size and unweathered ap- pearance) was observed for 12 of the 14 unionids collected in the quadrats. The three specimens of P. cooperianns were obtained near Ohio River Mile 967.2 by the diver during two of the three 30-min qualitative searches of the bar. The diver had been in- structed to concentrate on pustulate organisms or species judged uncommon by our earlier work. On the second search, he obtained eight species, including one P. cooperianiui and two Q. ptistulosa. During the final search he retrieved eleven species, including two P. cooperianus and two Q. pustulosa. The Orange-footed Pimple- back was not found with the brail, on any of the shoreline searches, or in the quadrat samples. Conchological data on the three live P. cooperianus are as follows: Length (mm) Height (mm.) Width (mm) 74 63 37 68 60 38 71 64 42 By gently prying the valves apart, it was ob- served that soft tissues were healthy looking and certainly not moribund. Discussion This gravel bar was dominated both in num- bers and biomass by Corbicula; evidently this had no effect on the unionids or P. cooperianus. In addition, it appears that requirements for food, water quality, and substrate for adult P. cooperianus must differ very little from the 25 other unionids inhabiting this bed. In a study of 21 species in Michigan streams, Strayer (1983) concluded that microhabitat differences were not discernible for 21 of the unionids. In addi- tion, two other species, the fairly uncommon Vol. 100(1) January 31, 1986 THE NAUTILUS 17 Cumberlandia monodonta and the endangered Lampsiliti higginsi, have also been collected from very diverse and densely populated mussel beds in the Upper Mississippi River. Cumber- landia monodonta was usually taken in samples where 50% of the sites had at least 20 species of common or wide-ranging mussels. Lampsilis higginsi was in association with 17 or more com- mon unionids at 50% of the reported sites in the upper Mississippi River (Nelson and Freitag 1979). Wilson and Clark (1914) judged P. cooperia- nus to be "not rare" during their survey of the Cumberland River. However, Neel and Allen (1964) were unable to locate this species during a follow-up study on the same river. It is ap- parent that the historical range of P. cooperia- nus has diminished and the species is in danger of becoming e.xtinct. Some of the reasons often cited for the loss of mussels in large rivers in- clude: sedimentation, navigation activities, pollution, reservoir construction (many with deoxygenated, low pH, and cold water releases), and loss of fish hosts (Fuller 1974). Conversion of large, free-flowing rivers such as the Ohio to navigable waterways altered the habitats of many benthic organisms. However, some extensive areas with suitable substrate, flow, water quality, and nutrients still exist which support recruiting unionid communities. For the endangered P. cooperianus, the studied gravel bar on the Ohio River appears to meet requirements necessary for maintenance of adults. The range reduction for P. cooperianus is probably related to species-specific problems of reproduction or recruitment. Conversion of the Ohio River from a freely-flowing river to a con- trolled waterway probably modified the avail- able habitats and could have had detrimental effects on host fish. However, it is also possible that this species is slightly more sensitive to altered habitats than the other thick-shelled unionids at the mussel bed. Perhaps conditions have degraded to a point that P. cooperianus can maintain itself but does not possess energy reserves necessary for production of adequate numbers of glochidia. Similar problems may also be blamed for loss of species in the genus Dysnamia, which were once commonly collected in riffles or shoals in large rivers (Stansbery 1970). Riffle and shoal habitats are now almost non-existent in large rivers in North America; their disappearance could be blamed for loss of recruiting popula- tions oi Dysnotnia and Plethobasus. A possible solution for long-term maintenance of P. cooperianus is artificial propagation (Isom and Hudson 1982) and translocation of im- mature forms to suitable habitats. Projects such as these will be important for separating direct effects of habitat alteration from indirect effects such as reduction in the availability of the host fish. Acknowledgments Funds for this work were provided by the Navigation Planning Support Center, U. S. Army Engineer District, Louisville in Louisville, Kentucky. LITERATURE CITED Clarke, A. H., and Fuller, S. L. H. 1983. A Field Guide to the Endangered Mussels. Technical Report in Preparation, U.S. Army Engineer Waterways Experiment Station, Vicksburg, Miss. Fuller, S. L. H. 1974. Clams and Mussels (Mollusca: Bivalvia), In: Hart, C. W., Jr., and S. L. H. Fuller, eds.. Pollution Ecology of Fresh-Water Invertebrates. Academic Press, New York. Isom, B. G,, and Hudson, R. G. 1982. In vitro Culture of Parasitic Freshwater Mussel Glochidia. The Nautilus 96(4):147-151. Neel, J. K., and Allen, W. R. 1964. The Mussel Fauna of the Upper Cumberland Basin Before Its Impoundment. Mala- rologia 1:427-459. Neff, S. E., and Pearson, W. 1980. Aquatic Ecology and Water Quality Studies for the Lower Ohio River Naviga- tion Study, Unpublished report submitted to the Louisville District Office, Louisville, Ky. Nelson, D. A., and Freitag, T. M. 1979. Ecology, Identifica- tion, and Recent Discoveries of Higgin's Eye {Lampsilis higginsi]. Spectacle Case (Cumberlandia monodonta). and Fat Pocketbook (Potamiius capax) Mussels in the Upper Mississippi River. In: Rasmussen, J. L., ed., Proceedings of the UMRCC Symposium on Upper Mississippi River Bivalve Mollusks. J. L. Rasmussen, ed. pp. 120-145. Stansbery, D. H. 1970. Eastern Freshwater Mollusks (I) The Mississippi and St. Lawrence River Systems. Malacologia 10(l):9-22. Strayer, D. L. 1983. Notes on the Microhabitats of Unionid Mussels in Some Michigan Streams. Amer. Midi Nat. 106(2):411-415. Wilson, C. B., and Clark, H. W. 1914. The Mussels of the Cumberland River and Its Tributaries. Report U. S. Com- mission Fisheries for 1912 and Spec. Papers, pp. 1-63 (separately issued as Bureau Fisheries Document No. 781). Williams, J. C. 1969. Mussel Fishery Investigation, Ten- 18 THE NAUTILUS January 31, 1986 Vol. 100(1) nessee, Ohio, ;ind (Sreen Rivers. Final Report. State of Kentucky Project Number 4-19-R, Report to U. S. Fish and Wildlife Service, 107 pp. Williams, J. C, and Schuster, G. A. 1982. Fresh-Water Mussel Investigations of the Ohio River, Mile 317.0 to Mile 981.0, Report Submitted to the U. S. Army Engineer Dis- trict, Louisville; Kentucky Department of Fish and Wild- life Resources; Division of Fisheries, U. S. Fish and Wild- life Service, Areas 3 and 4. CHARACTERIZATION OF A BIVALVE COMMUNITY IN THE TANGIPAHOA RIVER, MISSISSIPPI Andrew C. Miller Barry S. Payne Environmental Laboratory U.S. Army Engineer Waterways Exp. Station P.O. Box 631, Vicksburg, MS 39180 and David W. Aldridge Department of Biology North Carolina A&T State University Greensboro, NC 27411 ABSTRACT On 5 July 1983 a quantitative collection of bivalves was made at the Tangipahoa River in southern Mississippi. Although the river water was extremely soft and the substrate and water quality were good, a diverse and healthy mollv^k com- munity was found. In addition to the Asian Clam. Corbicula, five species of unionids fFusconaia flava, Quadrula pustulosa, Elliptic crassidens, Tritogonia verrucosa and Villosa sp.J were taken. Numerically, Corbicula (8.93/m.^), out- numbered total unionids (1.2Jt/m.'). However the dominant feature of the benthic biomass were the unionids with a total tissue dry mass of 170^.5 m.g/m' as com- pared with 1015.0 mglm} for Corbicula. habitats. The purpose of this research was to characterize, using quantitative field and laboratory techniques, a bivalve community in the Tangipahoa River in southern Mississippi. Mississippi bivalves were first investigated by Hinkley (1906), later by Grantham (1969), and then Stern (1976) who concentrated on the unionids of the Lake Maurepas-Pontchartrain- Borgene drainage system. Recent mollusk studies in Mississippi include an investigation of mussels in the Big Black River (Hartfield and Rummel 1984), distribution of Corbicula (Hartfield and Cooper 1983), and a discussion of diversity and abundance of mussels in the south- west portion of the state (Hartfield and Ebert 1984). The above-cited studies, while providing useful information on species composition and ecology, have been directed toward qualitative assessments of the molluscan fauna. Quantita- tive unionid studies, either in Mississippi or other parts of the country, are relatively uncom- mon. The primary reason for this is the diffi- culty of collecting significant numbers of live mussels from a measured area. Benthic grab samplers usually do not retrieve enough sub- strate to provide reliable population estimates; in addition, they do not function well in gravel Study Area The Tangipahoa River originates in Lincoln County, flows south through Amite and Pike counties, then enters Lake Tangipahoa approxi- mately 4 miles southeast of McComb in southern Mississippi. Below Lake Tangipahoa the river flows in an easterly direction into Louisiana and ultimately enters Lake Pontchartrain near New Orleans. The study site (latitude 31°07'15"N., longitude 90°29'25"W., elevation 85.3 M. Mc- Comb S, Miss., US Geological Survey Quad- rangle) was located south of Lake Tangipahoa and 2 miles south of the town of Magnolia. In this area the river consisted of pools, riffles, and runs with steep, partially eroding banks. Canopy cover was usually complete and sur- rounding land was either undeveloped riparian forest or pasture. At the time of collection the Vol. 100(1) January 31, 1986 THE NAUTILUS 19 water depth was no greater than 1.0 m, except in pools, and widths ranged from 4 to 10 m. Based upon preliminary surveys, this site was judged to be one of the richest for bivalves when compared with adjacent rivers or other portions of the Tangipahoa River. Substrate at the study site consisted of gravel- ly sands. On 5 July 1983 chemical conditions of the river water were as follows: calcium hard- ness, 3.2 mg/1; pH, 6.8; and turbidity, 8.9 NTU. On 12 August at 1300 hr the air temperature was 27°C and water temperature was 24°C. Additional chemical data from the Tangipahoa River at Hwy 190 in Louisiana (taken from the Louisiana Stream Control Commission as cited by Stern 1976) further characterize this river as clear, well oxygenated, and with low dissolved solids and turbidity. Methods and Materials Mollusks were collected from two adjacent riffles by two workers during a 6-hr period on 5 July 1983. Unionids were obtained from both sites (214 m^) and Corbicula were taken only from a subsection of the downriver site (44 m^). Bivalves were the dominant feature of the com- munity; no snails or fingernail clams and only a few immature insects were collected. Collec- tions were made by hand or with a basket dredge, an open-ended wire basket that was pulled through the substrate and retained only objects equal to or greater than 1 cm in dia- meter. All mollusks were kept cool and returned to the laboratory alive. The majority of the specimens were used in a series of laboratory experiments on navigation effects that were part of the Environmental and Water Quality Operational Studies at the Waterways Experi- ment Station. In the laboratory, total shell length and height were recorded. Anterior and posterior mussels were cut with a scalpel or small knife and the viscera removed. Shells and viscera were dried for 24-48 hr at 75 °C before weighing to the nearest 0.1 mg on a Mettler balance. Regression equations for length, height, or tissue dry mass (TDM) were derived after transforming data to logio or logj with an IBM 4331 computer using the Statistical Analysis System. Results and Discussion At the study area on the Tangipahoa River, Corbicula, which was common in Mississippi by the late 1960's (Grantham 1969), represented 87.8% of the total bivalve fauna (Table 1). Asian Clams were common throughout the downriver site, even in shallow water on unanchored sands where unionids were never collected. Length- frequency analysis (Fig. 1) indicated that two major cohorts were present in early July. About 40% of the Corbicula were approximately 14-mm long; the second cohort ranged from 20 to 30 mm. Based on Corbicula collections made in May in the Altamaha River, Georgia, Sickle (1979) identified a cohort at 14 mm and a second at 22 mm. In that river the first year spawn had achieved 4 mm by August of the same year. In the Tangipahoa River the second year cohort was more slow growing and exhibited a wider range of lengths than did the first year group, findings which are consistent with Sickle (1979). A power curve (Fig. 2) provided the highest r^ value for the relationship between shell length and TDM. Sickle (1979) concluded that the rela- TABLE 1. Summary statistics for bivalves collected from two adjacent sites on the Tangipahoa River Pil^eC 3unty, Mississippi, 5 July 1983. Total Numbers Tissue TDM/m^ 1015.0 Dry Mass % 37.3 Shell Len gth, mm Species No. Collected 393 No./m 8.93 % 87.8 X 21.6 CV* 23.6 min 10.0 max Corbiau la 33.0 Fueconaia flava 158 0.74 7.3 531.1 19.5 46.0 16.1 31.3 70.0 Quadrula pustulosa 55 0.26 2.5 296.8 10.9 47.6 11.8 36.6 64.2 Eltiptio crassidens 31 0. U 1.4 605.5 22.3 105.9 10.9 84.0 126.0 Tritogonia vevruoosa 18 0.08 0.8 271.1 10.0 94.0 13.1 69.0 108.0 Villoea sp. 4 0.02 0.2 - - 52.9 15.3 43.8 61.1 Total unionids 266 1.24 12.2 1704.5 62.7 Total bivalves 659 10. 17 2719.5 Coefficient of variation 20 THE NAUTILUS January 81, 1986 Vol. 100(1) CORBICULA Lr 15 20 SHELL LENGTH. MM FK;. 1. Shell length-frequency distributions for Corhiaila collected from the Tangipahoa River. Pike County. Mississippi. 400 300 2001- 100 80 60 40 CO 20 GO S2 10 CORBICULA Y = 0.0047 X3 21 r2 = 0.97 _L J L J 12 5 10 SHELL LENGTH. MIVI 15 20 30 40 FIG. 2. Relationship between tissue dry mass and shell length for CorbicuUi collected from the Tangipahoa River. Pike County, Mississippi. >- CJ a LU az 30 r 25 20 h 15 10 o ac >■ o a o cc o CJ cc >■ o ¥- A QUADRULA PUSTULOSA tfk 0 20 30 40 50 60 70 SHELL LENGTH, MM 17 15 h 10 0 ^ FUSCONAIA FLAVA EL 0 30 40 50 60 70 80 SHELL LENGTH, MM 25 r 20 15 h 10 5 0 Uj 1 III I" TRITOGONIA VERRUCOSA 0 50 60 70 80 90 100 110 SHELL LENGTH, MM 15 r 10 ELLIPTIO CRASSIDENS 0 80 90 100 110 120 130 SHELL LENGTH, MM FIC. 3. Shell length-frequency distributions for four species of mussels collected from the Tangi[)ahoa River. Pike County, Mississippi. Vol. 100(1) January 31, 1986 THE NAUTILUS 21 tionship between length and TDM was best described with a power curve for Altamaha River fauna and collections made in the Mud River, West Virginia, by Joy and McCoy (1975). Although Corhicula numerically dominated the Tangipahoa River bivalve community, because of their smaller size they constituted only 37.3% of the total molluscan TDM (Table 1). Ficsconaia Jlava' , a widely distributed unionid in southern Mississippi and Louisiana (Stern 1976), was the most abundant unionid in the Tangipahoa River with densities equal to 0.74/m^ The second most abundant species, Quadrula pustulosa. (this southern form called refidgens Lea by some) has been collected in mud, sand, and gravel substrate in flowing and slack water habitats in this region (Stern 1976). Total shell length for F. Jlava ranged from 31.3 to 70.0 mm; length-frequency distributions (Fig. 3) suggest that a major cohort existed from 36 to 44 mm. For Q. pustulosa a cohort was identi- fied at approximately 45 mm with at least one below and possibly two cohorts above this range. For both of these Tangipahoa River unionids, the highest r' for length and TDM was 'Alias cerina Conrad. 1838. See Hartfield and Rummel (198.5). a function of X' (Fig. 4). Quadrula pustulosa ex- hibited a slightly higher ratio of TDM to shell length, an indication of greater tissue mass com- pared to length of shell, than did F. flava. While F. JJava and Q. pustulosa comprised only 7.3% and 2.5% of the bivalve community, because of their larger size they represented 19.5% and 10.9% of the total bivalve TDM, respectively. While Elliptio crassidens and Tritogonia ver- rucosa exhibited densities of 0.14 and 0.08/m^ respectively, the contribution of these larger bivalves to the total TDM was approximately equal to that of the previous two species (22.3% and 10.0%, respectively). Grantham (1969) reported that the former species had sporadic distribution in Mississippi, and Stern (1979) found E. crassidens common in headwater streams in this region. Average shell length of E. crassidens (105.9 mm) exceeded that for T. verrucosa (94.0 mm) by approximately 10%. A linear plot of total shell length to total shell height (Fig. 5) demonstrated a greater ratio of height to length for E. crassidens then T. verru- cosa at sizes greater than 97 mm. For individ- uals larger than 97 mm, the reverse was true; height to length was greater for T. verrucosa than for E. crassidens. In this section of the Tangipahoa River, F. 2250 2000 1750 1500 1250 1000 750 500 250 QUADRULA PUSTULOSA Y = 0.68 X2 - 411.4, r2 = 0.86 FISCONAIA FLAVA •• ,,^., •.: Y = 0.52 X2 - 416.1. r2 = 0.82 ^,l-J I I I I I I I _L 0 32 34 36 38 40 42 44 46 48 50 52 54 56 58 60 62 64 66 68 SHELL LENGTH. MM FIG. 4. Relationship between tissue dry mass and shell length for Q. fnistulosa and F.Jhim collected from the Tangipahoa River, Pike County, Mississippi. 22 THE NAUTILUS 80 - 70 - s 60 - S 50 OD 40 30 20 January 81, 1986 ELUPTIO CRASSIDENS Y = 0.46X + 12.3. r2 = 0.76 Vol. 100(1) TRITOGONIA VERRUCOSA Y = 0.68X - 11.7. r2 = 0.90 h^ _L _L _L _L _L 50 60 110 120 130 70 80 90 100 SHELL LENGTH. MIVI FIG. .5. Relationship between shell height and shell length for E. crassideiis and T. verrucosa col lected from Tangipahoa River, Pike County, Mississippi. flava, because of its numbers, and E. crassidens. because of its size, dominated the unionid TDM. The other three unionids, Q. pustulosa, T. verru- cosa, and Villosa sp., were a minor part of the unionid community. While the unionids domi- nated the bivalve biomass (62.7%), they were outnumbered by Corbicula (87.8%). In addition, the Asian clams, because of their rapid growth rates, exhibited higher productivity in terms of tissue growth per unit time than did the unionids. Since dissolved calcium is of obvious impor- tance for shell maintenance, it is significant that the Tangipahoa River supported a healthy com- munity regardless of soft water (3.2mg/( as CaCOj). In central New York, Clarke and Berg (19.59) collected no unionids in water with total alkalinity less than 47 ppm. Harman (1969) found unionids in water with total alkalinity ranging between 20 and 30 mg/f and stated that sudden changes in pH were probably more detri- mental than low dissolved solids. Tangipahoa River fauna live in water at the extreme lower end of the range of dissolved calcium concentra- tions usually reported for the freshwater Mollusca. A site on the Tangipahoa River in southern Mississippi was identified which supported a dense, fairly diverse mollusk community. Bivalves were found in good quality substrate in water that was clear, well oxygenated but with extremely low dissolved calcium levels. While freshwater mollusks are usually found in medium to hardwater habitats, it appears that reduced dissolved calcium concentrations are not limiting the Tangipahoa River fauna. LITERATURE CITED Clarke. A. H., Jr. and Berg, C. 0, 19.59. The Freshwater Mussels of Central New York, with an Illustrated Key to the Species of Northeastern North America. University Agricultural Experiment Station Memoir 367:1-79. Grantham, B. J. 1969. The Freshwater Pelecypod Fauna of Mississippi, Doctoral Dissertation, University of Southern Mississippi, Hattiesburg, Miss. Harman. W. N. 1969. The Effect of Changing pH on the Unionidae. The Nautilus 83:69-70. Hartfield, P. D. and Cooper, C. M. 1983. Distribution of Corbicula Jluminea the Asiatic Clam, in Mississippi. The Nautilus 97:66-68. Hartfield, P. D. and Ebert, D. 1984. Factors Affecting the Diversity and Abundance of Mussels in Southwest Missis- sippi. Presentation for Mississippi Academy of Sciences, Biloxi, Miss., March 1984. Hartfield, P. D. and Rummel, R. G. 198,5. Freshwater Mussels (Unionidae) of the Big Black River, Mississippi. The Nautilus 99(4):\\6-\l9. Hiiikley, A. A. 1906. Some Shells from Mississippi and Alabama. The Nautilus 20:52-55. Vol. 100(1) January 31, 1986 THE NAUTILUS 23 Joy, J. E. and McCoy, L. E. 1975. Comparison of Shell Dimensions and Viscera Mass Weights in Corbicula manilensis (Philippi, 1844). The Nautilus 89(2):51-54. Sickle, J. B. 1979. Population Dynamics of Corbicula in the Altamaha River, Georgia. In J. C. Britton, ed., Proceed- ings. First International Corbicula Symposium, The Texas Christian University Research Foundation, Ft. Worth, Texas. Stern, E. M. 1976. The Freshwater Mussels (Unionidae) of the Lake Maurepas-Pontchartrain-Borgne Drainage Sys- tem, Louisiana and Mississippi. Ph.D. Thesis, Louisiana State University, Baton Rouge, La. NOTES ON SPECIES OF BROCCHINIA (GASTROPODA: CANCELLARIIDAE) Richard E. Petit' Research Associate Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 ABSTRACT Available replacement names are discussed for Voluta mitraeformis Brocchi, 181Jt, the type species of Brocchinia Jousseaume, 1887, and a primary junior homonym of Voluta mitraeformis Lamarck, 1811. Cancellaria pusilla Adams, 1869 is also preoccupied. This Recent species is redescribed as Brocchinia clenchi, n. sp. Voluta mitraeformis Brocchi, 1814, a well- known cancellariid of the European Tertiary, is the type species of the genus Brocchinia Jousseaume, 1887. Although this taxon is fre- quently cited, its status as a junior primary homonym of Voluta mitraeformis Lamarck, 1811 has been noted in the literature only by Malatesta (1974:376) and DavoH (1982:61). Malatesta (1974:377) considered Cancellaria pusilla H. Adams, 1869 to be available as a replacement name for the preoccupied Voluta mitraeformis Brocchi, although he considered the Recent and Tertiary populations to be sepa- rable. He further stated that the "fossil sub- species" {sotfospecie fossile) had as an available name Cancellaria cerithiopsis Almera & Bofill, 1887 {sic). Malatesta's conclusions are incorrect as C. pusilla H. Adams is not an available name, being a junior primary homonym of Cancellaria pusilla Sowerby, 1832, and C. cerithiopsis 'Mailing address: P.O. Box 30, North Myrtle Beach, SC 29582. Almera & Bofill dates from 1898, not 1887. Earlier synonyms of V. mitraeformis Brocchi exist, as shown below. Davoli (1982:61) stated that Voluta mitrae- formis Brocchi, 1814 and Voluta mitraeformis Lamarck, 1811 should not be regarded as pri- mary homonyms inasmuch as Brocchi pointed out that his species belonged to Lamarck's genus Cancellaria. Davoli referred to the Preamble of the International Code of Zoologi- cal Nomenclature in his plea for retention of Brocchi's V. mitraefoifmis, but the applicable portion of the Code in this instance is Article 60a which states: "A junior homonym must be rejected . . ." (italics added). This is unequivocal. Brocchinia Jousseaume, 1887 Type, by subsequent designation of Sacco, 1894, Brocchi- nia mitraeformis (Brocchi) ( = Voluta mitraeformis Brocchi, 1814 non Voluta mitraeformis Lamarck, 1811; = Brocchinia parvula tauropama Sacco, 1894). Brocchinia has been cited as a subgenus of Narona H. & A. Adams, 1854 by various authors (see synonymy in Malatesta, 1974:377). 24 THE NAUTILUS January 31, 1986 Vol. 100(1) The type of Narona is Cancellaria clavaiula Sowerby, 1832 from the tropical eastern Pacific. There is no evidence for a subgeneric relation- ship between the two genus-level taxa. Narona is characterized by a somewhat tabulate shell having a distinct anterior canal; its columella bears two strong, sharp folds, the posterior one being superior. Brocchinia has a small non- tabulate shell with a rounded anterior; the short siphonal canal evidenced only as a slight curving of the columella; its two columellar folds are weak and rounded. Brocchinia parvula parinila (Beyrich, 1856) Synoyiymy- Cancdlaria parvula Beyrich, 18.56:58, pi. 3, figs. 82a-b. Cancellaria bicarinata Homes & Auinger, 1890:281, pi. 33, figs. 16a-c. Brocchinia mitraefnrmis parvula (Beyrich), .Jansseii, 1983:9, pi. 1, figs.' 5-7; pi. 5, fig. 3, text-fig. 4. Cancellaria parvula Beyrich is apparently the earliest named taxon considered to be a synonym or subspecies of Valuta vfiitraeformis Brocchi, and therefore becomes the nominotypi- cal subspecies of the species group (I. C.Z.N. Article 47). This German Miocene species was cited by Gilbert (1952:130, pi. 9, fig. 16; 1960:4) as a form of C. mitraeformis (Brocchi), his figure agreeing well with Beyrich's original figures. Brocchinia parvula has been treated as a valid species, separable from B. mitraeformis. by Kautsky (1925:137), Sacco (1894:70) and Seiber (1936:93). Davoli (1982:61) mentions C. parvula in his discussion but does not include it in his synonymy of B. mitraeformis. Malatesta (1974:376), in his synonymy, lists Kautsky's cita- tion of B. panmla but does not list the original description, a possible indication that he consid- ered the two to be separable. In the most recent paper treating the subject, Janssen (1983:9, 10) considers B. parvula to be a subspecies of.S. mitraeformis (Brocchi) and states that B. par- vula occurs in the Mediterranean Pliocene as well as in the northern European Miocene. This interpretation of the relationship between B. parvula and B. mitraeformis agrees with this writer's opinion. Brocchinia parvula tauroparva Sacco, 1894 Synonymy - Volula milraiiformis Brocchi, 1814:fi45, pi. 15, fig. 13 (noii Valuta mitraeformis Lamarck, 1811:73). Brocchinia ynitraeformix var. tauroparva Sacco, 1894:68, pi. 3, fig. 82. Narona (Brocchinia) mitraejhi-viis (Brocchi), Davoli, 1982: 61, pi. 7, figs. 3-6. Sacco (1894:68-70) named eight varieties of Brocchinia mitraeformis (Brocchi) from the Italian Pliocene. The first of these, Broc- chinia mitraeformis tauroparva. is not distinct from the typical form, and is here considered to be a subjective synonym of B. mitraeformis (Brocchi). Brocchinia tauroparva (Sacco) thus replaces the preoccupied V. mitraeformis Brocchi. Brocchinia clenchi, n. sp. ' Figs. 1-4 Synonymy - Cancellaria pufiiJla H. Adams, 1869:274, pi. 19, fig. ]2(non Cancellaria puxilla Sowerby. 1832:6, fig. 34). Cancellaria mitraeformis Brocchi, .Jeffreys, 1885:49. Narona (Brocchinia) p^isilla (H. Adams), Nordsieck, 1968: 151, pi. 25, fig. 87.,3(). Df'.srnj9^iOK-Paucispiral nucleus smooth, heliciform, consisting of about IV2 whorls; tran- sition from protoconch to teleoconch almost im- perceptible. Faint peripheral spiral cords arise midway on the first postnuclear whorl. The spiral sculpture consists of 3 to 5 extremely weak cords, with 3 usually visible on the spire whorls, and 5 or so on the body whorl. Axial sculpture, when present, consists only of weak nodes on the periphery that are crossed by the spiral cords. Suture distinct, slightly impressed. Teleoconch of about 4 whorls; nonumbilicate. Aperture ovate with a weak but noticeable col- umellar callus. Columella with two distinct, rounded folds of approximately equal size, the posterior one being slightly larger and more pronounced. Siphonal canal indicated by the angled base of the columella and a minute depression in the base of the aperture. Outer lip prosocline, usually denticulate within. The den- ticles, which do not extend to the edge of the lip, vary in strength and number. Shell white or horn colored. Animal unknown. Type Material: Holotype, 4.5 mm x 2.3 mm (USNM 849002); Paratype, 5.2 mm x 2.6 mm (USNM 189694); Paratype, BM(NH) 1855.4. 4.202, 6.0 mm x 2.5 mm (possibly the type of C. pu.'iilla Adams). Type locality: Josephine Bank, 340-430 fms. Josephine Bank is situated at approximately Vol. 100(1) January 31, 1986 THE NAUTILUS 25 FIGS. 1-4. Brricrhima rlenchi new species. 1, Paratype, BM(NH) 185.S.4.4.202, Oratova, Canary Islands. 15x. 2. Paratype, USNM 189694, Josephine Bank, 621-786 m, 20x. 3, Holotype, USNM 849002, Josephine Bank, 621-786 m, 20 x. 4, Apical view of protoconch of holotype, 65 x. 37°N, 14°W, due West of the southern tip of Portugal. The type and paratype in the USNM are from the Jeffrey's collection. Etymology: This species is respectfully dedi- cated to the memory of the late Dr. William J. Clench. Discussion: This Recent species was previous- ly described as Cancellaria pusilla Adams, 1869, but that name is preoccupied by C. pusilla Sowerby, 1832. Jeffreys (1885:49) placed C. pusilla Adams in the synonymy of C. rnitrae- formis (Brocchi), while Sykes (1911:332), Harmer (1918:396) and Dautzenberg (1927:73) considered the Recent species to be distinct. The Recent B. clenchi is easily separable from the Tertiary species by its much smaller size and by its comparatively weak sculpture. The illus- tration accompanying Adams' description of C. pusilla is misleading, as the fine, almost in- distinct spirals are shown to be quite strong. 26 THE NAUTILUS January 31, 1986 Vol. 100(1) The drawings of Nordsieck (1968:pl. 25, fig. 87.30; 1979:pl. 37, fig. 6) are too poor to be recognizable. Although included by Nordsieck & Talavera (1979:152) with the notation that its occurrence in the Canaries may be accidental, specimens have now been found in fish traps in Tenerife South at a depth of 43 m (Talavera, pers. comm.). As the type of C. piisilla Adams cannot be positively identified, the species has been re- described so that a holotype can be designated, rather than simply proposing a nomen novum. The BM(NH) specimen listed as a paratype is from the R. M. McAndrew collection. Acknowledgments This paper would not have been possible with- out considerable assistance. Dr. Franco Davoli, Universita di Modena, Italy, furnished a transla- tion of his work, and corresponded regarding the desirability of retaining Brocchi's taxon. Mr. Piero Piani, Bologna, Italy, furnished transla- tions, comments and literature. Mrs. Kathie Way, British Museum (Natural History), London, located the possible type of Adams' C. pusilla and made it available for study. The late Dr. Joseph Rosewater, National Museum of Natural History, Washington, D.C., made the Jeffreys specimens available and located pertinent references. Dr. Francisco G. Talavera, Museo Insular de Ciencias, Tenerife, Canary Islands, furnished in- formation and literature. Dr. A. G. Beu, N.Z. Geological Survey, Lower Hutt, New Zealand, read and commented on an early draft of this paper. Dr. M. G. Harasewych, National Museum of Natural History, Washington, D.C. made the photographs, prepared the plate, and reviewed the manuscript. The cooperation and assistance of all is grate- fully acknowledged. LITERATURE CITED Adams. H. 1869. Descriptions of a new genus and fourteen new species of marine shells. Froc. Zool. Soc. Loiidtni. 1869^112-21^, pi. 19. Adams, H. & A. 1853-1858. The genera uf Recent Mollusca. 3 vols. London. Almera, J. & A. Bofill. 1898. Moluscos fosiles recogidos en los terrenos pliocenicos de Cataluna. Bol. Com. Mapa geoL Espana. IV (ser. 2): XII + 223, 14 pis., Madrid. Beyrich, E. 1856. Die Konchylien des norddeutschen Ter- tiargebirges. Zeitschr. d. Deutsch. geol. Ges. 8. Brocchi, G. B. 1814. Conchiologia fossile subappenniria con osservazioni geotogirhe sugli Appenniyii. 2 vols., Stamperia Reale, Milano. Dautzenberg, Ph. 1927. Mollusques provenant des cam- pagnes scientifiques du Prince Ibert I de Monaco dans rOcean Atlantique et le Golfe de Gascogne. Result. Camp. Scient. Pr. Alberta I de Monaco, fasc. LXXII, 400 pp., 9 pis. Davoli, F. 1982. Cancellariidae (Gastropoda). In E. Monta- naro Gallitelli (ed.). Studi monografici sulla malacologia mioceniea modenese. Parte I - I Molluschi tortoniani di Montegibbio. Pobieont. Ital 72 (n.ser. vol. 42);5-73, 7 pis., Pisa. Glibert, M. 1952. Faune malacologique du Miocene de la Belgique. II. Gastropods. Inst. Roy. Sci. Nat. Belg., Mem. 121:1-197, 10 pis. 1960. Gastropodes du Diestien. du Scaldisien et du Merxemien de la Belgique. 4me Note. Inst. Roy. Sci. Nat. Belg.. Bull. 36(33): 1-44, 2 pis. Harmer, F. W. 1918. The Pliocene Mollusca of Great Britain. Part III. Pnheontographical Soc. 70:303-461. pis. 33-44. Horncs, R. & M. Auinger. 1890. Die Gastropoden der Meeres-Ablagerungen der ersten und zweiten Miocanen Mediterranstufe in der ostereich-ungarischen Monarchie. Abhdlg. Geol. Reichsanst. 6. International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomenclature, 3rd Ed. pp. i-xx, 1-338. London. Janssen, A. W. 1983. An account of the Cancellariidae (Gastropoda) of Winterswijk-Miste (Miocene. Hemoorian), The Netherlands. Scripta Geol. 68:1-39. 6 pis. Jeffreys, J. G. 1885. On the Mollusca procured during the 'Lightning' and 'Porcupine' Expeditions, 1868-70. (Part IX). Proc. Zool. Soc. London. 1885:27-63 , pis. 4-6. . Jousseaume, F. P. 1887. La famille des Cancellariidae. Le Natural., ami. 9. 2e ser., pp. 155-157, 163, 192-194, 213-214, 221-223. Kautsky, F. 1925. Das Miocan von Hemmoor und Basbeck- Osten. Abh. preuss. geol. LandesanM.. N.F. 97:1-255, 12 pis. Lamarck, .J.B.P.A. 1811. Suite de la determination des especes de Mollusques testaces. Valuta. Ann. Mus. Nat. d Hist. Nat. 17:54-80. Malatesta, A. 1974. Malacofauna pliocenica umbra. Mem. Carta Geol. It 13:XI-i-498, 32 pis.. Roma. Nordsieck, F. 1968. Die europaischen Meeres-Gehausesck- necken (Prosobranrhia) com Eisnifer bis Kapverden U7id Mittelmeer. 268 p. Gustav Fischer. Stuttgart. Nordsieck, F. & F. G. Talavera. 1979. Moluscos Marinas de Canarias y Madera (Gastropoda). 208 p., 46 pis. Tenerife. Sacco, F. 1894. I mollu.schi dei terreni terziarii del Piemonte e Liguria. Pt. XVI (Cancellariidae). Mem. R. Accad. Sc. Torino, Clausen Ed., 78 pp.. 3 pis.. Torino. Sieber, R. 1936, Die Cancellariidae des niederosterreichis- chen Mioziins. Arrhir. Mollu.fk. 68:65-115. Sowerby, G. B. 1832-33. The Conchological Illustrations. Cancellaria. Pts. 9-13. London. 5 pis. with explanations -I- catalogue, 10 pp. (Pts. 9-12, figs. 1-35, published 1832; Vol. 100 (1) January 31, 1986 THE NAUTILUS 27 pt. 13, figs. 36-44 and catalogue published 1833). Sykes, E. R. 1911. On the mollusca procured during the "Porcupine" Expeditions, 1869-70. Supplemental Notes, Part IV. Proc. Malac. Soc. London 9(6):331-348. ON THE TYPE SPECIES OF METULA H. & A. ADAMS, 1853: BUCCINUM CLATHRATUM A. ADAMS AND REEVE, 1850 (GASTROPODA: BUCCINIDAE) William K. Emerson Department of Invertebrates American Museum of Natural History New York, New York 10024 ABSTRACT The type species o/Metula H. and A. Adams. 1853: Buccinum clathratum A. Adams and Reeve, 1850 (not Kiener, 183J,, nor Anton, 1839) is determined to be referable to Metula amosi Vanatta, 1913, from, the tropical eastern. Pacific. The genera Acamptochetus Cossmann, 1901; Antemetula Rehder, 19J,3; and Colubra- rina Kuroda and Habe, in Kuroda, Habe, and Oyama, 1971, are placed in the synonymy of Metula. The "Metula problem" has long been the con- cern of workers dealing with these buccinid gastropods owing to the uncertain nomencla- tural and taxonomic status of the type species of this genus-group taxon (E. A. Smith, 1904; Woodring, 1928; Tomlin, 1927; Rehder, 1943; Altena, 1949; Knudsen, 1956; Cernohorsky, 1971; Olsson and Bayer, 1972, Kilburn, 1975; and Houbrick, 1984). This note undertakes to solve these questions. The type of Metula, Buccinum. clathratum A. Adams and Reeve (1850, p. 32, pi. 11, fig. 12), was stated to have been dredged off the Cape of Good Hope, in 136 fathoms [248 meters] during the 1843-1846 voyage of the H.M.S. "Samarang". TomHn (1927, p. 160), in his review of the South American mollusks of the "St. George" Expedition, however, noted that this species was: "Originally described from deep water off the Cape, but that locality, like many others in the 'Samarang' work, is certainly erroneous." Furthermore, Tomlin (1927, op. cit.) concluded that this taxon was referable to speci- mens in his collection from Balboa, Panama, which he identified as "Metula clathrata (A. Ad. & Rve.)". Because later workers have largely ig- nored or questioned the identity of Buccinum clathratum A. Adams and Reeve, 1850, the type species oi Metula H. and A. Adams, I examined the holotypic specimen of this taxon, which is in the British Museum (Natural History). The specimen labeled as holotype of Buccinum clathratum. (BM(NH), 1874. 12. 1'l. 145; here illus- trated, figures 1, 2) is 25.3 mm in height. The original polychrome illustration of the type (A. Adams and Reeve, 1850, pi. 11, fig. 12) depicts a specimen of the same size as the holotype, which is now faded and has a chip on the anterior por- tion of the outer lip. As Cernohorsky (1971, p. 149) has pointed out, Bu.cci.num clathratum (A. Adams & Reeve, 1850) is twice preoccupied (not Kiener, 1834, p. 101, nor Anton, 1839, p. 91). Fortunately, a re- placement name is not required, as Metula amosi Vanatta (1913, p. 22, figs. 1, 2; Keen, 1971, p. 566, fig. 1133; Olsson and Bayer, 1972, pp. 906, 907, figs. 1, B-D) is an available junior synonym of this taxon. The most common of the Panamic Metula, this species ranges offshore from the Gulf of California to Panama Bay. Although large specimens attain more than 44 mm in height, mature examples of M. amosi in the American Museum of Natural History (AMNH) collection from Mexico and Panama 28 THE NAUTILUS .lanuarvSl, 1986 Vol. 100 (1) are nearly as diminutive as the lioiotype of M. clathratn. Tomiin's (1927, op. cit.) referral of this species to Pacific Panamanian waters is thus confirmed, and the type locality of M. amosi is here restricted to Balboa, Bahia de Panama. It should be noted that Knudsen (1956, pp. 39, 40, pi. 1, fig. 1) referred a specimen dredged off "Spanish Guinea", West Africa to Metula clatkrata (A. Adams & Reeve, 1850) (not Kiener, 1834, nor Anton, 1839). The specimen figured by Knudsen is 47 mm in height (here reproduced, fig. 3) and is referable to an uniden- tified specimen of Metula in the AMNH collec- tion (#198755; here illustrated, figs. 4, 5) from "West Africa", cx-Marcel Pin collection. Another record of this west African species, taken in 250 meters off Dakar, Senegal (Natal Museum J. 4152) was referred to me by Dr. R. N. Kilburn. These specimens, however, are not conspecific with Metula clathrata (=M. amosi Vanatta) on the basis of the type specimen of A. Adams and Reeve's M. clathrata (cf . figures 1 , 2 with figures 3-5), and they appear to represent a new species. Nor is Knudsen's west African specimen referable to Metula boswellae Kilburn (1975, pp. 594, 595, fig. 10b, 10c), from off "MoQamhique", East Africa. Kilburn (1975, p. 592) accepted Knudsen's (1956, op. cit.) referral of the "Spanish Guinea" specimen to M. clathrata (A. Adams & Reeve, 1850) and he noted that this taxon was preoccupied. Kilburn (1975, op. cit.), therefore, renamed this taxon, Metula knudseni (Kilburn, 1975, p. 592). This unnecessary replacement name thus becomes a junior subjective synonym of Metula amosi Vanatta, 1913. A number of authors have discussed the taxo- nomic status oi Metula H. and A. Adams (1853, p. 84). Rehder (1943, p. 199) appears to be the first to attribute Metula clathrata (A. Adams and Reeve, 1850) as the type species (subse- quent selection of Kobelt, 1876, p. 29, pi. 6, fig. 11 [ = "12"]). Woodring (1928, p. 286; 1964, p. 259; Keen, 1971, p. 566; Olsson and Bayer, 1972, p. 902) considered Buccinum m.etula Hinds, 1844, to be the type species by "hidden tautonymy," for which there is no provision in the ICZN Code. Thus, this designation must be rejected (Cernohorsky, 1971, p. 149). Kobelt's (1876, op. cit.) selection oi Metula clathrata, as the type species, therefore, appears to be the first available designation for Metula. Cernohorsky (1971, pp. 151, 152) and Houbrick (1984, p. 420) considered Acampto- chetus Cossmann (1901, p. 123) diwd Antemetula Rehder (1943, p. 199) to be congeneric taxa. The type species of Acamptochetus, by original designation, is Murex mitraeformis Brocchi, 1814, from the Neogene of Italy and is a typical Metula (Cernohorsky, 1971, fig. 46). Other Neogene fossils are known from Europe, Java, Sumatra, India and the Americas (Altena, 1949; FIGS. 1-5. 1 and 2, Metula amosi Vanatta, holotype of Buccinum clathrnia A. Adams and Reeve, BM(NII) 1874. lli. I 1 . 1 4;'.. x2. 3, "Metula clathrata" after Knudsen (19.56, pi. 1, fig. 1) from "Spanish Guinea", not Metula clathrata (A. Adams and Reeve), x IV2. 4 and 5, Metula sp., AiVINH 1987.5.5, ex-Marcel Pin coll., from "West Africa", x 1 V2. Vol. 100(1) January 31, 1986 THE NAUTILUS 29 Olsson and Bayer, 1972; Malatesta, 1974; and Grecchi, 1978). The type species oi Antemetula by original designation is Buccinum metula Hinds, 1844. This species was originally reported from the "West coast of Veragiia [Republica de Panama] . . . from a depth of a few fathoms, among mud." This locality, however, is apparently an error, as no additional specimens have been recognized in tropical American waters. The type specimen (Hinds, 1844, pi. 16, figs. 13, 14; refigured by Keen, 1971, p. 567, fig. 1134 and by Olsson and Bayer, 1972, fig. lA) is not in the British Museum (Natural History) teste Kathie Way, 1985). Furthermore, this specimen could not be located there some 35 years ago (Altena, 1949, p. 385). Many years ago Smith (1904, p. 465) noted a close resemblance of young specimens of Metula mitreUa (A. Adams and Reeve, 1850, p. 32, pi. 11, fig. 13, from the "China Sea; . . . ten fathoms"), to the illustration of the type specimen of M. metula. Smith (1904, op. cit.) concluded that Buccinum 'metula was conspecific with B. mitrella. a well- known Indo-West Pacific species, and that the original west American locality was probably erroneous. I agree with his conclusions {cf. the il- lustration of Cernohorsky (1971, p. 150, fig. 47) with that of Olsson and Bayer (1972, p. 905, fig. lA). If this interpretation is correct, M. mitrella (A. Adams and Reeve, 1850) is a junior subjec- tive synonym of M. metula (Hinds, 1844) and this taxon should be removed from the eastern Pacific faunal list. Kilburn (1975, p. 592) assigned the genus- group name, Colubrarina Kuroda & Habe, in Kuroda, Habe & Oyama (1971, p. 173) to the synonymy of Metula. The type species of this taxon by original designation is Antemetula (Colubrarina) metulina Kuroda & Habe, in Kuroda, Habe & Oyama, 1971 {op. cit.. p. 173, pi. 46, fig. 9), from Honshu and Shikoku, Japan, in 90-200 m. Described as a subgenus oiAnteyne- tula Rehder, the type species of Colubrarina was said to differ from the nominate subgenus by having a larger and more coarsely sculptured shell. In summary, the buccinid genus Metula H. and A. Adams, 1853, with the type species: M. clathrata (A. Adams and Reeve, 1850) [ = M. amosi Vanatta, 1913 (synonym; M. knu(lse7ii Kilburn, 1975)], is available for an assemblage of Neogene species from the Old and New World tropical belt and for several Recent species sur- viving in the tropics. Acamptochetus Cossmann, 1901 (type species; M. mitraeformis (Brocchi, 1814), Antemetula Rehder, 1943 (type species; M. metula Hinds, 1844) and Colubrarina Kuroda and Habe, 1971 (type species: M. metulina Kuroda and Habe, 1971) are junior synonyms. Acknowledgments I am very much indebted to Ms. K. M. Way of the Department of Zoology, British Museum (Natural History) for the loan of the holotype of Buccinum clathratum. Dr. Richard S. Houbrick of the National Museum of Natural History, Smithsonian Institution kindly read a draft of the manuscript and allowed me access to the United States National Museum malacological collections. I am indebted to Dr. Richard H. Kilburn of the Natal Museum for reading the manuscript and lending me a specimen critical to this study. I thank Dr. Philippe Bouchet of the Museum National d'Histoire Naturelle, Paris for his comments on the "Metula problem". My AMNH colleagues kindly provided valu- able contributions as follows; Mr. Walter E. Sage, HI, technical assistance, Mr. Peter J. Harries, photography, and Mrs. Stephanie Grooms, word processing. I also wish to join others in congratulating The Nautilus on its 100th anniversary of its found- ing and to express my particular appreciation of having known and benefited from the friend- ship of three past Consulting Editors- Karl Jacobson, Joe Rosewater and Bill Clench. LITERATURE CITED Adams. H. and A. 18.5.3 [-1854]. The Genera of Recent Mollusca, Arranged According to Their Organization London l(1.3):65-96, pis. 9-12 (June, 1853). Adams, A. and L. Reeve. 1850. Mollusca, pt. 2:25-44, pis. 10-17. In. The Zoology of the Voyage of H.M.S. "Samarang". Under the Command of Captain Sir Edward Belcher During the Years 1843-1846. London, 1848-1850. Altena, C. 0. van Regteren. 1949. The genus. .Antemetula Rehder in the Indo-West Pacific area, with a description of two new fossil species. Bijdragen Tot Dierkunde 28: 385-393. 7 figs. Anton. H. E. 1839. Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard Anton befinden. Halle, xvi -i- 110 pp. Cernohorsky, W. O. 1971. Indo- Pacific Pisaniinae (Mollusca: Gastropoda) and related buccinid genera. Rec. Auckland 30 THE NAUTILUS January 31, 1986 Vol. 100(1) Inst, and Mus. 8:131-167, 95 figs. Cossmann, M. 1901. Essais de Paleochologie Comparee, pt. 4, 293 pp., 10 pis. Grecchi, G. 1978. Problems connected with the recorded occurrence of some mollusks of Indo-Pacific affinity in the PHocene of the Mediterranean area. Riv. Ital. Paleont. 84(3):797-812, pi. 82, 8 figs. Hinds. R. B. 1844 [-1845], Mollusca, pt. 2, In The Zoology of the Voyage of H.M.S. Sulphur, under the command of Capt. Sir Edward Belcher . . . during 1836-1842. London, pp. 25-48. pis. 8-14 (Oct. 1844). Houbrick, R. S. 1984. A new "Metula" species from the Indo-West Pacific (Prosobranchia: Buccinidae). Proc. Biol. Soc, Wash. 97(2):420-424, 1 fig. Keen, A. M. 1971. Sea Shells of Tropical West America. Second Ed., Stanford Univ. Press, viii -^ 1064 pp., illus. Kiener, L. C. 1834 [-1841]. Genre Buccin. In. Species general et inconographie des coquilles vivantes . . . Paris, vol. 9, 112 pp., 31 pis. Kilburn, R. N. 1975. Taxonomic notes on South African marine Mollusca (5): including descriptions of new taxa of Rissoidae, Cerithiidae, Tonnidae, Cassididae, Buccinidae, Fasciolariidae, Turbinellidae, Turridae, Architectoni- cidae, Epitoniidae, Limidae, Thraciidae. Ann. Natal Mus. 22(2):577-622, 25 figs. Knudsen, J. 1956. Marine prosobranchs of tropical West Africa (Stenoglossa). Atlantide Report No. 4:7-110, pis. 1-4. Kobelt, W. 1876 [-1881]. Illustriertes Conchylienbuch, Nurnberg, vol. 1, pp. 1-143, 50 plates. Kuroda, T., T. Habe and K. Oyama. 1971. The Seashells of Sagami Bay. Tokyo, 1-485 + 55 pp. (English Text), pi. 1-22. Malatesta, A. 1974. Malacofauna Pliocenia Umbra. Mem. Carta Geol. Italia. Rome. vol. 13. 499 pp., 32 pis. Olsson, A. A. and F. M. Bayer. 1972. American metulas (Gastropoda: Buccinidae). Bull Mar. Sci. 22(4):900-925, 14 figs. Rehder, H. A. 1943. New marine mollusks from the Antil- lean region. Proc. U. S. Nat. Mus. 93(3161):187-203, pis. 19, 20. Smith, E. A. 1904. On Mollusca from the Bay of Bengal and the Arabian Sea. In. Series III. no. 1. Natural History Notes from H. M. Indian Survey Steamer "Investigator", Commander T. H. Hemming, R. N. Ann. Mag. Nat. Hist. 7(13):453-473. Tomlin, J. R. le B. 1927. The Mollusca of the "St. George" Expedition. (I) The Pacific coast of S. America. Jour. Conchology 18(6): 153- 170. Vanatta, E. G. 1913. Descriptions of new species of marine shells. Proc. Acad. Nat. Sci. Philadelphia 65(l):22-27, 3 text figs., pi. 2. Woodring, W. P. 1928. Marine mollusks from Bowden, Jamaica. Pt. 2. Gastropods and discussion of results. Carnegie Inst. Washington, Pub. no. 385, vii + 564, pis. 1-4, 3 figs. 1964. Geology and Paleontology of Canal Zone and adjoining parts of Panama. Description of Tertiary mollusks (Gastropods: Columbellidae to Volutidae). U.S. Geol. Surv. Prof. Pap. 306-C:241-297, pis. 39-47. SOME ADDITIONAL TAXONOMIC UNITS THAT FIRST APPEAR IN PUBLICATIONS BY J. G. COOPER Eugene Coan Research Associate Department of Invertebrate Zoology California Academy of Sciences San Francisco, CA 94118 ABSTRACT Seventeen taxonomic units that first appeared in publications by James Graham Cooper and that were not given in my earlier table of his taxa are listed here. All but three are nomina nuda. It is said that the more complete a list is, the more glaring are its omissions. So it is, perhaps, with my listing of the taxonomic units that first appeared in the publications of James Graham Cooper (Coan, 1982:148-185). Seventeen names that should have been in that list have come to my attention since my book was published. Most of them were well hidden, being credited to other workers. Fourteen of these names are nomina nuda, only three actually being validated by Cooper. Two replace previous homonyms. The third is a synonymous generic unit. It is not impossible that other similarly hidden taxa will come to light in the future. Vol. lOU(l) January 31, 1986 THE NAUTILUS 31 In the listing of mollusks below, I follow the format used in my previous table. The Cooper papers cited refer to the Literature Cited of this article, but I also provide, in brackets, references keyed to the complete bibliography of Cooper's papers given in my book (Coan, 1982:223-281). Class Bivalvia heringi. Thracia - Cooper, 1894 [18;)4b|: Suppl., 12th sheet, ex Dall MS [nomen nuduvi]. Later validated by Dall, 1915: 442. compacta. Sanguinolaria - Cooper, 1894 [1894b]: 126th sheet [nowen nudum]. Not subsequently validated. romijlectens. Tellina - Cooper, 1894 [1894b]: Add. & Corr., 4th sheet, ex Carpenter MS [nomev nudum]. Not sub- sequently validated. Philohrya - Cooper, 1867 [1867b]: 12, ex Carpenter MS. Type Species - Bryophila .-ietosii Carpenter, 1864a: 314, by monotypy. Remarks - Bi-yophila Carpenter, 1864a:314, proved to be a homonym (reon Treitschke, 1825;.57), and Carpenter him- self later renamed it as P/ii7obn/a (Carpenter, 1872: index, p. 21). By then, however, it was too late; Cooper had already validated the name. Cooper did not expressly propose Philohrya as a replacement name and it must therefore be taken as a new genus. Class Gastropoda bimaculata Clypidella - Cooper, 1867 11867b]: 24, ex Dall MS ]noinen nudum]. Later validated by Dall, 1871:132; 160; pit. 15, fig. 7), as Fissurellidaea bimaculata. callomarginata. Clypidella - Cooper, 1867 [1867b]: 24, ex Carpenter MS ]nomen nudum]. Later validated by Dall, 1871:133; 160; pit. 15, fig. 8. clathftita. Ocinebra interfossa "var." - Cooper, 1870 [1870f]:69, ex Carpenter MS [nomen nudum]. Later vali- dated by Dall, 1919:334, as Tritonalia interjmsa clathrata. compactum. Buccinum - Cooper, 1894 ]1894b]: 15th sheet, ex Dall MS [noinen nudum]. Not subsequently validated. gabbiana. Chemnitzia - Cooper, 1867 [1867b]: 34. New name for Turbonilla graciUiyna Gabb, 1865:186, non Chemnitzia gracillima Carpenter, 1857:431. hai'pa. Tomatina - Cooper, 1870 [1870tl: 56, ex Dall MS ]nomen nudum]. Later validated by Dall, 1871:136-137; 160: pit. 15, fig. 11. Leptothyra Cooper, 1867 [1867b]: 25. Type Species - "L. sanguinea Cpr.," = Turbo sanguinewi Linnaeus, 1758: 763 (subsequent designation herein). Remarks - A synonym of Homalapoma Carpenter, 1864b:537, 588, 627, 652, which has the same type species. Previous workers have evidently missed Cooper's validation of Leptothyra. dating it instead from Dall, 1871:130. It thus becomes a senior homonym of Leptothyra Pease, 1869:70, and if workers wish to save this name, a petiton would have to be filed with the International Commission on Zoological Nomenclature. muricata, Ocinehra interfossa "var." - Cooper, 1870 [1870f]: 69, ex Carpenter MS ]nomi-u riuduin]. Not sub- sequently validated. paueieostata. Leptothyra - Cooper, 1867 [1867b]: 25, ex Dall MS [nomen nudum]. Later validated by Dall, 1871: 131; 160; pit. 15, fig. 10. sitkensis, Odostomia - Cooper, 1894 [1894b]: 100th sheet, ex Dall MS [nomen nudum]. Not subsequently validated. steamsiana. La.mel.la.rin - Cooper, 1870 [1870f]: 67, ex Dall MS [nomen nudum]. Later validated by Dall, 1871: 122; 160; pit. 15, fig. 6, but as L. utearnsii. Class Polyplacophora fimbriatus, Callochilun - Cooper, 1867 [lS67b]: 23, ex. Carpenter MS [ncmwn nudum]. Later validated by Keep, 1887:112, as Callistochiton fimbriatus. latior. Planphorella - Cooper, 1894 [1894b]: 113th sheet, ex Carpenter MS [nomen nudum]. Not subsequently validated. LITERATURE CITED Carpenter, Philip Pearsall. 1857. Catalogue of the collection of Mazatlan shells, in the British Museum: collected by Frederick Reigen . . . London (British Museum) i-iv + ix- xvi 4- 552 pp. (1 Aug. 1857) [Warrington ed., publ. simultaneously] [reprinted, Paleo. Resh. Inst., 1967]. 1864a. Diagnoses of new forms of mollusks col- lected at Cape St. Lucas by Mr. Xantus. Ann. Mag. Natur. Hist. (3) 13(76):31 1-315 (April); (78):474-479 (June); 14(79):4.5-49 (.July) [reprinted in Carpenter, 1872: 207-221]. 1864b. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Brit. Assn. .Adv. Sci.. Rept. 33[for 1863]:517-686 (post-1 Aug. 1864) [reprinted in Carpenter, 1872:1-172]. 1872. The mollusks of western North America. Embracing the second report made to the British Asso- ciation on this subject, with other papers: reprinted by permission, with a general index. Smithsonian Inst. Misc. Colin. 10(252):xii + 325 + 13-121 pp. (Dec. 1872). Coan, Eugene V. 1982. James Graham Cooper; pioneer western naturalist. Moscow, Idaho (Univ. Press of Idaho) 255 pp.; 31 figs. (22 Feb. 1982). Cooper, James Graham. 1867 [1867b in Coan, 1982]. Geo- graphical catalogue of the Mollusca found west of the Rocky Mountains, between latitudes 33° and 49° north. San Francisco, Calif. (Geol. Survey of Calif.) 40 pp. (post- April 1867). 1870 [1870f in Coan, 1982]. Notes on the Mol- lusca of Monterey, California. Amer. Journ. Conch. 6(1): 42-70 (7 July 1870). 1894 [1894b in Coan, 1982]. Catalogue of west North American and many foreign shells, with their geo- graphical ranges. For labels, exchange, and check lists. With a supplement. Sacramento, Calif. (Calif. State Mining Bureau) 181 unnumbered pages grouped in stapled sections [see Coan, 1982, for details], main body: Title page + Preface -i- 16(1 pp.; Siippl.: Title page + 13 pp.; Add. & Corr.: 5 pp. (April 1894). Dali, William Healey. 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and 32 THE NAUTILllS January 31, 1986 Vol. ]()0(1) the North Pacific Ocean, with notes on others already described. Amer. Journ. Conch. 7(2):9,3-160; pits. 13-16 (2 Nov. 1871). 1915. A review of some bivalve shells of the group Anatinacea from the west coast of America. United States Nation. Mus., Proc. 49(21 16):441-4.56 (27 Nov. 1915). 1919. Descriptions of new species of Mollusca from the North Pacific Ocean in the collection of the United States National Museum. United States Nation. Mus., Proc. 56(2295):293-371 (30 Aug. 1919). Gabb, William More. 1865. Descriptions of new species of marine shells from the coast of California. Calif. Acad. Sci., Proc. (1) 3(3):182-190 (.Ian. 186.'3). Keep, .Josiah. 1887. West coast shells. A familiar descrip- tion of the marine, fresh water, and land mollusks of the United States, found west of the Rocky Mountains. San Francisco (Bancroft Bros) 230 pp.; 182 figs.; frontis. (post-July 1887). Linnaeus, Carolus. 1758. Systema naturae per regna tria naturae . . . editio decima, reformata 1 [Regnum animale], Stockholm (Laurentii Salvii). 824 + iii pp. Pease, William Harper. 1869. Descriptions of new species of marine gasteropodae inhabiting Polynesia. Amer. J. Conch. 5(2):64-79 (7 Oct. 1869). Treitschke, Friedrich. 1825-1826. Die Schmetterlinge von Europa 5. Leipzig (G. Fleischer) [pts. 1, 2 - 1825; pt. 3 - 1826). NOTES ON THE HABITS AND ANATOMY OF THE INTRODUCED LAND SNAILS, RUMINA AND LAMELLAXIS (SUBULINIDAE) Dee S. Dundee^ Department of Biological Sciences University of New Orleans New Orleans, LA 70148 ABSTRACT Marking experiments showed that Rumina decollata moved no more than 25 inches from points of release in a six-month period, demon.'it rating, as with most introduced forms, that passive distribution prevails. Egg-to-egg span was slightly over a year in the lab. While normally considered carnivorous, R. decollata ivill feed on a variety of vegetation, and were routinely fed lettuce. Average growth is l.J,6 mm, in width/ week and average whorl addition is 1.18 whorls/week. At 10 mm high, the snails lose the upper 3. 0-3.. 5 whorls and fill in the top with a .' 9 _ ^ . i ue. 9 - - t • - 4 ■ - . . ( .... ^i:;j^;iJ 1 8 1 ' ■ ' . flit 7- ;j- -':'-'[ ; K - - '.^ ^:: -i » t4 ■ ■ -1 ^ « « ' ' -jt?^ - -.<»E ! ' 1 + 4. ^ *- TI I--. ^ .:^..:.|Ti 41 ^ ""' 1 J: : 1 ^ _, 9 . ^- ^- _j.^j_4- --.-, , — 1 n- -+- -1— J— -AU- i|ilij^ ' ''^' ^ ^ F ^ UI^ ^ WHO R L S FIG. 1. Rumina ikcollnta. This graph shows in terms of height and number of whorls. At the 10 mm high point, the snails become decollated thus having fewer whorls but con- tinuing growth in height. Open circles represent non- decollate individuals; solid dots represent decollate ones. 34 THE NAUTILUS January 31, 1986 Vol. 100(1) examines a 9-mm-long, non-decollate specimen (Fig. 2), one notices a peculiar-appearing apex area. The first 3.0-3.5 whorls have a smooth, glassy appearance, whereas the remaining whorls have some striations. The second major whorl below the protoconch is inflated thus giv- ing the top of the shell a ))uffy appearance. The shell decollates just l)eneath the whorl below the inflated one. I have been unable to observe how the breakage occurs although it has been re- ported that the snail bangs its upper whorls violently against a hard object to get rid of them (Cooke, 189,5). Batts (1957) reported observing them striking the shell against rocks or the aquarium wall to aid in decollation. A calcareous septum, resembling the protoconch, is formed as a cover for the top of the broken shell (Fig. 3). It is easily seen in the decollate forms. If one breaks off the top whorls of a 9.5-mm-long speci- men, one discovers that the top 3.0-3.5 whorls break much easier than those below and that there is no septum present; it is apparently secreted after decollation occurs. Figure 3 is a view of a septum as one looks down on the top of a decollate shell. Notice that the umbilicus is retained. Decollation occurs in widely separated taxa from pulmonates to prosobranchs. It occurs in terrestrial as well as aquatic forms (brackish water). It seems to have evolved independently several times. Examination of a serial section of a 9-mm-long R. decollata reveals nothing unusual about the internal anatomy in the area which is destined to break off. One can only speculate at this point on the selective advan- tage of casting off part of the shell. Perhaps the upper whorls serve as a storage for toxins dur- ing early growth; perhaps decollation is a behavorial matter related to habitat (e.g., ability to get under shelter); perhaps it is only a non- selective mutant. The entire matter needs fur- ther study. Longevity. The longest I have l)een able to keep this species alive in the lab is just over a year. Rascop (1960) reports a longevity record of 12 years but does not cite a reference. That seems a very long time considering longevities of various other snails (Hyman, 1967). Years of observations by the author have shown that mollusks tend to reproduce just prior to the ex- tinction of the colony. This colony, which was followed from egg-to-egg, began reproducing at ten months which was approximately two months prior to the death of the last colony member. Observations on Lamellaxis gracilis Shell. Figure 4 shows a typical adult shell. The maximum size seen in the New Orleans area F'UiS. Z and 3. Riiniiiid di'mUata. 2, A spi'cmifn of ;i|j|jnix- irriatoly 9 mm showing the smooth quahly of the top three whorls as opposed to the striations on the remaining ones. 3, A view into the top of a large decollate specimen. Vol. 100(1) January 31, 1986 THE NAUTILUS FIGS. 4-7. Lamelhxif: gracili.'i. 4, Adult shell. 5, An about-to-hatch snail dissected from its egg case. 6, The jaw, 7, Central and lateral teeth. was 13 mm in length, 3.5 mm in width at the broadest point, and with 9.25 whorls. The maxi- mum size of Lamellaxis micra (Orbigny, 1835) which also has been introduced to this area was, by contrast, 7.2 mm x 2.2 mm, and had 6.75 whorls. The color of L. gracilis is light tan and all whorls except the protoconch are finely striated. A small umltilicus is partially hidden by a minor reflection of the lip (Fig. 4). Unhatched ova (Fig. 5) removed from the female system show clearly the beginnings of striation except on the protoconch. Jaw. This structure is found at the entrance to the pharynx in a mid-dorsal position. As can be seen in Figure 6, it is a single, curved device which has coarse transverse ribbing. Radula. The radula (Fig. 7) is relatively nar- row with little variation among central, lateral, and marginal teeth. The teeth are reminiscent of marginals of Helix aspersa except the lateral cusps are more pronounced. Digestive system. The esophagiis, lined by tall, ciliated columnar epithelium and underlain by a thin circular muscle layer surrounding an equally thin longitudinal layer. The esophagus leaves the pharyngeal area, passes through the nerve ring and posteriorly into the stomach; (Fig. 9 ST). In keeping with other Stylommato- phora, no crop was found. The stomach is lined by tall, columnar, ciliated epithelium underlain by a double muscle layer (longitudinal and cir- cular) and surrounded by connective tissue. The lining is folded in several areas. The digestive gland (Fig. 8, D) empties into the stomach via two ducts -one from each section of gland. A dissection showing the two parts is very difficult and can only be done with fresh (non-preserved) material. Histological observations reveal two cell types making up the digestive gland (also called liver and hepatopancreas). A tall colum- nar cell type which contains numerous vacuoles is often seen with the lumen in a state of disinte- gration implying secretion. The second type is a low, non-ciliated triangular form. Carriker and Bilstad (1946) reported similar cells and called them "digestive" and "calciferous" and deter- mined that the latter contain calcium phosphate granules. Digestion is apparently both intra- and extra-cellular. Sections reveal partially digested material in the stomach lumen (Fig. 8, ST) and also in some the cells of the digestive gland. An intestine leaves the stomach, loops around in the same whorl as the stomach, descends to the right dorso-lateral of the head- 36 THE NAUTILUS January 31, 1986 Vol. 100(1) FIG. 8. Lamt'lliiji.^ ^innms. Parasagittal section of a young snail. D = digestive gland; ST= stomach; I = intestine; A = albumen gland. foot where the anus is located. The intestine (Fig. 8, I) is lined by tall, ciliated columnar epithelium containing mucous cells and under- lain by an inner longitudinal and an outer cir- cular muscle layer. Nervous system. The ganglia form a ring around the anterior portion of the digestive tract just posterior to the pharynx. This is the result of detorsion wherein the parietal (in- testinal) and visceral ganglia migrate forward near the pleural ganglia. In Lamellaxis the end result is two cerebral ganglia on the dorsal of the ring and two pleurals, two parietals, and a visceral (two fused ganglia) forming the ventral portion of the ring and innervating the entire visceral mass. The total nervous system was not traced. A pair of eyes, located on the dorsal tips of the posterior tentacles, are typical pulmonate vesi- cular types (Hyman, 1967, p. 584). The stato- cysts, located adjacent to the pedal ganglia in the foot, are small spherical vesicles each con- taining calcareous granules. Each is covered by a layer of connective tissue and lined internally by a squamous-type epithelium with a layer of cilia equally spaced around the inner periphery. Excretory system. A single nephridium is located near the dorso-posterior of the pulmon- ary cavity. If one holds the snail so that the aperture is facing him, the reno-pericardial mass is on the extreme lower right of the whorl above the aperture. The pericardial cavity has a duct from it into the sac-like nephridium. That nephridium, in most Pulmonates, opens into the |.)ulmonary cavity directly via a slit or a short ureter. I could not determine which is present in this snail either by dissection or sections. Reproductive. System. Lamellaxis is herma- phroditic. The ovotestis, a grapelike cluster embedded in the digestive gland proximal to the columella, contains both oocytes near the peri- phery of each follicle and spermatocytes and sperm more centrally located near the duct of each. A coiled hermaphroditic duct, lined by low ciliated cuboidal epithelium, drains both the ovary and testis. At the lower end of that duct a small saclike structure, the seminal vesicle, ap- pears to branch off and serves as a sperm stor- age area. It is seen full of sperm during the breeding season. Near that point, the male and female systems part company with each system continuing downward, closely applied to and paralleling each other (spermoviduct of some authors) through the whorls towards the anterior end. The male system consists of a sperm duct (vas deferens) with a ciliated cuboidal lining. It receives a prostatic secretion via a small connec- tion between the prostate and vas deferens. The prostate consists of cuboidal cells around an ir- regular lumen. The cells closest to the lumen disintegrate as a result of secretion. Nuclei are basal. The vas deferens continues downward and enters the verge. Just prior to that entry the duct liecomes lined by tall, ciliated glandular epithelium. The system empties on the right side of the head near the base of the right tentacle. The verge is simply a long attenuated tubule with no secondary structures on it. At the point where the systems become separate the oviduct appears to enter a large albumen gland but, actually, the gland is large enough that it has the oviduct "embedded in" it (Fig. 9, A). The allmmen gland has a varied histological struc- ture ranging from cuboidal to low, ciliated, Vol. 1(H)(1) January 31, 1986 THE NAUTILUS 37 epithelial cells. This gland, in section, always ap- pears striated and somewhat "crumbled". The lumen ends of the cells appear to disintegrate as secretions are formed. Just ventral to this albumen gland, at the separation of the male-female systems is an area where fertilization occurs (fertilization pouch or carrefour in other pulmonates: e.g., (Petrellis and Dundee, 1969). A distinct pouch could not be located in L. gracilis. Only a slightly swollen area in the hermaphroditic duct exists. It is dif- ferent histologically from the initial part of the duct in that the epithelium is ciliated columnar in contrast to low, non-ciliated cuboidal in initial portion of the duct. The change in histological structure indicates that the area is the equiva- lent of the fertilization pouch and is doubtlessly the site of fertilization. Fertilized eggs receive their albumen coat as they pass by the albumen gland. As they con- tinue anteriorly downward, the shell is added by the mucus gland (also called capsule gland) which is a continuation of the large glandular mass. The albumen and mucus glands can be dis- tinguished microscopically by change from whit- ish, coarse appearance (albumen) to a smooth yellowish one with the entire mass convoluted (mucus). Microscopically the mucus gland dif- fers from the albumen gland described above by being a highly glandular mass lined by cuboidal cells which, in many cases, are secreting as evidenced by the disintegrated inner cell mass. Non-secretory cells are ciliated, and cell boun- daries are indistinct. Often as many as 6-8 large ova can be seen lined up in the oviduct in the body whorl and the one immediately above it. Just at the point where ova emerge from near the mucus gland, the vagina receives a duct from the spermatheca (also called seminal receptacle or bursa), a sac which stores sperm from copulation. Histologi- cally the spermatheca is composed of non- ciliated, tall columnar epithelium encapsulated in a muscle-connective tissue layer. The vagina is a muscular tubule lined by ciliated epithelium. It ends at the gonopore which is found in the right dorso-lateral area of the head. LITERATURE CITED Batts, Jeanne H. 1957. Anatomy and life cycle of the .'^nail Rumina decollata. Southwest Nat. 2:74-82. Biswas, S. K.. R. Rahman and T. R. Mitra. 1976. Observa- tions on the breeding habits of Opeas gracile (Hutton) (Gastropoda: SubuMnidae). .Jour. Conch. 29:69-70. Carriker, Melbourne and N. M, Bilstad. 1946. Histology of the ahmentary system of the snail, Lymnnen stagnalis (ippressa. Trans. Amer. Micr. Soc. 64:250-275. Cooke, A. H. 1895. In Cambridge Natural History Vol. HI (Molluscs and Brachiopods). Macmiilan & Co. (Reprint edition: 1-ix + 535; 334 figs.; 1 map. Wesley, Ltd., England, 1959). Dundee, Dee S. 1970. Introcr (Linnaeus, 1758) living in the Mediterranean, Init that species lacks nodes and enlarged scales and dif- fers in hinge details, as discussed above in the comparison of genera. None of the extant or extinct species of Nodipectcn have as closely spaced commarginal lamellae, and they have much more massively developed dorsal and intermediate hinge teeth. Ecdiogy: The habitat of the new species is known only from the data provided by Taiwanese fishermen, who said that the shells were trawled off Somalia at depths of 150 to .'500 m. Because they have not divulged the exact locality, it is not known whether the specimens are from the Gulf of Aden or the Indian Ocean side of Somalia. Mr. T. C. Lan of Taipei, Taiwan, who obtained the specimens from the Taiwanese fishermen, has provided a list of associated species, some of which would appear to shed light on the locality of the pectinids. Strombu>! otdi Emerson, 1965, was originally described from the Indian Ocean side of Somalia in the vicinity of Obhia and Mogadiscio (Emer- son, 1965), and recently it also has been found off Oman in the Arabian Sea (David Hargreave, /)cr.s-. com.. 1986). It is not known to occur in the Red Sea or the Gulf of Aden. Cypraea hroderipi Soweri)y, 1832, Cypraea marginalis Dillwyn, 1SI7, and Vanum Iruncafum (Sowerby, 1892) are known mainly from the Indian Ocean and not the Red Sea (Abbott and Dance, 1982), and Mi))>ulio[i. I'mr. Malacol. Sor. London 18:lfir,-2()9. pis. 11-13. Eames, F. E., and L. R. Cox. 1950. Some Tertiary Peclina- cea from East Africa, Persia, and the Mediterranean Region. Pror. Malacol. Sor. London 32:1-68, 20 pis. Emerson, W. K. 1965. Strombun (Tricornis) oldi New Species. Indo-Pncifir MoUuxm l(6):.397-398, pi. 294. Criiffydd, LI. D. 1981. Observations on the Rate of Produc- tion of External Ridges on the Shell o{ Pecten marimi/a in the Laboratory. Jour. Mnrinf Biol. Aax. U.K. 61:401-4 1 1 . Helm. N. E.. and R. E. Malouf. 1983. Rate of Production of External Ridges in the Bay Scallop, .4 n/opcc^'H irradtmix. {.■\bstract) Amer. Zoologist 23(4):1024. Jones, D. S. 1981. Repeating Layers in the Molluscan Shell Are Not Always Periodic. Jour. Pateo. 55(5):1076-1082. Matsukuma, Akihiko. 1978. Fossil Boreholes Made by Shell- Boring Predators or Commensals. I. Boreholes of Capulid Gastropods. Jupnnexe Jour. Molncol. lVpnu.-<) 37(l):29-45. Ohno, Terufumi, 1985. Experimentelle Analysen zur Rhythmik des Schalenwachstums einiger Bivalven und ihre Palaobiologische Bedeutung. (Experimental Analyses Concerning the Rhythm of Shell Growth in some r^ivalves and its Paleobiological Implications). Pnldconlo- unipliira. Abt. A, 289:63-123, 7 pis. Orr, Virginia. 1962. The Drilling Habit or ('apulux daniM (Crosse) (Mollusea: Gastropoda). The Vrliger 5(2):63-67, pi. 7. Waller, T. R. 1969. The Evolution of the Argopi'rtcn gibbux Stock (Mollusea: Bivalvia). with Emphasis on the Tertiary and Quaternary Species of Eastern North America. Polf(„il. Sor. Mfw. 3 (Jour. Palco.. 43(5), suppl.), 125 p., 7 pis. 1972. The Pectinidae (Mollusea: Bivalvia) of Eniwetok Atoll, Marshall Islands. The VcIkjct 14(3): 221-264, 8 pis. 1978. Morphology, Morphoclines and a New Classification of the Pteriomorphia (Mollusea: Bivalvia). Phil. Trans. Royal Soc. London. B, 284:345-365. _. 1984. The Ctenolium of Scallop Shells: Func- tional Morphology and Evolution of a Key Family-level Character in the Pectinacea (Mollusea: Bivalvia). Malarolo ijia 25(1):203-219. A NEW SPECIES OF TRITONIA (NUDIBRANCHIA) FROM SOUTHERN CALIFORNIA AND BAJA CALIFORNIA Hans Bertsch' Biological Sciences, National University, Los Angeles, CA and Antonio Mozqueira Osuna Escuela Superior de Acuacultura, Centro de Estudios Superiores del Estado de Sonora (CESUES), Navojoa, Sonora, Mexico ABSTRACT Tritonia myrakeenae is descrihed as a new species from the Califomian marine famial province and is ilisfinguished anatomically froni the north Pacific and north Atlantic species q/" Tritonia. Although the opisthohranch gastropod fauna of the northeastern Pacific has been mono- graphed extensively by numerous authors (e.g., recently by Marcus, 19(il a; MacFarland, 19(i(v, Keen, 1971; McDonald, 1983; et at.), there are still numerous new distributional records (e.g., Bertsch, 1981; Behrens, 1982; Gosliner & Millen, 1984) and new species descriptions (e.g.. 'Mailing address: 6056 Beeman Ave.. No. Hollywood, CA 91606 Gosliner, 1981; Behrens, 1984; and Millen, 1985) being published. These add significantly to the known species of opisthobranchs that are endemic to this coastline or its various marine faunal provinces, or that are shared with other marine zoogeographic regions. In this paper we describe a new species of Tritonia that occurs in the Californian marine faunal province; the ex- istence of this species was first reported over 5 years ago (Behrens, 1980). Vol. 100(2) April 30, 1986 THE NAUTILUS 47 Order Nudibranchia Suborder Dendronotacea Family Tritoniidae Lamarck, 1809 Tritonia myrakeenae Bertsch & Mozqueira, n. up. Syno)ii/>ny: Tritonia sp. Behrens, 1980: 102-103 (includes a color photograph of the liv- ing animal). Type Locality: Southeast end of Isla Cedros, Baja California, Mexico, rocky reef west of lighthouse, in Bahfa Sudeste: 28°2'30"N; 115°12'W. Type Material: Holotype: Approximately 7 mm long while alive, 4.5 mm preserved; inter- tidal zone, rocky reef SE end of Isla Cedros; leg. H. Bertsch and Soa Tsung, 26 December 1985. Deposited in the collection of the Los Angeles County Museum of Natural History, Dept. of Malacology, No. LACM 2135. Paratype: Approximately 6 mm long while alive, 4 mm preserved; intertidal zone, rocky reef SE end of Isla Cedros; leg. H. Bertsch and Soa Tsung, 26 December 1985. Deposited in the collection of Los Angeles County Museum of Natural History. Dept. of Malacology, No. LACM 2136. Para types: Three specimens, 3.5, 4, and 4.5 mm preserved lengths; intertidal zone in front of Hotel Puesta del Sol, El Sauzal, approximate- ly 8 km north of Ensenada, Baja California, Mexico (3r52'N; 116°41'W); leg. A. Mozqueira, September 1984. Deposited in the collections of California Academy of Sciences, Dept. of Inver- tebrate Zoology, No. CASIZ 061379. Additional Records and Range: The authors and Jesus Pineda have found specimens infre- quently in the intertidal region at El Sauzal (in front of Hotel Puesta del Sol), Baja California, during June, September, October and Novem- ber 1983, and February and March 1984. Behrens (1980: 102) reported the occurrence of Tritonia myrakeenae (as Tritonia sp.) from "Santa Barbara to San Diego, California." All known records of this species are only from the intertidal region. At this time, the known r'ange of Tritonia myrakeenae is from Santa Barbara, southern California. U.S.A., to the SE tip of Isla Cedros, near the southern limit of the state of Baja California, Mexico. Description: A thin, elongate, delicate- appearing tritoniid, 6-18 mm in total length (alive). Oral veil with 4-7 (rarely branching) ten- tacular processes. Foot broadly expanded laterally beyond sides of body; anterior margin gently rounded, bilabiate; posteriorly extending out flat past the body, terminating in a pointed tip. Retractile rhinophores surrounded by basal sheath; the rhinophore stalk bears 10-22 vertical processes which encircle the shaft below the tip. Dorso-lateral edge with a delicate, undulating free margin from which grow the branchial pro- cesses (often 9 on each side). Each branchial ap- pendage branches distally into 3-5 filaments. Color a very pale dirty orange-brown (see color photo in Behrens, 1980: 103); oral veil and sides of foot clear of color, partly opaque. Dis- tinctive white patches occur on the dorsum (often as a matched pair on each side of the midline); Behrens (1980: 103) illustrates an animal with 4 transverse groups of opaque white patches (or streaks) arranged down the length of the back. Specimens from Ensenada and Isla Cedros (Figiu'e 1) had fewer white splotches (usually just a pair located about Vs of the body length distant from the anterior end). Radula (Figure 2 A) about 1.1 mm long, 0.65 mm across at widest portion. Radular formulae of 2 specimens collected 26 September 1983 at El Sauzal were: 22 (10.1.1.1.10) and 25 (8-10.1.1.1.8-10) (widest row damaged but ap- peared to have 12 outer lateral teeth). Central rachidian tooth prc.iminent (a|)proximately 63 microns wide), with 3 posteriorly-directed cusps (Figures 2B and 3B). First lateral tooth well dif- ferentiated from the others, its structure a hooklike cusp curling over the side of the rachi- dian (Figiu'es 2B and 3B). Succeeding laterals all fairly straight, lilade-like cusps. Jaws elongate (P^igin-e 3A), each about 1.3 mm long and 0.37 mm wide; a row of pointed mam- 5 mm FlCi. 1. Tntiiiiin iiiijrakfi'xiif. xpt'c. tii)i\, dorsal view of liv- ing animal. Drawing by C. Yanes. 48 THE NAUTILUS April 30, 1986 Vol. 100(2) 2 A 2 B FIG. 2. Camera lucida drawings (scale represents 50(j) of: 2A. Major shape and features of entire radula, and 2B. Raehi- dian tooth, innermost lateral, and four outer lateral teeth of Tri.toiiia niyra- keenae. FIG. 3. Tritonia myrakeenae. sketches of jaws (3A) and selected radular teeth (3R, rachidian, innermost lateral tooth, and two outer lateral teeth). millate denticles begins on the inner margin at the joined region of the jaws, continuing as a row along the entire inner edge, increasing about Va of the length to 3-5 rows; the inner den- ticled margin terminates distally in a small blunt process or free flange. Discusfiion: Tritonia myrakeenae is readily distinguished from other north Pacific species of Tritonia. Tritonia hawaiiensis Pease, 1860, has much more prominent and more highly branched branchial processes and tints of orange-pink and dirty light-purple (Bertsch & Johnson, 1981: 84-85) which are absent in our new species. Tritonia insulae (Baba, 1955) has vermilion-tinted gills and a different radular for- mula: 30 (40-46.1.1.1.40-46). Tritonia diomedea Bergh, 1894, has a distinct white line bordering the edge of the foot, and a larger radular for- mula. Tritonia festiva (Stearns, 1873) has a series of white lines and loops reticulating on the dorsum and a radula with more teeth in each row. Tritonia pickensi Marcus & Marcus, 1967, has a distinct white swath running down the center of the dorsum with lateral extensions to the branchial processes; although it has a similar radular formula, the shape of the rachidian tooth (narrower and deeper indentation be- tween the cusps) and the presence of small den- ticles on the cusp of the first lateral tooth (Bertsch & Gosliner, 1984) are differentiating features. Species of Tritonia in the North Atlantic are also easily distinguished from T. myrakeenae. The tropical T. welhi Marcus, 1961 (b), is pink and white, and the shapes of the teeth are differ- ent (cf. Marcus & Marcus, 1967: 100, fig. 130), and T. bayeri Marcus & Marcus, 1967, has an opaque white diffuse network over its dorsum. Tritonia nilsodhneri Marcus, 1983, is rose-pink; T. m.anicata Deshayes, 1853, has red, black or olive-green spots on the dorsum; T. plebeia Johnston, 1828, has white stippling but is darker colored with brown mottling and has more teeth per half row (21-33); T. lineata Alder & Hancock, 1848, has 2 conspicuous longitudinal white lines running lengthwise down the dor- sum; although T. homhei-gi Cuvier, 1803, has white splotches, its body shape is different (tubercled dorsum, more prominent bilobed oral veil) and the radula is completely different (den- ticles on teeth of young specimens and 27-158 teeth in a half row). Thompson & Brown (1984) give further anatomical characteristics that distinguish these latter 5 species from T. myrakeenae. Mrs. Marcus (1983) details the anatomy of other Tritoniidae, none of which are comparable with Tritonia myrakeenae. Etymology: This new species is named in honor of the distinguished malacologist, Dr. A. Myra Keen (1905-1986), colleague and friend, who included nudibranchs in Sea Shells of Tropical West America, second edition (this was the first major review of all known nudibranchs and other opisthobranchs from the Panamic marine faunal province). Her persistent scholar- Vol. 100(2 Af)ril 30, 1986 THE NAUTILUS 49 ship and encouragement of research has helped us to know (and hopefully to protect) the mollusks of western North America. Acknowledgments Field work was partially supported by a grant (to the senior author) from Have Mule Will Travel (Alpine, California), and with the co- operation of the Escuela Superior de Ciencias Marinas, Universidad Autonoma de Baja California, Ensenada (Guadalupe Ballesteros, Director). We especially thank Jesus Pineda for collect- ing assistance in Ensenada, and the Cedros Island Have Mule Will Travel research expedi- tion team (including Miss Soa Tsung) of December 1985-January 1986. The illustration of the living animal of Tritonia myrakeenae (Figure 1) was drawn by Senorita Clara Yanes. LITERATURE CITED Behrens, D. W. 1980. Pacific coast nudibranchs: a guide to the opisthobranchs of the northeastern Pacific. Sea Chal- lengers, Los Osos. 112 pp. 1982. Sakuraeolis en.osimensis (Baba, 1930) (Nudibranchia: Aeohdacea) in San Francisco Bay. Veliger 24(4):359-363. . 1984. Notes on the tergipedid nudibranchs of the northeastern Pacific with a description of a new species. Veliger 27{l):6b-7l. Bertsch, H. 1981. Rectification of the generic placement of Sclewdoris tanyn (Marcus, 1971). comb. tior.. a nudi- branch from southern California, with a range extension to the Gulf of California, Mexico. Veliger 23(3):217-220. Bertsch, H., and T. Gosliner. 1984. Tritoyiin pickeiisi (Nudi- branchia: Tritoniidae) from Baja California, Mexico. Shells and Sea Life 16(9):138-139. Bertsch, H., and S. Johnson. 1981. Hawaiian nudibranchs. Honolulu, Oriental Publ. Co., 112 pp. Gosliner, T. M. 1981 . A new species of tergipedid nudibranch from the coast of California. Jour. Moll. Studies 47:200-20,5. Gosliner, T. M., and S. V. Millen. 1984. Records oiCuthoria pustulata (Alder & Hancock, 1854) from the Canadian Pacific. Veliger 26(3):183-187. Keen, A. M. 1971. Sea shells of tropical west America, sec- ond edition. Stanford, California. Stanford University Press, 1064 pp. MacFarland. F. M. 1966. Studies of opisthobranchiate mol- lusks of the Pacific coast of North .-Vmerica. Mem. Calif Acad. Sci. 6:546 pp. Marcus. Er. 1961 a. Opisthobranch mollusks from Califor- nia. Veliger 3 (Suppl.):l-85. 1961 b. Opisthobranchia from North Carolina. Jour. Elisha Mitchell Scient. Soc. 77:141-151. Marcus, Ev. du Bois Reymond. 1983. The western Atlantic Tritoniidae. Bolm. Zool. Univ. S. Paulo 6:177-214. Marcus, Er. and Marcus, Ev. 1967. American Opisthobranch Mollusks. Studies Trop. Oceanogr. 6:1-256. McDonald, G. 1983. A review of the nudibranchs of the Cali- fornia coast. Malacologia 24(1-21:114-276. Millen. S. V. 1985. The nudibranch genera Onchidons and Diaphorodoris (Mollusca, Opisthobranchia) in the north- eastern Pacific. VV/((/fv28(l):80-93. Stearns, R. E. C. 1873. Descriptions of a new genus and two new species of nudibranchiate mollusks from the coast of California. Proc. Calif Acad. Sci. 5:77-78. Thompson, T. E., and G. H. Brown. 1984. Biology of Opisthobranch Molluacs. Volume II. London, The Ray Society, 229 pp. UNIONIDAE OF THE UPPER CONNECTICUT RIVER, A VANISHING RESOURCE' Arthur H. Clarke ECOSEARCH, Inc., 325 East Bay view Portland, TX 78374 Introduction It was my good fortune to meet William J. Clench in the winter of 1949-50, shortly after having become irrevocably committed to shell collecting. His personality was so enthusiastic and friendly and the Mollusk Department was so 'In memory of my good friend and teacher, William J. Clench. impressive that I soon made frequent trips to the Museum of Comparative Zoology at Har- vard, known familiarly as the "MCZ", and later became a Saturday volunteer. It was not long after that I resolved to seek a career in malacology. I shall always be grateful to Bill Clench and to Ruth Turner for the inspiration to make that decision and for their numerous acts of kindness since that time. 50 THE NAUTILUS April 80, 1986 Vol. 100(2) I well remember our first collecting trip to the Connecticut River. The occasion was the 1952 annual field trip of the Boston Malacological Club; the site was the Connecticut River at South Hadley, Massachusetts; the leader was Dr. Clench; the weather was warm and sunny; and the water was low. Bill, as his friends called him with affection and respect, led us to a large sandbar in the middle of the river. Beautiful mussels were everywhere in 2 or 3 feet of water and the Lampsilis cariosa were the most massive that I had ever seen. During" the following years Arthur and Louise Clarke collected widely in New England and at several localities in the Connecticut River System. There we found Alasmidontu hetero- don. atrangeAhismidonta uruiulata. and several other species which, to my neophyte eyes, were especially marvelous. Although the Connecticut River fauna had recently been reported upon (Clench and Russell, 1939, 1940), it still merited additional investigation. It was not until nearly 30 years had passed, however, that I was able to do "it. It is appropriate that the results of that survey, although principally of regional interest, should be dedicated to Bill Clench. In a larger sense, however, all of my research has been in- fluenced by his teaching and it will always con- tinue to be so. I would also like to thank The Nature Conservancy for funding the field work in 1983 and my wife Judith, and Drs. L. L. Master and T. French, for valuable field assistance. Preparation of the manuscript was supported by ECOSEARCH, Inc. Methods and Results The primary objective of the 1983 program was to determine the present distribution and abundance of Alasruidovfa hetcrodon (Lea) in the Connecticut River System of New Hamp- shire and Vermont. During the course of that work tabulation of habitat attributes and of the identity and abundance of all other mollusks en- countered were also made. The detailed results concerning/!, keterodori will be reported as part of a status survey, now underway, of that species throughout its entire range, but the general results for 1983 are reported here. The principle field activities were carried out over an 18-day period in August and early September, 1983. Collecting equipment in- cluded a motor-assisted canoe, wet suits, view- ing boxes, and a small Kolkwitz dredge. A total of 36 survey areas were studied. These are shown in Map 1 and, to conserve space, are ab- breviated and listed below in narrative form. Station List: 1, Third Conn. L, E side, N.H.; 2, Conn R between Third and Second Conn. L; 3, Second Conn. L, S side; 4, First Conn. L, N side, N.H.; 5, Conn. R just above Francis L, N.H.; 6, Conn. R above Canaan, Vt.; 7, Conn. R 1.0 mi N of Tinkerville, Columbia Twp., N.H.; 8, 9, Conn. Mass. FU;. 1. The upper Connecticut River System with 1983 study sites indicated. Vol. 100 (:; April 30, 1986 THE NAUTILUS 51 R, by canoe from N Stratford to 2.4 mi above Stratford, N.H.; 10, Conn. R, 2.4 mi above Stratford, N.H.; 11, Conn. R, flooded marginal pool, above bridge at Guildhall, Vt.; 12-14, Conn. R, by canoe from Comerford Dam to bridge at Barnet, Vt.; 15, Conn. R, S Newbury, Vt., V4 mi above Oxbow Bridge; 16, Conn. R, 1.0 mi NW of Piermont, N.H.; 17, Conn. R, Hart- land, Vt.; 18, Conn. R, Weathersfield Bend, 4 mi S of Ascutney, Vt.; 19, Conn. R, below bridge from N Walpole, Vt. to Bellows Falls, N.H.; 20, Nulhegan R, 0.5 mi W of Bloomfield, Vt.; 21, Paul Stream, 1.1 mi above mouth, Brunswick Twp., Vt.; 22, Upper Ammonoosuc R, Groves- ton, N.H.; 23, Forest L, Dalton Twp., N.H.; 24, outlet of Hazen's Pond, 2.5 mi E of Whitefield, N.H.; 25, lake at town park, Thetford Twp., Vt; 26 Mud Pond, Thetford Center, Vt.; 27, Ompom- panoosuk R, 0.5 mi S of Union Village, Vt.; 28, Croydon Branch, 1.0 mi S of Croydon, N.H.; 29, Sugar R, 2 mi SW of Croydon; 30, Williams R, 0.5 mi N of Rockingham, Vt.; 31, Ashuelot Pond outlet, 3.0 mi NE of Marlow, N.H.; 32, Ashuelot R, 0.4 mi W of Ashuelot Pond; 33, Asheulot R just above Marlow, N.H.; 34, Ashuelot R, be- tween Shaw's Corner and Roundy's Corner, N.H.; 35, Ashuelot R, 3.0 mi S of Surry Moun- tain Dam; 36, Ashuelot R, 2.4 mi NW of center of Keene, N.H. In most cases living specimens were tabulated and promptly returned to their habitats. In the tabulation below, each station number is fol- lowed by a syml)ol which reflects the number or abundance of specimens seen there. Numbers of living (no parenthesis), recently-living (in paren- thesis), and subfossil (in parenthesis, with S) specimens are cited separately. Other symbols are: A = abundant and C = common. No mollusks were found at stations 4, 5, 6, 12, 13, 14, 20, 22, 24, 27-31, 33, and 34. Species List Unionidae. M. inargaritifera: sta. 32, (2S). E. complnmita: 8,(1S); 15,3; 'l6,(25); 17,ca.500-i- (77); 18,2-H(1); 19.(5S); 23,12-h(l); 35,A; 36,A. An. cataracta: l,12-i-(6); 2,6-i-(2); 3,41 -i-(3); 11,(1); 23,2-^(1).^/. heterodon: 17,(5); 35,1-h(2); 36,(1). AL undulata: 1,1 -(-(9); 3,2; 9,(1S); 17,(24); 35,2-^(3); 36,4. St. undulata: 3,1; 17,(4); 35,1; 36,3-f(l). La7rip. r. radiata: 17,ca.300-i-(28); 19,(4S). Sphaeriidae Sphaerium striafinuni: 17,11. Viviparidae. Canipclonid decisum: 10,3; 16,27; 17,2; 36,18. Cipanqopaludina chinerhsis: 25, A; 26,A. Lymnaeidae. Juv. lymnaeid, sp.?: 7, 1. Physidae. f^hi/.^n heterostropha: 11, A; 16,1. Planorbidae. Helisovia anceps: 7,1; 8,1; 11,C; 16,4; 17,4. Ancylidae. Ferrisia rivularis: 9, A; 18, A. Discussion and Conclusions According to Johnson (1915), Clench and Russell (1939, 1940), and Clarke (1981), and specimen-associated data in the MCZ, USNM, and ANSP, 10 species of freshwater mussels oc- curred in the Connecticut River prior to and during part of the first few decades of this cen- tury. They are Elliptio complanata, Anodonta cataracta. An. implicatn. Alasmidonta hetero- don, Al. undulata. Al. varicosa, Strophitus undulatus, Lampsilis cariosa, L. ochracea and L. r. radiata. Three of these (An. implicata, L. cariosa and L. ochracea) were not known to ascend the river above Massachusetts. Many widely-distributed species of Sphaeriidae, proso- branchs, "and pulmonates were also known to occur there (Johnson, 1915; Clench & Russell, 1939) and another unionid (Ligumia nasuta) is also known from the vicinity of Keene, N.H., presumably in the Ashuelot River drainage (Walker and Coolidge, 1908). The principle ob- jective of this paper is to discuss the unionids of the main river, but the records listed above from some tributaries (e.g. the introduced gastropod, Cipangopaludina chinensis) may also be of interest. Early in the course of our work it became ob- vious that the rnolluscan fauna, and especially the unionids, of the Connecticut River has been greatly harmed by human activities. Dams, the construction of which began in the 1930's, have altered this once free-flowing river so that it is now chiefly a series of impoundments within which water levels are drastically manipulated by computer controlled dams for the generation of electricity and for flood control. Pollution from the pulp and paper industry has further ex- acerbated the situation. The probable distribu- tions of the unionids in the upper Connecticut River, as they existed prior to historical degredation of the river, are shown in Figau-e 2. 52 THE NAUTILUS April 80, 1986 Vol. 100(2) c < ■D C 3 ■D C 3 V (0 4 5" P ■ • 44' • — 4 3' • o • o • • Vt/NH Mass FIG. 2. Probable distribution of unionids in upper Connec- ticut River prior to pollution and impoundment (white bars and open circles) contrasted with their 1983 distributions (black bars and black spots). Locations are expressed in terms of north latitude. Pre-impoundment distributions are based on subfossil specimens (stations 8 and 19), live specimens collected by W. .J. Clench from Conn. River near M(mroe, N.H., and Wells River, Vt., in 1940, other records at MCZ, I'SNM, and ANSP, and Clench and Russell (1939. 1940). The di.strihutioii of those species as observed in 1983 are also shown on that figure. Based on the presence or absence of fresh- water mussels, it is useful to consider the upper Connecticut River as now comprised of 3 faunal regions. Region 1 (stations 1-3) includes the uppermost part of the system, i.e., the Second and Third Connecticut Lakes and the interven- ing portion of the Connecticut River. (The u{)- permost Fourth Connecticut Lake is small and inaccessible by road; it was not sampled). Signif- icantly, the water levels in this part of the system are not manipulated but are left to fluc- tuate moderately and naturally. This region con- tains large poy)ulations oi Anodonta rnturnrta. (specimens of which have a i)eculiar subarctic fades), of Alasmidontd undulata morph sower- hyuna. (an ecophenotyj)e which has been shown (Clarke, 1981) tiir. Inrert. Pnth. 17:.53-i58. 1973. Genetics of Bior/ijihulnrin glahmta (Gas- tropoda: Planorbidae). Mtilaeol. Reriew 6:199-202. and Merritt. J. W. 1972. Genetic factors in the susceptibility of juvenile Bumiphalnrin qlabrnta to Schistosomii in5 will be considered. Although almost all species could occur in lakes, this water body type was the major type of habitat for all of the amni- colids, and ValiHita spp., Stagnicola catasco- pium, Fossaria decampi and Helisoma carnpa- nulatum. Ponds were the major water body type for Stagnicola caper ata, Fossaria exigua, Planorbula campestris, Armiger crista, and to a lesser extent, Stagnicola palustris. Aplexa hypnormn and Physa jennessi (ssp. skinneri Taylor, 1953). Bulim.nea megasowM appeared to favor rivers above other habitat types when compared to the overall sampling distribution. In general, running waters were preferred over lentic habitats by Stagyiicola rejlexa. Bulimnea megasoma. Ferrissia rivularis and Planorbula armigera. Gyraulus deflectus occurred often in both running waters and lakes but appeared to avoid ponds. Distribution of gastropods with respect to substrate type (Table 2) showed that Ferrissia rivularis and Stagnicola reflexa were commonly found in granitic Precambrian Shield basins, although they also occurred on clay and coarse sediments respectively. Planorbula campestris occurred mostly on coarse and silt sediments. Sand appeared to lie preferred by Valvata since7'a. V. tricarinata., Cincinnatia cincinna- tiensis. Marstonia decepta. Probythinella lacustris, Stagnicola catascopium, Fossaria decampi, Helisoma campanulatum and Pro- nienetiis umbilicatellus. Fossaria exigua and Stagnicola caperata favored both sand and clay, while clay was the major sediment type for Stagnicola. palu.stris, Fossaria modicella and Helisoma pilsbryi. Physa jennessi preferred both silt and clay. Physa gyrina, Lymnaea stagnalis. Helisoma anceps, H. trivolvis, Planor- bula armigera and Gyraulus parvus appeared to 58 THE NAUTILUS April 80, 1986 Vol. 100(2) TABLE I . Percc'ntai;<' ciislriliiiti10 ha <10 ha -Zm deep '2ni deep Campeloma decisum Say, 1816 Valvata sincere Say, 1824 V. tricarinata Say, 1817 Cincinnatia cincinnatiensi s Anthony, 1840 Marstonia decepta F.C. Baker, 1928 Probythinella lacustris F.C. Baker, 1928 Amnicola 1 imosa Say, 1817 A^ walkeri Pilsbry, 1898 Lymnaea staqnal is Linne, 1758 Acella haldemani "Deshayes" Binney, 1867 Bui imnea megasoma Say, 1824 50 50 57 29 84 14 2 73 4 23 100 86 71 80 40 100 30 Stagnicola palustri s Muller, 1774 29 S_. catascopium Say, 1817 S^ caperata Say, 1829 S^ reflexa Say, 1821 Fossaria da11i F.C. Baker, 1907 1896 92 14 29 50 75 7 35 20 47 36 22 F. decampi Streng, F. exigua Lea , 1841 F. modicel 1 a Say, 1825 F^ parva Lea, 1841 Physa qyrina Say, 1821 P^ jennessi Dall, 1919 Aplexa hypnorum Linne, 1758 Ferrissia paral lela Haldeman, 1841 F_^ rivularis Say, 1817 45 Hel isoma trivolvis Say, 1816 42 H. pi 1 sbryi inf racarinatum F.C. Baker, 1932 42 FL corpulentum Say, 1824 50 ti. campanulatum Say, 1821 81 H^ anceps Menke, 1830 47 Planorbula arniigera Say, 1821 41 P^ campestris Dawson, 1875 29 Promenetus exacuous Say, 1821 51 P. umbili eaten us Cockerell, 1887 43 Armiger crista Linne, 1758 33 Gyraulus parvus Say, 1817 43 G^ circumstriatus Tryon, 1866 41 G. deflectus Say, 1824 59 14 4 7 44 26 14 7 7 19 3 69 20 5 47 6 7 246 3 35 26 9 23 57 6 8 170 8 12 86 7 29 29 13 7 25 25 4 12.5 12.5 8 86 7 14 52 9 4 46 80 5 36 10 7 253 58 6 33 62 5 11 37 100 3 19 23 13 31 43 7 8 189 37 21 19 50 4 4 15 75 44 3 6 106 29 16 14 68 71 7 31 11 7 55 43 14 7 67 9 45 5 7 161 48 4 7 27 16 14 11 44 TOTAL SITES 40.5 43.0 9.0 7.5 412 be substrate-indifferent. The mean gastropod community diversities where each species was found were higher than the mean overall site diversity (Table 3) because all species except Sfngnicola rcjlexa occurred to some extent in a proportion of highly diverse comnuinitii's. For species with N>5, Fossaria ilcrnmiii and Frohnthinclla hirustris. both primarily lacustrine species, showed the highest diversity values. Low mean (liversities were seen for communities in which Stagnicola rejlexa, Ferrissia rivularis. Physa gyrina. Stag- Vol. 100(2) April 30, 1986 TABLE 2. Perceiitaije ilislriliutimi acconiing to sul)strate type in thf study area. THE NAUTILUS 59 Species Gravel/ Bedrock coarse Granitic Litnestone Shale sand Sand Silt Clay Organic Campeloma decisum 75 25 4 Valvata sincera 14 57 29 7 V. tricarinata 4 2 11 46 2 20 15 46 Cincinnatia cincinnatiensi s 15 4 62 19 26 flarstonia decepta 88 12 8 Probythinel la lacustris 29 57 14 7 Amnicola limosa 19 12 39 1 16 13 69 A. walkeri 33.3 33.3 33.3 6 Lymnaea stagnalis 6.5 2 1 17 24 6.5 23 20 246 Acella haldemani 100 3 Bulimnea megasoma 19 19 24 14 24 21 Staqnicola palustris 2 2 2 21 23 9 29 12 172 S. catascopium 8 8 8 50 18 8 12 S. caperata 57 43 7 S. reflexa 43 43 14 7 Fossaria dalli 50 25 25 4 F. decampi 12.5 75 12.5 8 F. exiqua 7 7 43 7 36 14 F. modicella 2 2 2 13 22 11 34 14 45 F. parva 20 20 20 40 5 Physa gyrina 12 2 1 15 28 5 22 15 251 P. jennessi 9 21 21 34 15 33 Aplexa hypnorum 3 3 34 18 5 26 11 38 Ferrissia parallela 33 67 3 F. rivularis 37 7 13 33 10 30 Helisoma trivolvis 4 2 1 18 24 9 25 17 189 H. pilsbryi inf racarinatum 16 4 32 32 16 19 H. corpulentum 25 25 50 4 H. catnpanulatum 16 1 10 49 4 6 14 73 H. anceps 8 19 28 2 19 24 107 Planorbula armijera 9 3 3 15 21 3 24 22 67 P. campestris 43 14 29 14 7 Promenetus exacuous 15 4 13 16 7 29 16 55 P. umbilicatellus 14 14 58 14 7 Armiger crista 11 23 33 33 9 Gyraulus parvus 5 1 1 16 26 9 21 21 161 G. ci rcumstriatus 8 4 23 34 4 8 19 26 G. deflectus 12 2 9 35 5 12 26 43 TOTAL SITES 10.8 2.0 0.9 15.9 27.0 5.9 19.6 17.9 408 nicoln pdlusfris, Lymnaea stagnalis, Helisoma anceps. Bulimnea m.egasnma. Physa jennessi. Fossaria exigua and Helisoma trivolvis were present. The maximum ret-onled gastro|)od community diversity was 17. Mean macrophyte community diversity was high for Valvata sincera. Gyrauluji deflectus. Hciisonut campanulatum, Bulimnea megasonta and all amnicolids except Prohythine.lla lacuMris. Low mean macrophyte diversities {<6) were seen for Physa jennessi, Fossaria modi- cella, Probythinella laciistris. Fossaria exigua. Stagnicola palustris and S. capcrala. Of tlie above species, mean diversity values were low for both gastropod and macrophyte communi- ties for Physa, jennessi. Fossaria exigua and Stagnicola i)alu.' 4 pH units). The narrowest ranj^es (< 2 units) were seen for Planorbula campestris, Mar- stonia decepta and Fossaria decampi; these three species liowever were sampled at relative- ly few sites. The highest sampled pH of \()Ji was tolerated by Stagnicola palustris, Fassaria modicella, Helisoyna trivohns, Promenetus exacuous, Aryniger rj-ista and Gyraulus parvus, while the lowest uH of 5.0 was observed for Bulimnea megasonia. Ferrissia rivularis, Helisoma trivolvis, H. anceps and Gyraulus dejlertus. The widest ranges for total dissolved solids, encompassing nearly the total range sampled, were seen for Fo.nsaria modicella. Stagnicola fiatustris and Physa jennessi (Table 4), followed by somewhat smaller ranges (approx. 4500 mg/1) for Lymnaea. stagyialis and Helisoma trivolvis. Narrow ranges (300 mg/1 or less), all towards the bottom end of the scale, were seen for Vol. 100(2) April :;o, i;)S6 THE NAUTILUS HI TABLK 1. Minimuni. iikixiiiiuiii ami luraii \aliK'S fcir pH ami tutal disscilvt'd solids for ffaslnipods in the stud\ area. Species pH Total dissolved solids, mg/1 Min. Max. x H Min. Max. x H Campeloma dec i sum 7.1 8.1 7.8 3 58 123 87 3 Valvata sincera 6.2 9.2 7.4 6 134 384 264 6 v. tricarinata 6.7 9.8 8.3 48 33 1794 432 47 Cincnnnatia ci nci nnat lens i s 7.3 9.8 8.3 25 33 875 170 24 Marstoma decepta 7.7 9.2 8.4 9 60 208 134 7 Probythinel la lacustris 7.9 9.5 8.5 5 104 1300 494 5 Amnicola limosa 6.1 9.8 8.1 80 16 1794 235 79 A. wall 700mg/l CaCOj) for Stagnicola palustris. Gyraulus deflectus, G. circumstriatus and Physa jennessi. Narrow ranges (<100 nig/l CaCOs), all again towards the bottom end of the scale, were seen for Stagyiicola reflexa and Valvata sincera. both of which were repre- sented by small numbers of samples. Thirteen species occurred at alkalinity values as low as 0-4 mg/1 CaCOa, while only three were observed at the maximum sampled value of 800 mg/1 CaCOs. All species tolerated chloride values of 0 mg/1 (Table 5), but only Stagnicola palustris, Fossaria modicella and Physa jennessi could be found throughout the complete range sampled. Valvata sincera and Stagnicola reflexa ap- parently avoided chloride altogether, although again these were represented by small sample numljers. Fossaria decampi and Marstonia decepta showed chloride ranges of < 10 mg/1. 62 THE NAl'TILUS April 30, 1986 Vol. 100(2) TABLE 5. Mininiuiii. niaxinium and mi'aii valufs tor total alkalinity and chloridi- for tjastrupods in the study area. Species Total alkal i ni ty . , mg/1 ) Chloric le, mg/1 CaCO., Min. Max. " X N Min . Max. x N Campeloma decisum 64 128 97 3 0 0 0 3 Valvata sincera 70 166 119 6 0 0 0 6 V. tricarinata 0 448 144 47 0 480 70 47 Cinci'nnatia cincinnatiensis 14 239 91 24 0 18 3 23 Marstonia decepta 70 220 122 8 0 8 1 8 Probythinel la lacustris 70 159 132 5 0 238 48 5 Amnicola limosa 4 448 96 78 0 480 34 77 A. walkeri 11 172 63 5 0 8 2 5 Lymnaea stagnalis 0 560 135 255 0 592 25 254 Acella haldemani 72 100 81 3 0 0 0 3 Bulimnea megasoma 9 172 54 29 0 56 3 28 Stagnicola palustris 4 800 180 172 0 1234 51 172 S. catascopium 84 184 134 11 0 430 130 11 S. caperata 106 224 173 3 0 7 2 3 S. reflexa 0 72 31 7 0 0 0 7 Fossaria dalli 72 202 129 3 0 4 1 3 F. decamgi 70 284 138 6 0 2 0. 3 6 F. exigua 36 560 183 13 0 77 10 13 F. modicella 48 448 194 48 0 1234 115 48 F. parva 108 230 167 4 0 21 5 4 Physa gyrina 0 560 122 263 0 592 24 262 P. jennessi 62 768 196 28 0 1234 79 28 Aplexa hypnorum 4 448 179 33 0 438 17 33 Ferrissi'a parallela 128 172 150 2 0 0 0 2 F. rivularis 4 220 68 38 0 238 9 37 Hell soma trivolvis 0 654 144 195 0 480 22 195 H. pilsbryi inf racarinatum 10 316 106 15 0 261 18 15 H. corpulentum 10 41 29 5 0 0 0 5 H. campanul atum 10 280 71 89 0 15 0. 5 88 H. anceps 4 448 115 111 0 470 23 111 Planorbula armigera 0 560 147 67 0 480 49 67 P. campestris 148 280 210 6 0 13 3 6 Promenetus exacuous 4 654 153 60 0 480 45 59 P. umbilicatellus 12 266 107 6 0 22 4 6 Armiger crista 80 448 241 10 0 480 146 10 Gyraulus parvus 0 654 134 174 0 592 28 174 G. circumstriatus 25 800 222 26 0 602 46 26 G. deflectus 20 800 99 47 0 394 13 46 TOTAL SITES 0 800 122 440 0 1234 24.4 440 All species (except Stagnicola caperata) were observed in the absence of measurable sulphate (Table 6), but, as for chloride, only Sttignicota paliiHtris, Fossaria modicella and Physa. jen- nessi were found throughout the entire range sampled. Valvata. sincera. Stagnicola reflexa, Fossaria decampi and Bulimnea megasoma ap- peared to be relatively intolerant of sulphate. For nitrate and nitrite (Table 6), Cincinnatia cincinnatiensis, Planorbula armigera and Stag- nicola palustris occurred throughout all or most of the range sampled. Narrow ranges towards the lower end of the scale were seen for Stagnicola re.jlexa. and Marstonia decepta. Eleven species were observed to tolerate the highest phosphorus levels (Table 7). Bulimnea megasoma and Stagnicola rcjlexa appeared to favor low phosphorus concentrations. li Discussion the study area only niarginallv more species were found in lakes than in ponds. Although none of the species appeared to be restricted to either lakes or ponds, many Vol. 100(2) April 30, 1986 THE NAUTILUS 63 TABLE (). Miiiinuiiii, niaxiiiiuni .uid mean values fur sulphate and nitrate/nilrile for t;astrn|ioils in the study area. Species Sulphate, mg/l Min. Max. x Mi trate/nitri te, mg/l N Min. Max. x N Campeloma decisum 0 0 0 3 0 0.7 3 Valvata sincera 0 2 0. 5 7 1.0 3.3 1.6 6 V. tricarinata 0 170 51 54 0 2.4 1.3 48 Cincinnatia c i nci nnat lensis 0 170 21 25 0 7.5 1.1 25 Marstonia decepta 0 109 13 9 0 1.5 0.8 9 Probythinel la lacustris 0 170 69 6 0 1.2 0.5 5 Amnicola limosa 0 170 23 85 0 2.7 0.9 78 A. walkeri 0 15 4 5 0 1.5 0.9 5 Lymnaea staqnalis 0 334 28 263 0 5.5 1.2 258 Acella haldemani 0 0 0 3 0.8 1.0 0.9 3 Bulimnea megasoma 0 15 3 30 0 5.3 0.7 28 Stagnicola palustns 0 3403 65 182 0 7.2 1.6 174 S. catascopium 0 113 32 12 0.1 1.7 1.1 12 S. caperata 1.6 157 61 3 1.0 2.3 1.5 3 S. reflexa 0 10 2 7 0 1.4 0.5 7 Fossaria dalli 0 39 13 3 0.8 1.3 1.1 3 F. decampi 0 14 4 6 0 2.4 1.2 6 F. exigua 0 161 29 14 0.2 3.3 1.0 14 F. modicella 0 3403 129 49 0 5.6 1.6 49 F. parva 0 148 37 4 0.2 2.4 1.1 4 Physa qyrina 0 334 24 271 0 5.5 1.1 265 P. jennessi 0 3403 190 28 0 6.8 2.2 28 Aplexa hypnorum 0 165 32 37 0 2.4 1.3 35 Ferrissia paral lela 0 104 42 3 0.9 1.5 1.2 2 F. ri vulari s 0 161 18 36 0 5.3 0.6 36 Helisoma trivolvis 0 443 38 204 0 5.2 1.3 197 H. pilsbryi i nf racari natum 0 64 5 15 0 2.8 0.7 15 H. corpulentum 0 5 2 5 0.1 1.4 0.6 5 H. campanul atum 0 108 3 92 0 2.4 0.8 90 H. anceps 0 167 18 114 0 5.3 1.1 112 Planorbula armiqera 0 157 25 70 0 7.5 1.3 68 P. campestns 0 115 45 6 0.9 1.6 1.2 6 Promenetus exaciious 0 158 42 64 0 5.2 1.3 59 P. umbi 1 icatel 1 us 0 159 28 6 0.3 2.0 1.2 6 Armiger crista 0 128 63 11 0.1 2.9 1.4 10 Gyraulus parvus 0 170 32 181 0 5.2 1.2 176 G. CT rcumstri atus 0 158 35 26 0.7 6.8 2.0 26 G. deflectus 0 108 8 49 0 5.3 1.2 47 TOTAL SITES 0 3403 31 456 0 7.5 1.1 445 gastropods showed strong tendencies to occui- in one of tiiese liahitat types. Aho rt al. (19. F^colnKX <>t' freshwater Molhisca in the Engiish Lake District. ./..»/•. Anini. EciA. 19:124-146. 19.')7. Chemical analysis in ecokigj' illustrateii front Lake District tarns ami lakes. 3. Faunistic differ- ences. Priir. Lnm. Sm- Lniiilmi 167: 172-I7"i. Mackie, G. L. and L. A. Khppance. 1983. Intra- and inter- specific variations in calcium content of freshwater mol- lusca in relation to calcium content of the water. Jitiir. imill. &7h(//>.s' 49:204-212. Okland. J. 1979. Distriliution of environmental factors and fresh-water snails ((lastropoda) in Norway: use of Rum- pean invertebrate survey principles. Muliu-olnijii: 18: 211-222. Okland, .1. and K. A. Okland. 19,S(l. pH level and food organisms for fish ■ studies of UKK) lakes in Norway. In Proc. Internat. Conference on the ecological im|:iact of acid precipitation. Norway. 1980 (D. Drablos and \. Tollan, eds.). SNSF Project, pp. 326-327. Pip, E. 1978. A survey of the ecology and composition of submerged aquatic snail-plant ciminiunities. Cniuididn ■Inn,-. Zoi.l. 56:22(;.3-2279. 1979. Survey of tlie ecology of submerged aquatic macro|ihytes in central Canada. AijikiHc Hatdiiy 7:339-357. 1984. Ecogeographical tolerance range variation in ai|uatic macrophytes. Hydrobidloyid 108:37-48. 1985. The ecology of freshwater gastropods on the southwestern edge of the Precambrian Shield. Cariddinri Fii'hl-Nitt. 99:76-85. PSEUDOSUCCINEA COLUMELLA (LYMNAEIDAE) IN THE DOMINICAN REPUBLIC AND TRANSMISSION OF FASCIOLIASIS IN THE CARIBBEAN REGION Jose Gomez, Mercedes Vargas Departanieiitd dv Micrdhioiogia y Parasitologia Universidad Autonoma de Santo Domingo Dominican Reiuihlic and Emile A. Malek Depaftment of Tropical Medicine Tulane University New Orleans 70112-2699 ABSTRACT Pseudosuccinea columella (Sai/. 1817) is reported for the first time from the Dominican Republic and from the island of Hispaniola. It occurs together with Fossaria cuhensis (Pfeiffer, 1839) in Puerto Rico, Cuba. Jamaica, and Venezuela in addition to the Dominican Republic in the Caribbean region. The two lymnaeid species can be differentiated by shell and animal characteristics. FascioiiasiH is transmitted in this region p7-incipaUy by F. cubensis, but the occurrence of P. col- umella in some countries and it.^ jtrobaJite introduction into others may e.rlend the range of this disease. Pseudosuccinea columella (also referred to as Lymnaea columella and Lymnaea (Pseudosuc- cinea) columella), a North, Uentrtd and South American species, has a[)parenUy been intro- duced into certain islands of the Caribbean and in other regions such as Europe, Africa, Australia and New Zealand. Maiek and Chrosciechowski (1964) iind Malek and Cogswell (1980) reported its occurrence in Venezuela, Colombia, Costa Rica, Pantima and Brazil and reviewed its geogrtiphical distribution. F'araense (1982) also outlined its distribution in the neotropical region. In our freshwater snail survey in the Dominican Re{)ublic we encountered P. col- umella, but it was less common (3 localities) than the other lymnaeid, Fossaria cubensis (Pfeiffer, 1839) (28 localities). Previous reports of the molluscan fauna of the Dominican Republic did not include P. columella (see Crosse, 1891), nor did Alvarez and Cordero (1977) in their work on fascioliasis report P. col- umella; they only discussed F. cubensis. The two lymnaeid species can be differen- tiated morphologically as follows: The shell of P. columella has a low spire, much shorter than the Vol. 100(2) April 30, 1986 THE NAUTILUS «7 high and elongated aperture, with a large and expanded body whorl. The surface is sculptured with microscopic raised spiral periostracal threads. Lateral teeth of the radula are tri- cuspid. The penis sheath is short, less than half the length of the prej)utium. The prostate is long and narrowly, roundly cylindrical, with the proximal end somewhat enlarged. P. rohonella is aquatic, with a tendency to be amphibious. The adult shell of Fossarm cubensis is small, with well-impressed sutures and usually in- distinct spiral lines. The spire is conic, broadly pyramidal or acute, the aperture is much smaller when compared to that of P. columella. The columella is smooth; the inner lip of the aperture is folded backward, forming a moderate or broad flat expansion. The lateral teeth of the radula are bicuspid. The penis sheath is two-thirds the length of the preputium; the prostate is ovate and rounded at the ex- tremities. F. eubensis is amphibious. Reports of P. colitmelht in addition to F. cubensis from other Caribbean islands are from Puerto Rico (van der Schalie, 1948; Harry and Hubendick, 1964; Leon-Dancel, 1970) and from Cuba (Aguayo and Jaume, 1949). Specimens received from Ms. Charmine Crawford from Jamaica are also P. columella and F. cubensis. The two species are also known to occur in Venezuela (Malek and Chrosciechowski, 1964). P. columella, has not been reported from St. Lucia (Malek, 1965; Barnish el ai. 1980), Guadeloupe (Pointier, 1974, 1976), Martiniciue (Guyard and Pointier, 1979), or Haiti (Robart et ai. 1976); however, F. cubensis is present in these countries. Pilsbry (1910) described the subspecies F. citbensis aspiraus as Liitunaea cubensis aspirans from Barbados, West Indies. The liver tluke Fasciola hepatica. which causes fascioliasis in the Americas, occurs in cattle and sheep-raising countries, parasitizing these animals and other herbivores. Human fascioliasis has also been reported from alnwst every American country, including islands of the Caribbean. There are several early reports of high prevalence rates on Cuba and records of some human cases. There are also records of fascioliasis in the Dominican Republic (Ueno et ai, 1973; Alvarez and Cordero, 1977), Puerto Rico (Frame and Bendezu, 1978; Hillyer, 1981; Bendezu et a I.. 1982), Martinique (Gretillat, 1966, 1967), St. Lucia (Malek, 1965, 1980; Barnish et ai. 1980), and .Jamaica (Anonymous, 1960). Malek (1965) signaled the occurrence of fascioliasis in St. Lucia and later (1980) gave details of prevalence rates in several estates in the north, center and south and in the abattoir in Castries, the capital. Malek (1965) predicted the occurrence of lymnaeid snails on the island and shortly after the report was written F. cubensis was encountered in 4 localities. The snails from St. Lucia were infected exjierimen- tally with F. hepatica miracidia of St. Lucian origin, and were later photographed (Malek, 1980). F. cubensis from St. Lucia was also susceptible to the schistosome, Heterobilharzia aniericana (Malek, 1967). From all reports it seems that in the Carib- bean region the principal snail intermediate host of F. hepatica is F. cubensis. P. columella, although susceptible to experimental infection (Leon-Dancel, 1970), has not been found naturally infected where it occurs in the Carib- bean region, although natural infection has been reported in Brazil (Ueta, 1980). There is also a record by Van Volkenberg (1939) (cited in Leon- Dancel, 1970) and one by Hillyer (1981) in which P. cdhanella was stated to be an intermediate host in Puerto Rico. The natural and experimen- tal infections in P. columella indicate that its presence in some Caribbean countries, and its likely introduction in others, may extend the range of fascioliasis in the region. References in the literature suggesting that snails other than lymnaeids, for example Pliysa cuboixis in Cuba or terrestrial snails, can act as intermediate hosts for F. hepatica are er- roneous. Some authors tend to implicate snails other than lymnaeids as transmitters of fascioliasis when they fail to locate lymnaeids such as F. cubensis or P. cnhouclla in the Carib- bean region. Moreover, Leon-Dancel et al. (1971) tested the susceptibility of Physa cubensis and Aple.ra marmorata in Puerto Rico and found them refractory to infection with F. hejiatica. LITERATURE CITED Atrunv". ('. ('.. anil .lauriie. M. L. U)41). Faniilia Lyninaeidae. (in i:i47-l'.l.'>:i) ' 'atiihyi, ili /.,s Molliisms ,!>■ Ciiha, No. 4;t, Ciilia. 68 THE NAUTILUS April 30, 1986 Vol. 100(2) Alvarez. .1. M. V. and Conlcro. E. T. C. 1977. Fasciolosis bovina en la Republica Domiiiioana. PuU. Universidad Aulonoma S. Domingo, Vol. 224, Col. Cienciay Techti. No. 5:1-189. Anonymous. 1960. Annual report Ministry Agriculture .lanuiica, 31 December I9.')8. Barnish, 0.. Prentice, M. A. and Harris. S. 1980. Fasciola h<'i>(ittc. A. 1965. Freshwater and land snails of St. Lucia, the West Indies. Arner. Malncol. Union. Annual Reports, p. 38. 1967. E^xperiniental infection of several lym- naeid snails with Hetero))ithnrzi(i nmi'ricono. Jour. Pariiiiitol. 53:700-702. 1980. Snail Trnnaniitted Paraaitic Diseases, Vol. n, CRC Press, Boca Raton, Florida. 324 pp. and Chroscieehowski, P. 1964. Lymnaea (Pseu- dosuccinea) columella from Venezuela, and notes on distri- bution <){ P.'ieudosuecinea. The Nautilus 78:54-56. and Cogswell, F. B. 1980. Lymnaea (Pseudosuc.- cinea) columella in Colombia. The Nautilus 94:112-114. Paraense, W. L. 1982. Lymnaea riatrix and Lymnaea col- umella in the Neotropical region: A distributional outline. Mem. Inst. Clswaldo Cruz 77:181-188. Pilsbry, H. A. 1910. Lymnaea cubensis nspirans n. subsp. The Nautilus 23:\2{)'. Pointier, .1. P. 1974. F^iune malacologique dulcaquicole de I'ile de la (Hiadeloupe (Antilles Francaises). Bull. Mus. Hist. Natur.. Paris, 3rd Ser., No. 235, Zool., 1.59:905-933. 1976. Repartition locale el biogeographie des mollusques dulcaquicoles de la Guadeloupe (Antilles Francai.ses). Malacol. /?e<'i>H' 9:85-103. Robart, G.. Mandahl-Barth. G. and Ripert. C. 1976. Inven- taire, repartition geographique et ecologie iles mollusques dulcaquicoles d'Haiti (Caraibes). Haliotis 8:159-171. Heno, H.. Alvarez. .J. M. V., Mergen. A. M. R. de and Sanchez, V. M. 1973. Observation on the prevalence of parasitic diseases in cattle especially fascioliasis, in the Dominican Republic. Nat. Inst. Animal Hlth. Quarterly (Tokyo) 13:59-68. Ueta, M. T. 1980. Ocorrencia de infecvao natural dv Faxciola hepatica Linnaeus. 1758 em Lymnaea columella Say. 1817 no Vale do Paraiba. SP. Brasil. Rec. Saude Publico 14: 230-233. van der Schalie. H. 194cS. The land and fresh water niollusks of Puerto Rico. Misc. Publ. Mus. Zool. Univ. Michigan, No. 70. ANNOUNCING THE CO. A. 19S6 CONVENTION The Concholo^ists of America's 14th annual convention will he held at the Sheraton \'ankt'e Trader Hotel at Fort I Lauderdale, Florida, from July 1') throujjjh July 19, 198(5. The convention will he hosted hy the Broward Shell Cluh, and the CO. A. welcomes all shell collectors, whether novice or advanced, as well as shell dealers and scientists from aroinid the country and the world. For further information and pre-registration packets, please contact Ruth ('hesler. Conven- tion Chairman, at 7401 S. W. 7th Street. Planta- tion, Florida 8:5317. NEWS JULY 1986 AMU MEETING The American Malacological Union and the Western Society of Malacolojjists will have a joint meetin,ti; from July 1-5 at the Sheraton Hotel in the heart of historical Monterey, Calif. The three symposia will be on the biolojjy of opisthohranchs, life history of cephalopods and molluscan morphological analyses. Housing, travel and registration details, already sent to AMU members, may also be obtained from President Dr. James Nybakken, Marine Labora- tories. P.O. Box 450, Moss Landing, CA 95039-0450. Vol. 100(2) April ::5(), 1986 THE NAUTILUS 69 SOME ASPECTS OF THE EVOLUTION OF THE RUDIST PELECYPODS David Nicol Box 14376, University Station Gainesville, FL 32604 ABSTRACT The evolution of the rudist pelecypods was remarkably rapid even as compared to many mammalian groups. One new genus appeared every 1.900,000 years in the Late Jurassic, hi the Early Cretaceous, one neiv genus appeared every 1,370,000 years, hut iii the Late Cretaceous the rate of evolution accelerated great- ly when one new genus appeared every 3^5,000 years. For the entire time-span for the existence of the rudists, one new genus appeared every 738,000 years. Like the dinosaurs, the first rudists were of modest size. Beginning in the AUiian stage, some rudists attained a size as large as any pelecypods that have ever lived. Some additional aspects of the evolution of the rudists are given here because they are either not mentioned or inadequately covered in a manuscript now in press (D. S. Jones and D. Nicol: Origination, survivorship, and extinction of rudist taxa). Some of the data used herein were updated by examining the Zoological Record from 1966 to 1981. The main source of data is the Treatise on Invertebrate Paleon- tology (C. Dechaseaux and A. H. Coogan, 1969). The aspect that is particularly emphasized here is the rapid increase in diversity at the generic level, especially during the Late Cretaceous or the last 29 million years of the ex- istence of the rudists. The 15 Jurassic and Cretaceous stages listed in Table 1 are based primarily on ammonite zones and are not all of the same duration. The duration in years of each stage was taken from Palmer, 1983. The 15 stages can be subdivided into three groups for studying the changes of rates of evolution: the three Jurassic stages (Oxfordian through Titho- nian), the Early Cretaceous stages (Berriasian through Albian), and the Late Cretaceous stages (Cenomanian through Maastrichtian). Rudist evolution began at a rather modest rate in the Late Jurassic with a new genus ap- pearing approximately every 1,900,000 years. The evolutionary rate did not accelerate greatly during Early Cretaceous time when a new genus appeared every 1,370,000 years. How- ever, beginning with the Cenomanian, the earliest stage of the Late Cretaceous, evolution speeded up greatly, and a new genus arrived every 345,000 years. The evolutionary rate for TABLE 1. The I.t Jurassic and Cretaceous stages with their duration in millions of years, number of new genera per stage, and new genera per year. Oxfordian -Tithonian. Late Jurassic; Berriasian -Albian, Early Cretaceous; Ceno- manian- Maastrichtian, Late Cretaceous. stages Duration Mew :e^ £-n-rd in years jenera per ye^r ■■■'aastrichtia.a ",000,000 17 470,000 Campanian 9,500,000 13 730,000 Santonian 2,500,000 15 100,000 Coniacian 1,000,000 10 100,000 Turonian 2,500,000 13 192,000 Cenomanian 5,500,000 lo 344,000 Albian 15,500,000 10 1 , 55o,000 Aptian 0,000,000 7 "57,000 Barresnian 5,000,000 9 025,000 Hauterivian 7,000,000 0 — 0— Valanginian 7,000,000 5 1 ,400,000 rerriasian 0,000,000 4 1 ,500,000 Tithonian 8,000,000 4 2 ,000,000 Kimmeridgian 4,000,000 0 — 0— Oxfordian 7,000,000 0 1 ,107,000 Totals 94,500,000 12'! 73 ",000 the total 15 stages, or the complete life span of the rudists, is the appearance of a new genus every 738,000 years. This would be considered exceedingly rapid evolution amongst the pelecy- pods in a stable marine environment and is com- parable to rapid evolutionary rates in many THE NAUTILUS April ;i(), 1986 Vol. 100(2) groups of mammals. The rapid acceleration of evolution of the rudists during the Late Cretaceous appears to be based on two phenomena. First, the three larger and more rapidly evolving families (Caprinidae, Radiolitidae, and Hippuritidae) either did not appear until the Late Cretaceous or did not become diverse until that time. Sec- ond, the rudist faunas became more provincial and ecologically complex in the Late Cretace- ous. Endemism of the rudist genera increased greatly in the Late Cretaceous, and the rudist communities were generically most diverse in the Caribbean region during the Maastrichtian (Kauffman and Sohl, 1974). As more paleontological study is done in the more remote areas with Jurassic and Cretace- ous rocks in the Tethyan realm, a greater number of new genera discovered will be in the Late Cretaceous rather than in the Early Cretaceous and Late Jurassic. In the rates of evolution within the L5 stages in Table 1, it is worthwhile to point out a few additional features. After a good start in the Ox- fordian Jurassic, the evolution of the rudists slowed somewhat with a gap of no new genera in the Kimmeridgian, a time span of four million years. This slowing of the rate of evolution con- tinued into the first three stages of the Early Cretaceous with a climax in the Hauterivian when, again, no new genera have so far been discovered. The unusual aspect of the Hauteri- vian gap is that it encompassed approximately seven million years. Beginning in the Barremian, evolution accel- erated greatly and this may have been caused partially by the onset of the two most diverse families, the Caprinidae and Radiolitidae. The rate of evolution then slowed in the next two stages which end in Early Cretaceous times. New genera appeared with exceeding rapidity in the Cenomanian and continued unabated until the Campanian and then accelerated again in the Maastrichtian. In the latest Cretaceous, new rudist genera were still appearing at a very rapid rate. Although extinction of genera occurred rapid- ly in the Late Cretaceous, the large number of new genera in each stage tended, with some ex- ceptions, to increase the total number of genera in each stage to a maximum of 43 in the Maastrichtian. This total of 43 includes the holdovers from earlier stages as well as the new arrivals. The present total of rudist genera is 128 so that within a period of no more than eight million years, one third of all the rudist genera became extinct. Five of the seven rudist families also became extinct in the Maastrichtian. The decline and extinction of the rudists oc- curred in slightly less than 8.5% of their total geologic history. This sudden demise of the rudists at their zenith at the end of the Cretaceous is unlike the extinctions of the trilobites, graptolites, conodonts, and rostro- conchs where there was a long period of decline in diversity before extinction. Even the am- monites and dinosaurs experienced a longer period of decline than the rudists. It is most dif- ficult and highly speculative to explain such a sudden extinction in a group of marine pelecy- pods that had been so highly successful. The rudist pelecypods, like the dinosaurs, began with small-sized species. Some of the early rudists were about 20 mm long, and small- sized rudists are also found at the end of the Cretaceous in Maastrichtian strata. Particularly among the diverse families Caprinidae and Radiolitidae, some stocks attained large size as early as the Albian, and large species continued to occur into the Maastrichtian. However, even by Albian time, 50 million years had passed, which is more than half of the total time span of the rudists. Titanosarcolites attained a length of more than 2 m, Bournonia reached a diameter of more than 0.6 m, and the shell wall of Durania attained a thickness of at least 10 cm (B. F. Perkins, 1969, p. N751). These large rudists were among the largest pelecypods that have ever lived. Acknowledgments I am indebted to Gary S. Morgan of the Florida State Museum for making some helpful suggestions that improved this paper. LITERATURE CITED Ilechaseaux. ('., and A. H. Coogan. 1969. Hippuritacca- Systeniatic descriptions, p. N776-817. In R. C. Moore (ed.). Treatise on Invertebrate Paleontology, Part N, Vol. 2, Mollusca 6, Bivalvia. Univ. Kansas Press, Lawrence. Jones, D. S., and D. Nicol. Origination, survivorship, and extinction of rudist taxa. Jour. Paleo. 60:107-115. Kauffman. E. ti.. and N. F. Sohl. 1974. Structure and evolu- Vol. 100(2) April 30, 1986 THE NAUTILUS 71 tion of Antillean Cretaceous rudist frameworks. Verhand- lungen Naturfor.'ichende Gesellschaft. Basel. 84:399-467. Palmer, A. R. 198,3. The decade of North American geologT,- 1983 geologic time scale. Geology ll:.503-504. Perkins. B. F. 1969. Rudist morphology' -Shell size ami shape, p. N7.")l. In R. C. Moore (ed.). Trcntixc on hirci-tr- bmte Paleontology, Part N, Vol. 2, Mollusca 6, Bivalvia. Univ. Kansas Press, Lawrence. Zoological Record. 1966-1981. The Zoological Society of London. A NEW SOMATOGYRUS FROM THE SOUTHWESTERN OZARKS WITH A BRIEF REVIEW OF THE HYDROBHDAE FROM THE INTERIOR HIGHLANDS (GASTROPODA: PROSOBRANCHIA) Mark E. Gordon Department of Zoology University of Arkansas Fayette vi"lle, AR 72701 ABSTRACT A previously unknown Somatogyrus is described from the Elk River, McDoridld County, Missouri. With this description, the number of Hydrobiidae presently recorded from the Interior Highlands increases to 15, including 11 regional endemics. Nine species are endemic to the Ozark Plateaus, while two Somatogyrus are restricted to the Ouachita Mountains. Fire of the Ozark endemics are obligate troglohites. Troglobitic Amnicola antroecetes (Hubrichf) is considered distinct from epigean A. aldrichi (Call and Beecher). Excepting the Unionoidea, little study has been afforded the aquatic Mollusca of the In- terior Highlands. Among the poorer known groups, regional accounts of the Hydrobiidae are rare. The primary sources of pulilished records have been original descriptions of several endemic species (e.g. Walker, 1915; Hinkley, 1915; Hubricht, 1979), although the oc- casional collection of hydrobiids has been noted in a few species surveys (e.g. Sampson, 1913; Wheeler, 1918; Gordon, 1981). A recent faunal survey of drainage basins in southwestern Missouri and northwestern Arkansas (Gordon, 1980) recovered several species of Hydrobiidae. A previously unknown Somatogyrus. collected from the Elk River, McDonald County, Missouri, is here described. Family Hydrobiidae Troschel 1857 Subfamily Lithoglyphinae Fischer 1885 Somatogyrus rosewateri new species Shell descripfiori of holotype-SheW sub- globose, medium size for Somatogyrus. solid, dextral, 4.25 whorls, turbinate, subhyaline; body whorl .90 times axial height, greatly inflated; [penultimate whorl .34 times width of body whorl (Fig. 1; see Table 1 for holotype measurements and range of paratypes). Spire very short, depressed, broadly conic; sutures moderately impressed; protoconch flattened (Fig. 2). Periostracum greenish (may be masked by color of visceral mass showing through shell) with shallow, oblique growth striae; protoconch or- namented with fine, spiral ridges (Fig. 3). Aper- ture broadly ovate, .74 times axial height, .80 times height and .55 times width of body whorl, plane of aperture at 30° to shell axis. Peristome complete across parietal wall by a callus which continues and thickens across columella (callus in some individuals may be slightly reflected over columella, partially obscuring umbilicus); lip straight along parietal wall, curving con- cavely near the umbilicus and along columella, parietal-columellar junction non-angular. In lateral view (Fig. 4), periphery of parietal lip 72 THE NAUTILUS April 30, 1986 Vol. 100 (2) FIGS. 1-5. Suiniit(i(j!irus nuiruvlcn n. .sp. 1, apei'tural view (if paratype (3.9 x 3.8 mm.); 2, juvenile paratype; 3, suture and pr<)t(jconeh sculpture iif paratype. The liarely visible white bar in the upper left is 10 micrcms in length. 4, lateral view of paratype (4.2 x 3.8 nun.); 5, urnhilical view (if paratype (shell, 3.fi x 3.5 mm.). approximately parallel to plane of aperture, col- umella straight but recedes towards base, and parietal-columellar junction forma an obtuse angle. Umbilicus relativi'ly open for Somnfo- gyrus (Fig. 5); passes into a shallow, canal-like depression continguous to the columellar Iif). Basal lip slightly receded due to angle of aper- ture; area of base-columcllar junction projects forward about ecjual to the distance of the basal lip recession (Fig. 4). Operculum chitinous, ovate, horn-colored; paucispiral, consisting of about three whorls; nucleus subcentral, located slightly left of midline in lower third; outer surface sculptured with many fine growth striae, peripheral margin without striae and edge appears slightly ragged (Fig. 6). Soft paW.s- Living animals translucent (odon- tophore complex visible through proboscis), whitish. Mantle collar and dorsal aspects of pro- boscis, head, tentacles, and foot speckled with sub-epidermal gold to orangish gold chromato- cytes (Fig. 7); dorsal surface and V3 thickness of visceral mass infused with golden color. Dense concentrations of melanin at base of tentacles, extends dorso-medially in front of eyes a short distance and posteriorly on to dorsum of head (Fig. 8); a narrow line of melanin runs laterally from in front of eye forward about V4 to Vs length of tentacle; dorsal and lateral surfaces of mantle with mottled patterns of melanin (Fig. 7), 35% of paratypes (n = 52) lacked mantle melanin. Shells of living snails appear amber to orangish due to visceral and mantle pigmenta- tion showing through shell, darkens to brown in spire; melanistic snails have slightly browner tint. Penis unpigmented. Preserved animal white. Foot broad (Fig. 7); anterior truncated, ex- tending forward to eyes; posterior tapers to a rounded point. Head (Fig. 8) extends anterior of shell to expose eyes; proboscis relatively long and slender, curved ventrally; tentacles long and slender, eyes positioned ventro-laterally at base. Penis (Fig. 9) slender, simple; tapers to a fine, blunt point; dorso-ventrally flattened, dor- sum slightly convex, venter flatly concave; vas deferens positioned near posterior margin. Penis arises a short distance inside mantle cavi- ty above and behind the right tentacle, tends to curve counter clockwise; when extruded, emerges from shell directly behind right ten- tacle and extends below it (Fig. 8). Type locality -M\ssour\, McDonald County, Elk River at Missouri Department of Conserva- tion Mt. Shira Public Fishing Access, 1.6 km south of Ginger Blue (R33W, T21N, sections 1 and 2). Holotype: University of Colorado Museum of Zool()g>' (UCM) no. 32812; collected 24 October 1985 by Mark E. Gordon. Allotype: UCM no. 32813; same data as holotype. Para- types: ten specimens each deposited at UCM, U.S. National Museum of Natural History, Academy of Natural Sciences of Philadelphia, Vol. 100(2) A|:)ril 80, 1986 THE NAUTILUS 73 TABLE 1. Dimensions of holotype, allotype, and paratypes oi' SumiifiHii/nis niKPimferi (measurements in mm): H = height, W= width, BH = body whorl height, PW = penultimate whorl width, A = aperture, AH = aperture height, AW = aperture width. H W W/H BH BH/H W/BH PW PW/W AH AW AW/AH Plane of A Holotype (9i J. 9 i.8 .97 3,5 90 1.09 1. 3 .34 2.8 2.1 75 30° Allotype (o-) 3.4 3.4 1.00 3.1 91 1.10 1.0 .29 2.5 1.9 76 Paratype 3.7 3.6 .97 3.2 86 1.13 1.2 .33 2.3 2.1 91 „„ 4.0 3.7 .93 3.5 88 1.06 1.2 .32 2.7 1.7 63 34° „„ 4.1 3.9 .95 3.6 88 1.08 1.2 .31 2.8 2.4 86 29° „„ 4.0 3.8 .95 3.6 90 1.06 1.1 .29 2.9 2.2 /6 tin 3.9 3.7 .95 3.4 87 1.09 i.2 .32 2.5 2.1 84 34° II It 3.8 3.8 l.OU 3.1 89 1.12 1.3 .34 2.4 2.1 88 35° „„ 3.8 3.5 .92 3.4 89 1.03 1.3 .3/ 2.5 2.1 84 30° (Fig. 1) 3.9 3.8 .97 3.5 90 1.09 1.1 .29 2.5 2.1 84 30° (Fig. 3,4) 4.2 3.8 .90 J. 7 88 1.03 1.3 .34 2.8 2.2 79 30° (Fig. 5) 3.6 3.5 .97 3.3 92 1.06 1.0 .29 2.5 1.9 76 27° FIGS. 6-9. Somatogyrus rosewateri n. sp. 6, operculum; 7, animal extruded from shell (composite drawing from live and rela,\ed specimens; body slightly distended from relaxa- tion with sodium pentobarbital): C = chromatocytes, E = eye, F = foot, M = melanin. Mm = mantle melanin showing through shell. Mo = mouth, 0 = operculum, P = proboscis, T = tentacle; 8, dorsal view of head: Pn = penis; 9, penis: V = vas deferens. Museum of Comparative Zoology, University of Michigan Museum of Zoology, and FieW Museum; same data as holotype. Distribution -Found only in the middle sec- tion of Elk River (Fig. 10). To date, this species has been collected from only the type locality and below an old mill dam at Noel, McDonald County, Missouri. It was not recovered from 74 THE NAUTILUS April 30, 1986 Vol. 100(2) ARIANSAS FKi. Kl. Distriliution of Siini.atofiyrus riiaeiratcri (sdlid triangle is the type locality). areas upstream of the type locality nor the river channel downstream from the Missouri- Oklahoma border which has been inundated by a reservoir on the Neosho River (Grand Lake 0' the Cherokees). The known range of this species occurs in the Springfield Plateau region of the Ozarks. //rr6;7a/- Specimens were collected at the type locality from a series of riffles. Substrate was composed of gravel and cobble-sized materials (mainly chert). At Noel, habitat was below the outfall of a small mill dam. Substrates were primarily stepped and broken bedrock with some cobble. Gradients at both sites were fairly high and water quality was quite good (Table 2). Efyyuology -Th\i^ s|)ecies is named in memory of Dr. Joseph Rosewater, late Curator of Mollusks, U.S. National Museum of Natural History. Discussion The comparative difference between S. rosewateri and previously described species in the genus are shown in Table 3. TABLE 2. Physicd-cheniical nieasurenierits madi locality. Elk River; 20 November, 1979. at type Water temperature Tiirliiility Specific coruiuctance Alkalinity I'll Nitrate Color 15°C 1.6 NTU 2(j0 fjmhos/cm 140 mfT/l total CaCUj 8.1.-) 1..') mg/l neglij^ble Besides Somatoyyru,^ rosewateri. the only other hydrobioids collected from the Elk River basin were the nymphophiline Cincinnatia in- tegra (Say) and the pomatiopsid Pomatiopsi>i lapidaria (Say) (Gordon, 1980). Two other Somatogyrus are known from the Ozark Plateaux. Somatogyrus depressus (Tryon), an in- habitant of the upper Mississippi River valley, occurs in the Osage River, central Missouri (Thompson, 1984) and S. crassilahris Walker, an Ozark endemic, is apparently known only from its original collection from the North Fork of the White River, Arkansas. In discussing phylogenetic relationships among North Ameri- can Lithoglyphinae, Thompson (1984) noted dif- ficulties in pursuing such studies due to a relatively large number of undescribed species and a paucity of morphological data for de- scribed species. For Somatogyrus. he did list several characters useful for specific com- parisons. Available data for the three Ozark species and two Ouachita Mountains endemics (Table 3) suggests that the Ozark species may be fairly closely related. Walker (1915) also con- sidered S. wheeler i Walker closer to S. crassilahris than to its Ouachitan congener. However, both Ouachita Mountains endemics may have evolved from other lineages (e.g. Alabama River). Considerably more information is necessary before such speculative, phylogenetic relationships may be confirmed. In some instances, complete analyses may no longer be possible. Many hydrobiids, such as S. era.ssilabrls. S. amnicoloides Walker, and S. wheeleri. are known only from their type localities. Habitat alterations, such as reservoir construction, may have been deleterious for species with such highly restricted distributions. Fifteen species of Hydrobiidae have been documented from the Interior Highlands (Table 4). Four are wide-ranging species. In the case of Prohythvnella lacu.'^tris (Baker) (mistakenly listed as F. binneyana (Hannibal) in Gordon, 1981) and Somutogyrus depres.' 4 mm) ( 1) Spire depressed (>15% H)(0) 0 elevated (<15% H) (1) Protoconch flattened (0) 0 elevated (1) Par ietal-colume liar wall junction of lip smooth curve ( 0 ) 0 angular (1) Umbilicus narrow { 0 ) 0 open ( 1 ) Widest expansion of outer lip periphery above mid-height (0) 0 below mid-height (1) Aperture apex relation to parietal wall fused (0) 0 detached (1) .77-. 92 .90-1.00 .93 amnicoloides and S. ivheeleri are presently the only endemic hydrobiids that have been iden- tified from the Ouachita Mountains province. Four epigean and five troglobitic species are endemic to the Ozark Plateaus. The maj(jrity (eight) are restricted to the eastern half of the province. The only endemic, western Ozark hydrobiid presently recognized is S. ro^tewaferi: however, a single specimen of Somntogyrus has been collected from the headwaters area of the White River, Arkansas (Gordon, 1980). This specimen was too eroded to allow identification beyond genus and sulisequent collecting failed to recover additional specimens. Most of the Ozark endemic, obligate troglobi- tic Hydrobiidae have been referred to Amnicoli- nae: Anniicoln. Burch and Tottenham (1980) noted that this classification was tentative due to the lack of anatomical flata for these species. However, Antrobia culreri Hubricht was rele- gated to the Lithoglyphinae without justifi- cation by Burch and Tottenham (1980). The anatomy of this species remains all but unknown. The brief description of the radula presented by Hubricht (1971) does not appear to meet the radular criterion for Lithoglyphinae established by Thom()son (1984) and differs from that of Amniroln only by the size of the central tooth mesocone. Hubricht (1971) described the penis of Antrohid as "simple, tapering to a point, without appendages." This morphology is characteristic of the lithoglyphine penis (Thompson, 1984) and is prolialily the basis for the classification in Burch and Totten- ham (1980). Unfortunately, Hubricht (1971) did not describe the number of ducts penetrating the (lenis. Thompson (1984) considered the Amnicolinae, with two penial ducts, remote in 76 THE NAUTILUS April 30, 1986 Vol. 100(2) TABLE 4. Hydrobiidae known from the Interior Highlands. Hydrobiinae Prohythinctta iacu.stritt {Baker, 1928) Lithoglyphinae Som tations to be ecophenotypic variation implies an amazing genetic plasticity and an "use it or lose it" evolutionary interpretation. Ecophenotypic variation does not meet modern criteria for sub- specific status (Mayr, 1966); however, previous relegation of nntroeretes as a subspecies reflects the over-reliance on shell characters for con- structing phylogenies. Amnicola antroecefes should be considered a distinct species. In- terestingly, A. iintriieceteii has a rather wide distribution when compared to other regional troglobitic gastropods (restricted to single or connected caves). Other troglobitic taxa (e.g. Amblyopsidae; Isopoda: Caecidotea) exhibit similarly wide ranges in the Interior Highlands. This may be related to the karst geology of the region. Hubricht (1940) did note some size varia- tion at different localities liut correlated this to food availability. However, due to its wide range he suggested that .4. antn)ecefe>< may represent a composite group. Further investigation is ob- viously warranted for the A. aldrichi complex and, for that matter, the Hydrobiidae through- out the Interior Highlands. Acknowledgments The uniqueness of Somatogyrus roi^ewnteri was confirmed by Fred G. Thompson, Florida State Museum, University of Florida. SEM's were prepared with the assistance of Claudia S. Bailey and specimens o{ Amnicola aldrichi were provided by Michael L. Mathis, both Depart- ment of Zoology, University of Arkansas. LITERATURE CITED Baker. F. C. 1928, The fresh-water Mollusca of Wisconsin, part 1: Gastropoda. BulL Wiitc. Geol. Nat. Hist. Sun: 70:1-507. Burch. J. B. and J. L. Tottenham. 1980. North American Ireshvvatcr snails; species Hst. runnes, and illustratinns. W.ilbrniir, \:S]--l\r,. Clarke, A. 11. 197M. The freshwatei' molluscs of the Cana 26.5 mm.). 11 and 12, paratype Din the Donald Pisor collection (26.6x27.2 mm.; areas with moderately raised cords; varices ex- tending over shoulder obliquely to left and ter- minating at suture. Spiral sculpture as follows: one broad cord at the shoulder prolonged prominently into a blunt spine, the margin nearly closed on the leading side; a similar broad cord below extending beyond the varix, but not prolonged into a spine, up to the penultimate varix. On the mature apertural varix these two cords do not project beyond the remaining sculpture of the varix. Four secondary cords on the body whorl dimin- ishing in size anteriorly; varix deeply excavated between the cords on receding side; cords on body whorl more or less weakly raised; three minor cords on shoulder obsolete except near apertural varix. Leading side of varices re- curved and with scabrous lamellae producing a surface with squarish pits; weak scabrous lamellae on surface of spiral cords between varices. Color: Creamy white entirely except for the holotype with is a dead collected shell lightly stained with rust. Type Locality -Ma.\si\t2. Island, Solomon Islands. Dimensions- LenKth width Holotype SDNHM 8B944 .Ataa, Malaita Island, Solomon Is. 2t;.l) 21) 5 mm Paratype A, SDNHM 86942 Marau Sound, Guadalcanal, Solomon Is. 17,0 1,5,',) I'aratype B, ANSP 289.'JBK Malaita Island, Solomon Islands 2(1, .5 20,2 82 THE NAUTILUS April 30, 1986 Vol. 100(2) KKjS. lIMti. 13, Outline drawinn' 95%) of aragonite. Soft-parts Morphology External features: The soft-parts comprise 3 to 3V3 whorls, of which the digestive gland (Fig. 7, dg) occupies about 2V4 whorls, the kidney (Fig. 7, k) less than Vs whorl and the mantle cavi- ty just over V2 whorl. Preserved animals were orange tan in color, lacked any discernible markings and were retracted at most 'A whorl into the aperture. The foot is broad and ovate, lacking an operculum. Tentacles (Fig. 7, t) are long, tubular and symmetrical. The mantle edge is smooth, the siphon (Fig. 7, s) short but distinct. Mantle rarity: The mantle cavity is shorter and broader than in other cancellariids dis- sected, with pallial organs situated as in other higher prosobranchs. The osphradium (Fig. 7, os) is broad anteriorly (L/W = 3), tapers posteriorly, and consists of about 35 leaflets per side. Adjacent is the ctendium (Fig. 7, ct), twice as long and slightly narrower than the osphradium, composed of about 70 triangular leaflets. This organ and the voluminous, trans- versely pleated hypobranchial gland (Fig. 7, hg) Vol. 100(3) July 30, 1986 THE NAUTILUS 87 FIGS. 1-6. Features of the shells and jaw of Adniete viridula (Fabricius). 1, Apertural and right side views of specimen collected in the Saguenay River, Quebec. [48°24'48"N, 70°44'-70°48'W] sta. 62 30 (NMNS 43946) 3.0 x . 2, Scanning elec- tron micrograph of juvenile specimen dredged in 673 m, off Martha's Vmeyard, Massachussetts, U.S. Fish. Comm. sta. 994-97. (USNM 43232) 20.0 x . 3, Protoconch of specimen in figure 2, lateral view, scale bar = 200 ^jm. 4, Protoconch of specimen in figure 2, axial view, scale bar = 200 /jm. 5, Fracture surface. Plane of fracture parallel to outer lip, scale bar = 100 fjm. 6, Lateral view of jaw. scale bar = 50 jjm. 88 THE NAUTILUS Vol. 100(3) vd pap rmc FIGS. 7-10. Anatomical features of Admete oiridula. 7, Female specimen removed from shell, partially un- coiled and mantle cavity opened mid-dorsally to display contents. 8, Dissection of anterior portion of proboscis, opened mid-dorsally. 9, Diagrammatic representation of the alimentary system. 10, Male reproductive system. a, anus; ag, albumen gland; asg, accessory salivary gland; be, bursa copulatrix; bg, buccal ganglia; bm, buccal mass; eg, capsule gland; ct, ctenidium; dg, digestive gland; hg, hypobranchial gland; int, intestine; j, jaw; k, kidney; m, mouth; me, mid-esophagus; ng, nephridial gland; nr, nerve ring; opm, opening to mantle cavity; os, osphradium; ot, oral tube; pap, papilla; pc, pericardium; pen, penis; pr, prostate gland; r, rectum; rmc, rear of mantle cavity; s, siphon; sg, salivary gland; sto, stomach; sv, seminal vesicle; t, tentacle; tes, testes; vd, vas deferens; vl, valve of Leiblein. span the roof of the mantle cavity. The pallial gonoducts and rectum (Fig. 7, r) are located along the right side of the mantle cavity, while its rear is formed by the pericardium (Fig. 7, pc) and kidney (Fig. 7, k). The kidney is small, being only twice the size of the pericardium. The nephridial gland (Fig. 7, ng) occupies nearly half the kidney along the pericardium. Alimentary system: The broad, tubular pleu- rombolic proboscis can be extended nearly the length of the shell aperture, and is the same color as the foot. Broad retractor muscles ex- tend from the columellar muscle, while thinner strands originate from the body wall. A short oral tube (Fig. 8, ot) extends from the mouth (Fig. 8, m) to the cuticularized, tubular portion of the jaw (Fig. 8, j), which expands posteriorly to envelope the lateral surfaces of the minute buccal mass (Fig. 8, bm). The longitudinally grooved buccal mass lacks a radula and has only a vestigial subradular membrane. A pair of short, extremely thin accessory salivary glands (Fig. 8, asg) lie alongside the buccal mass, emptying into the posterior portion of the oral tube through fine ducts. The wider and much longer salivary glands (Fig. 8, sg) join the buccal mass laterally, via short ducts, just anterior to the esophageal opening. A small valve of Leiblein (Fig. 8, vl) and a pair of buccal ganglia (Fig. 8, bg) are situated at the rear of the bucal mass. The narrow mid-esophagus (Figs. 8, 9, me) runs posteriorly from the valve of Leiblein, passing through the nerve ring (Fig. 9, nr) before expanding and becoming sacular. The esophagus narrows at the rear of the cephalic sinus and leads to a simple U-shaped stomach (Fig. 7, 9, sto) that is embedded in the anterior face of the digestive gland. Longitudinal folds Vol. 100(3) July 30, 1986 THE NAUTILUS 89 TABLE 1. Admete rindutn (Fabricius), measurements of shell characters. Linear measurements in millimeters, (n = 10). Standard Character Mean Deviation Range shell length 16.87 1.91 13.82-19.02 shell width 9.90 1.13 8.18-10.97 aperture length 10.21 1.42 8.13-11.58 aperture length 0.604 0.023 0.567-0.641 shell length #whorls, teleoconch 4.78 0.35 4.33-5.33 spire angle .5.5.2° 2.9° 45.5°-.58.0° line the stomach and pass into the intestine (Fig. 9, int), which runs through the kidney and along the right wall of the mantle cavity, expanding slightly to form the rectum (Figs. 7, 9, r). Neither an anal gland nor identifiable gut con- tents were detected in any of the specimens ex- amined. Female reproductive system: As the only female specimen available to us was poorly pre- served, only the pallial portion of the oviduct is described. The oviduct passes through the kidney and expands to form a broad, laterally compressed albumen gland (Fig. 7, ag), which expands further to form the long, blunt, capsule gland (Fig. 7, eg). A small, ovate, bursa copula- trix (Fig. 7, be) is situated below the anterior margin of the capsule gland. An ingesting gland was not found. Egg capsules oi Admete viridula are figured by Bouchet and Waren (1985: fig. 687). Male reproductive system: The testes (Fig. 10, te) consist of a yellowish mass of ascinous tubules embedded in the right side of the diges- tive gland and extend nearly a full whorl from its anterior edge. These tubules converge to form a testicular duct that immediately forms a serpentine seminal vesicle (Fig. 10, sv). The duct straightens at the anterior limit of the digestive gland, passes through the kidney and enters the mantle cavity ventral to the rectum. It greatly expands to form the prostate gland (Fig. 10, pr), which extends along the posterior third of the mantle cavity and communicates with it by a long ventral slit (Fig. 10, opm). From the anterior portion of the prostate gland, a tubular vas deferens (Fig. 10, vd) leads to the base of the penis (Fig. 10, pen), which is ovate in cross- section, extends approximately % the length of the mantle cavity and has a terminal papilla (Fig. 10, pap). Nervous system: Admete viridula has a typical cancellariid nervous system (Bouvier, 1887: Harasewych and Petit, 1982, 1984), with the cerebral, pedal and pleural and subesophageal ganglia fused into a nerve ring, and the buccal ganglia (Fig. 8, bg) at the rear of the buccal mass. Discussion Cladistic analysis of the 20 shell and anatomical characters listed in Table 2 produced the cladogram in Figure 11. Despite the lack of agreement on the relationships of supraspecific taxa, nearly all classifications proposed to date separate the Admetinae from the remaining Cancellariidae (H. and A. Adams, 1853: Troschel, 1865; Fischer, 1883; Tryon, 1885; Cossmann, 1899; Thiele, 1929; Wenz, 1943), and this arrangement is supported by the present study. Of the characters used to distinguish the Admetinae from other cancellariids, most con- TABLE 2. Shell and anatomical characters used to distin- guish between Admete viridula, Cancellaria reticulata and Olssonella smithii. Primitive states denoted by (0), derived states by (1). 1) Shell: (0) thick: (1) thin. 2) Internal varices: (0) pronounced; (1) reduced or absent. 3) Shell ultrastructure of: (0) 2 orthogonal layers; (1) 1 layer. 4) Protoconch of: (0) 2 whorls; (1) 1 whorl. 5) Tentacles: (0) symmetrical, unmodified; (1) asymmetri- cal, left tentacle modified. 6) Osphradium with approximately: (0) 35 leaflets/side; (1) 70 leaflets/side. 7) Ctenidium with: (0) < 125 leaflets; (1) > 200 leaflets. 8) Distance between osphradium and ctenidium: (0) nor- mal; (1) large relative to either organ. 9) Kidney occupying; (0) > 'A whorl; (1) < Vs whorl. 10) Proboscis: (0) tubular; (1) ventrally flattened, papillose. 11) Jaws with: (0) short posterior lobes; (1) long posterior lobes. 12) Buccal mass: (0) small; (1) large, filling retracted pro- boscis. 13) Radula: (0) present; (1) absent. 14) Outer cusps of radular teeth: (0) simple, smooth; (1) with secondary dentition. 15) Accessory salivary gland: (0) shorter than; (1) longer than; salivary gland. 16) Mid-esophagus posterior to nerve ring: (0| sacular; (1) convoluted tube. 17) Anal gland: (0) present; (1) absent. 18) Sperm ingesting gland: (0) present; (1) absent. 19) Albumen gland: (0) dorsally recurved; (1) laterally com- pressed. 20) Prostate gland: (0) long, expanded; (1) short, con- voluted. 90 THE NAUTILUS July 30, 1986 Vol. 100 (3) 3 •o > CO w 3 o o CJ E FIG. 11. Cladogram of phylogenetic relationships of cancel- lariid taxa. Single slashes across tree branches represent transformations of the corresponding character from the primitive (0) to the derived (1) state. Double slashes indicate transformations that occur more than once. spicuous are their thin shells lacking pro- nounced surface sculpture and apertural lirae. This is likely a consequence of their polar habitat, as Graus (1974) has shown that de- creased calcification in high latitudes is due to reduced availability of calcium carbonate in colder waters. Admetines also lack such anti- predatory features as internal varices that are found in tropical and temperate cancellariids. The predominance of aragonite in the shell is plesiomorphic, as calcite is more easily formed (Wilbur, 1964) and more stable (Lowenstam, 1954) at low temperatures. In terms of anatomical organization, Admete viridula differs from ancestral cancellariid mor- phology, most nearly approximated by Olsso- nella snulhii of the taxa studied to date, in hav- ing a shallower mantle cavity, a reduced kidney, and in lacking color pattern. At least some of these may also be adaptations to cold water. Cancellariines and Trigonostomines have uni- serial "tricusped" radular teeth with elaborate secondary dentition on the outer cusps and a comparatively simple ventrally recurved central cusp (Harasewych and Petit, 1982, 1984; Petit and Harasewych, 1986). The radula of Notho- admete tumida, the only known radulate admetine, has barbed central cusps and simple bulbous outer cusps (Oliver, 1982: figs. 3, 5). Presence of central cusp barbs in Cancellaria atopodonta Petit and Harasewych (1986: figs. 15, 16) suggests that this feature is primitive and occurred in the ancestor of all Recent cancellariids. The jaws of admetines lack the long posterior lobes found in cancellariines and trigonostomines. Although the diet of cancel- lariids remains unknown, we have suggested that they are piercing suctorial feeders based on the functional morphology of their alimentary systems (Harasewych and Petit, 1982, 1984; Petit and Harasewych, 1986). The loss of radula and reduction in the size of the buccal mass in most admetines suggests that they feed suc- torially, but on a different group of prey organisms than other cancellariids. The lack of an anal gland in Admete viridula and Cancellaria reticulata is likely a com- paratively recent convergence, as this organ has been reported in Cancellaria cancellata (Graham, 1966). Additional differences between admetines and other cancellariids are found in the reproductive systems. The prostate gland ofAdynete viridula is long and greatly expanded, while in Olssonella smithii and Cancellaria reticulata it is short and convoluted. Admete viridula also lacks the separate sperm ingesting gland and dorsally recurved albumen gland found in cancellariines and trigonostomines. The phylogenetic arrangement in Figure 1 1 is supported by the fossil record, as several "admetines" have been reported from the Upper Cretaceous, and Waipaoa. known to be aradulate in the Recent fauna, dates back to the Oligocene. Cancellaria, sensu stricto, first ap- pears in the Miocene and is characterized by modifications to the left cephalic tentacle, pro- boscis, osphradium and ctendium, organs in- volved in tactile and distance chemoreception. Acknowledgments We thank Jane B. Topping, Invertebrate Zoology Division, National Museum of Natural Sciences, Ottawa, for making available the preserved specimens used in this study. Some of this work was done at the Smithsonian Marine Station at Link Port, Florida, and we thank Dr. Vol. 100(3) July 30, 1986 THE NAUTILUS 91 Mary Rice for making these facilities available. This is contribution number 158 of the Smith- sonian Marine Station at Link Port. Critical review of the manuscript by Dr. R. S. Houbrick, National Museum of Natural History, Smith- sonian Institution, is gratefully acknowledged. LITERATURE CITED Adams, H. and A. Adams. 1853-58. The Getiera of Recent MoUusca. 3 vols. London. Beu, A. G. and F. M. Climo. 1974. Mollusca from a recent coral community in Palliser Bay, Cook Strait. N. Z. Joum. Marine and Freshwater Research 8(2):307-332. Bouchet, P. and A. Waren. 1985. Revision of the Northeast Atlantic Bathyal and Abyssal Neogastropoda excluding Turridae (Mollusca, Gastropoda). Boll. Malacologico. Suppl. 1:121-296. Bouvier, E. L. 1887. Systeme nerveux. morphologie generale et classification des Gasteropodes prosobranches. Ann. Sci. nat. Zool. 3:1-510. Cossmann, M. 1899. Caneellariidae. Essais de Paleocon- chologie Ccmiparee 3:1-41, pis. 1-2. Dall, W. H. 1887. Supplementary notes on some species of mollusks of the Bering Sea and vicinity. Proc. U.S. Nat. Miis. 9:297-309, pis. 3-4. 1888. [in] Agassiz, A. Three Cruises of the United States Coast and Geodetic Survey Steamer Blake, 2(8): 62-75, figs. 282-312. 1918. Notes on the nomenclature of the mollusks of the family Turridae. Proc. U.S. Nat. Mus. 54:313-333. Dell, R. K. 1956. The Archibenthal Mollusca of New Zealand. Dominion Mus. Bull. 18:1-235, pis. 1-25, A, B. Fabricius, 0. 1780. Fauna Groenlandica. Hafnia et Lipsiae. 452 p. Fischer, P. 1880-1887. Manuel de conchyhologie et de paleontologie conchyliologique ou histoire naturelle des moUusques vivants etfossil.es. Paris, F. Savy. 1369 p. Graham, A. 1966. The fore-gut of some marginellid and can- cellariid prosobranchs. Stud. trop. Oceanogr. Miami 4:134-151. Graus, R. R. 1974. Latitudinal trends in the shell character- istics of marine gastropods. Lethaia 7:303-314. Harasewych, M. G. and R. E. Petit. 1982. Notes on the mor- phology of Cancellaria reticulata (Gastropoda: Caneellari- idae). The Nautilus 96(3):104-113. 1984. Notes on the morphology of Olssonella smithii (Gastropoda: Caneellariidae). The Nautilus 98(l):37-44. .Jay, J. C. 1839. A catalogue of the shells . . . contained in the collection of John C. Jay. M.D. Ed. 3, New York. 125 p., 10 pis. Jeffreys, J. G. 1876. New and peculiar mollusca of the Euli- midae and other families of gastropoda, as well as the pteropoda, procured in the 'Valerous' expedition. Ann. Mag. nat. Hist. 4(19):317-339. Linne, C. von. 1767. Systema naturae per regna tria naturae. Editio duodecimo reformata. Vol. 1 (2): 533-1327. Stockholm. Lowenstam, H. A. 1954. Factors affecting the aragonite: calcite ratios in carbonate secreting marine organisms. J. Geol. 62:284-322. Macpherson, E. 1971. The Marine Molluscs of Arctic Canada. Nat. Mus. Canada, Publ. in Biol. Oceanography, No. 3:1-149. Moller, H. P. C. 1842. Index Molluscorum Groenlandiae. Nat. Tidsskr. 4:76-97. Morse, E. S. 1921. Observations on living gastropods of New England. Peabody Museum, Salem. 29 pp., 9 pis. Oliver, P. G. 1982. A new species of Cancellariid gastropod from Antarctica with a description of the radula. Br. Antarct. Surv. Bull. 57:15-20. Petit, R. E. and M. G. Harasewych. 1986. New Philippine Caneellariidae (Gastropoda: Cancellariacea), with notes on the fine structure and function of the nematoglossan radula. The Veliger 28(4):436-443. Pilsbry, H. A. 1938. On the history and status oi Lora Gistel. The Nautilus 51(4):11.5-118. Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda. Discovery Repts. 26:47-196, pis. 5-10. Strebel, H. 1905. Beitrage zur Kenntnis der Mollusken Fauna der Magalhaen-Provinz. Zool. Jb., Abt. Syst., Jena 22:575-666, pis. 21-24. Thiele, J. 1903 [1904]. Die beschalten Gastropoden der deutschen Tiefsee-Expedition 1898-1899. B. Anatomisch- systematische Untersuchungen einiger Gastropoden. Deutsche Tiefsee-Exped.. Bd. VII:149-179, Taf. 1-9. 1929-35. Handbuch der system.ati.'ichen Weich- tierkunde. Jena, Gustave Fischer. 1154 p. Troschel, F. H. 1856-1893. Das Gebiss der Schnecken. zur Begriindung einer natilrlichen Classification. 2. Berlin. Tryon, G. W. 1885. Family Caneellariidae. Manual ofCon- chology 7:65-98, pis. 1-7. Wenz, W. 1938-1943. Handbuch der Paldozoologie (0. H. Schindewolf, ed.), Berlin, Band 6, Teil 1 (pt. 2) 1639 p. Wilbur, K. M. 1964. Shell formation and regeneration [in] Physiology of Mollusca, vol. 1, Wilbur, K. M. and C. M. Yonge eds. Academic Press, New York. BOOK REVIEW Washington Public Shore Guide: Marine Waters by James W. Scott and Melly A. Reuling. 1986. 348 pp., numerous photos, maps and habitat sketches. Cloth, $25.00; paperback, $14.95. If you are planning on studying or visiting the shorelines of the State of Washington, this is a great bargain with many useful facts about the 2,400-mile saltwater shoreline of 14 counties. It largely ignores mollusks, a fact which offers a challenge to conchologists.-7?. T. Abbott. 92 THE NAUTILUS July 30, 1986 Vol. 100(3) OBSERVATIONS ON THE FEEDING BEHAVIOR OF THE GASTROPOD PLEUROPLOCA PRINCEPS (FASCIOLARIIDAE) IN THE GALAPAGOS ISLANDS Ian Stupakoff Department of Invertebrates American Museum of Natural History Central Park West at 79th Street New York, NY 10024 ABSTRACT The feeding behavior of the gastropod. Pleuroploca princeps, (Fasciolariidae) is reported. Eight species of gastropods were found to be preyed upon by P. princeps; two species of crabs were eaten as carrion. No signs of cannibalism were noticed, but intraspecific competition was observed. While working at the marine laboratory of the Charles Darwin Research Station, Isla Santa Cruz, Galapagos, Ecuador, I had the opportuni- ty to collect the gastropod, Pleuroploca princeps (Sowerby, 1825), (Keen, 1971, p. 611, fig. 1324), and to study its feeding habits. The only literature citation on the feeding habits of P. princeps is by Sorensen (1943, p. 4), who observed P. princeps feeding on Phyllono- tus bicolor (now Hexaplex erythrostomus (Swain- son)) in Bahia San Carlos, Gulf of California. Other observations on feeding habits are reported for P. gigantea (Kiener) in the western Atlantic by Menzel and Nichy (1958, p. 144), Paine (1963 a, pp. 66, 67; 1963 b, pp. 402, 403), and Cornellisson (1985, p. 125), and on P. trapezium (Linnaeus) and P. filamentosa (Roding) in the Indo-Pacific by Risbec (1932, pp. 374, 375), Maes (1967, p. 51), and Reyes (1981, p. 4). The major prey items in the diet of members of the family Fasciolariidae are gas- tropods, bivalves, sedentary polychaetes, cirri- pedes and carrion (Taylor, Morris and Taylor, 1980, p. 377, table 1). Pleuroploca princeps is a characteristic faunal constituent of the Panamanian province and is one of the largest gastropods in the Colon Archipelago (Wellington, 1975, p. 59), attaining 250 mm. in length. This species is widely distri- buted in the Galapagos Islands. Several specimens were collected or observed around the islands of Santa Fe, Santa Cruz, Espahola, Floreana, Bartolome, Cousins Rock and San Cristobal. Living specimens were noted in the subtidal zone and to depths of 21 meters (per- sonal observation). They were found on lava rocks or on sandy bottoms but occur more fre- quently in areas where the lava boulders meet the sandy bottom. The specimens of P. princeps used in this study were collected on lava boulders and sand at Isla Santa Fe (July, 1984) and Isla Santa Cruz (Oct. and Nov. 1984). The animals were kept in aquaria at the marine laboratory. Feeding Behavior Since P. princeps is a typical neogastropod, it feeds mainly on other gastropods (Table 1). Dur- ing these observations no bivalves were offered as food. In aquaria, P. princeps were observed to feed on dead crabs such as Grapsus grapsus (Linnaeus) and Petrolisthes edwardsii (Saussure), (see Table 1 and Fig. 2). A specimen of P. princeps tried several times to attack a Nerita .^cabricosta (Lamarck) but could not reach the soft body of the prey with its proboscis because of the protection of the heavily calcified operculum of the nerite. When P. princeps was at rest, the body and half of the shell was buried in the sandy substra- tum (personal observation). It was at this resting time that five specimens of Thais melones (Duclos) were introduced into the aquarium as food. By using what are thought to be chemoreceptors (fide Kohn, 1961), the predator detected the presence of the prey and began its search. The eyes of P. princeps are poorly developed Vol. 100(3) July 30, 1986 THE NAUTILUS 93 TABLE 1. Species eaten by Plcurophca princeps in aquarium. Phylum Class Faml ly Species MoIIusca Gastropoda Cymati idae Cymatium partheopeum Thaididae Thais melones Purpura columellaris Purpura pansa Muricidae Muricanthus princeps Tonnldae Ma lea r ingens (pers. obs. J. de Roy) Cypraeidae Cypraea cervinetta Conidae Conus brunneus Arthropoda Crustacea Grapsidae Grapsus grapsus Porcellanidae Petrolisthes edwardsii and probably cannot visually detect prey. Conse- quently, this predator must feel its way to the prey with its massive foot (Fig. A). On contact, P. princeps wraps the prey in its foot and begins a series of usually two or three muscular con- tractions as observed in 20 cases. Raising its shell slowly and then lowering it very quickly against its prey (Figs. B and C), the predator succeeds in making the prey withdraw inside its shell. When the prey has withdrawn, the shell is maneuvered until the aperture is positioned up- ward and close to the predator's proboscis. At the same time, the predator's muscular foot closes the prey's siphonal canal which may last from a few minutes up to two days depending on the size, shape and strength of the prey (arrow in Fig. E). Figure 1 shows how P. princeps envelopes a Cymatium. partheyiopeum (von Salis) by the same method described above. When this series of movements is terminated, P. princeps prys up the prey's operculum with its proboscis, eventually penetrating the prey's body and devouring the soft parts (Fig. E). The whole animal is consumed and the muscular tissue attached to the operculum is rasped clean by radular action. Similar feeding habits were described for Fasciolaria hunteria (Perry), (Wells, 1958, p. 155) and for Pleuroploca gigantea (Paine, 1963 a, p. 67). Competition and Cannibalism In one of the aquaria, two adult specimens of P. princeps were deprived of food for a period of two weeks. During this time no signs of attack or cannibalism were noticed. No references to cannibalism by Pleuroploca have been located in the literature. In contrast, Fasciolaria tulipa (Linnaeus) (Snyder & Snyder, 1971, p. 257) and F. hunteria (Wells, 1958, p. 155) will cannibalize even when other sources of food are available. Nevertheless, after this period of starvation one specimen of T. melones was offered as food to these two specimens. At this moment, competi- tion was observed between the two P. princeps. The specimen (approx. 100 mm.) that had enveloped the prey was then attacked by the other larger and heavier specimen (approx. 125 mm.) which "unwrapped" the prey and removed it from its first predator in a period of two minutes. The smaller predator tried to recover its prey four times without success. Acknowledgments I thank the following people for their comments on the manuscript: Dr. William K. Emerson and Mr. Walter E. Sage, III, of the American Museum of Natural History, Dr. Jan Pechenick of Tufts University, and Dr. Ruth D. Turner of Harvard's Museum of Comparative Zoology, and to Dr. R. Tucker Abbott for editing this document. I am very grateful to Dr. Gunther Reck, Director of the Charles Darwin Research Station, for the wonderful opportunity to work on the Galapagos Islands. Support in 94 THE NAUTILUS July 30, 1986 Vol. 100 (3) FIG. 1, Adult Pleuroploca prinreps (.shell approx. 12r) mm.) ciivflciping a live Cynuitium parthenopeum and closing the prey's siphonal canal with its muscular foot (see arrow); 2, Adult Pleuroploca princeps feeding on a dead Porcelain crab. Petrolitkes edwardsii (natural size). Vol. 100(3) July 30, 1986 THE NAUTILUS 95 part was received by the Undergraduate- Graduate Research Program of the American Museum of Natural History supported by the Greenwall Foundation. LITERATURE CITED Cornellisson, H. 1985. Florida Shelling-Perseverance pays off! Texas Conchologist 21(4):124-126. Keen, A. M. 1971. Sea Shells of Tropical West America. (Second Ed.), Stanford University Press, Stanford, CA., xiv + 1064 pp. Kohn, A. J. 1961. Chemoreception in Gastropod Molluscs. A7yu;rican Zoologist l(2):291-308. Maes, V. 0. 1967. Radulae of Two Species of Pleuroploca (Fasciolariidae) From The Indo-Pacific. The Nautilus 81(2):48-54. Menzel, R. W. & F. E. Nichy. 1958. Studies of The Distribu- tion And Feeding Habits of Some Oyster Predators in Alli- gator Harbor, Florida. Bull. Mar. Sci. Gulf and Caribbean 8(2):125-145. Paine, R. T. 1963 a. Trophic Relationships of 8 Sympatric Predatory Gastropods. Ecology 44(l):63-73. 1963 b. Feeding Rate of a Predaceous Gastro- pod, Ple^iroploca gigantea. Ecology 44(2):402-403. Reyes, C. W. 1981. The Fascination of Live Shells. Carfel (Manila) 3(5):4, 10. Risbec, J. 1932. Note Sur Les Moeurs de Ricinula chaidea Duel, et de Fasciolaria filamentosa Lmk. Bull. Sac. Zool. Ft. 57:374-375. Snyder, N. F. R, & H. A. Snyder. (1971). Pheromone-Medi- ated Behaviour of Fasfio/aj-ia tulipa. Anim. Behav. 19(2): 257-268. Sorensen, A. (1943). Traveling and Collecting in Mexico. The Nautilus 57(l):l-5. Taylor, J. D., Morris, N. .J, and C. N. Taylor. 1980. Food Specialization and the Evolution of Predatory Proso- branch Gastropods. Palaeontology 23(2):375-409. Wellington, G. M. 1975. Medios Ambientes Marinos Y Costeros de Galapagos. Ms. on file in library of the Charles Darwin Research Station. Isla Santa Cruz, Galapagos, Ecuador. Wells, H. W. 1958. Predation of Pelecypods and Gastropods by Fasciolarta huntena (Perry). Bull. Mar. Sci. Gulf and Caribbean 8(2): 152- 166. FIG. 3 A-E, Sequence of capturing movements of Pleuroploca princeps. 96 THE NAUTILUS July 30, 1986 Vol. 100(3) A NEW SPECIES OF MORUM FROM THE ANDAMAN SEA (GASTROPODA: VOLUTACEA) William K. Emerson Department of Invertebrates American Museum of Natural History New York, NY 10024 ABSTRACT Morum (Oniscidia) ninomiyai, new species, is described from off Thailand in the Andaman Sea and is compared with closely related congeners. In 1981, the late Carl C. Withrow of Florida submitted to me for identification a specimen of an apparently new species of Morum from the Andaman Sea, off southern Thailand. This unique specimen was subsequently illustrated by Sally Diana Kaicher (1983) in her "Card Catalogue of World-Wide Shells", as an ap- parently undescribed Morum. The Andaman Sea specimen was again transmitted to me for study in 1983 by Taizo Ninomiya of Tokyo, who had obtained Mr. Withrow's collection of Morum. Mr. Ninomiya kindly permitted me to retain the specimen with the expectation that additional material might be forthcoming from his contacts in Bangkok. Fortunately, a second specimen resulted from the inquiries of Mr. Ninomiya, for whom I take great pleasure in naming this interesting discovery. The new species is a member of an Indo- Pacific group within the subgenus Oniscidia Morch, 1852, composed of the following taxa: cancellatum (Sowerby, 1824; type species); grande (A. Adams, 1855); uchiyamai Kuroda and Habe, in Habe, \^&\\ joelgreenei Emerson, 1981; and watanabei Kosuge, 1981. The genus Morum (sensu lato) has long been classified with the mesogastropods in the Cassidae (Thiele, 1929; Boss, 1982; Emerson, 1985). Anatomical studies of Morum, however, indicate that this genus is referable to the neogastropod family Harpidae (Hughes, 1986, and personal communications). Dr. Hughes has in preparation a manuscript in which he rectifies the present taxonomic misplacement of the genus. Recent observations on New and Old World species of Morum (sensu stricto and Oniscidia), moreover, confirm that these gastropods also autotomize the posterior por- tion of the foot (R. Goldberg, P. Williams and W. Liltved, personal communications), a well- known behavorial characteristic of the' genus Harpa (Rehder, 1973). It should be noted that Harpidae Bronn, 1849 (type genus Harpa (Roding, 1798); (Gastropoda), is a homonym of Harpidae Hawle and Corda, 1847 (type genus Harpes Goldfuss, 1839) (Trilobita). Raven (1985) has applied to the International Commission On Zoological Nomenclature to emend the spelling of this gas- tropod family to Harpaidae Bronn, 1849 and to place the emended taxon on the Official List of Family-Group Names in Zoology. Morum (Oniscidia) ninomiyai, new species Figs. 1-5 Morum sp. Kaicher, 1983, #3730, 3 figs., "off Thailand". (Illustrations of the paratype of this species.) Diagnosis: Member of the Morum (0.) cancel- latum (Sowerby, 1824) -M. (0.) watanabei Kosuge, 1981, complex. Distinguished by having the outer lip and the thin parietal shield sculp- tured by numerous, fine, thread-like ridges (Fig. 5). Description: Shell medium size for genus, at- taining 40-1- mm in height, pyriform, attenuated at base. Spire low, extended; protoconch erect and papillate, composed of IV2 glossy, smooth whorls. Postnuclear whorls 5 in number, low conical, weakly shouldered below the suture, first and second whorls weakly cancellated, sub- sequent whorls strongly cancellated, with 14 prominent spinose axial ridges per whorl crossed by 10 moderately weak spiral cords to form a blade-like, hooked spine at the juncture of the spiral and axial ribs; spines most promi- nent at the shoulder. Intervarical areas with 8 to Vol. 100(3) July 30, 1986 THE NAUTILUS 97 FIGS. .\-5.Morum (Oniscidia) ninomiyai, new species. 1 and 2, Holotype, AMNH no. 221241. 3-5, Paratype. T. Ninomiya col- lection (Fig. 5, coated to show sculptural details; photograph courtesy of S. D. Kaieher). 6-9. Morum (Oniscidia) cancellatum (Sowerby, 1821); after Emerson, 1985, pi. 1, figs. 15-18. 6 and 7, off Taiwan, AMNH no. 183783. 8 and 9, Lectotype, BM(NH) no. 197744, "China Seas" (photographs courtesy of A. Beu). All figures except figure 5 approximately x 1. 10 evenly spaced axial lamellae. Aperture nar- row, elongate, semicrescentic in outline. Parietal shield moderately large, thin with outer edge raised and covered by numerous, fine, ir- regular linear lirations, giving the appearance of fine threads. Outer lip thickened, crenulated and weakly toothed, with about 10 primary teeth forming inconspicuous linear projections on the inner labial margin, 2 secondary teeth formed between each pair of primary teeth. Anal sulcus shallow; siphonal canal short, widely open, weakly recurved. Operculum not known. Color: Nucleus shiny, buff; postnuclear whorls with whitish base color, overlaid with flecks of reddish brown; 4 widely spaced, interrupted brownish spiral bands on body whorl (1 band on shoulder, 2 on either side of midbody area, 1 near the base). Aperture milky white; outer lip and parietal shield white and surface thinly glazed (in holotype). Measurements: Holotype, 41.8 mm in height, 25.7 mm in width; paratype, 40.6 mm in height, 25.8 mm in width. Type locality: off Phuket Island, Thailand (8°N, 98.22°E), Andaman Sea, dredged in 50 to 150 meters. Type specimens: holotype, AMNH 221241 (Figs. 1, 2) from type locality; paratype, T. Ninomiya Collection, from "off Thailand, Andaman Sea, ex-Carl Withrow Collection, April 28, 1980", (Figs. 3-5). Distribution: Known only from the type locali- ty and adjacent waters. Remarks: The new species superficially resembles specimens oi Mo7iim (0.) cancellatum (Figs. 6, 7, and 8, 9 (lectotype) and M. (0.) watanabei Kosuge (see Kosuge, 1981, pi. 33, figs. 1, 2 holotype; Emerson, 1985, figs. 11-14), but differs in the number of axial and spiral ridges, the dentition of the outer lip and the unique sculpture of the parietal shield (Fig. 5). 98 THE NAUTILUS July 30, 1986 Vol. 100(3) Acknowledgments We are grateful to Taizo Ninomiya for depositing the holotype of this new taxon in the type collection of moUusks of the American Museum of Natural History. Richard L. Goldberg of Fresh Meadows, New York, William Liltved of San Francisco, California, and Peggy Williams of Sarasota, Florida kindly provided information on living Morum observed or photographed in the field and laboratory. Their generous cooperation is much appre- ciated. Alan G. Beu of the New Zealand Geologi- cal Survey, and Sally Diana Kaicher of St. Petersburg, Florida, generously provided the photographs for figures 8, 9 and 5, respectively. I am also grateful to Dr. Roger N. Hughes of the University College of North Wales for his kindness in keeping me informed of his anatom- ical studies of Morum. I thank my AMNH colleagues for their con- tributions to this study: Walter E. Sage HI for technical assistance, Stephanie Crooms for word-processing the manuscript, and Stephen Butler for the photography, except as noted. LITERATURE CITED Adams, Arthur 1855. Descriptions of new genera and species of gasteropodus |.s-(c"] Mollusca. Pror. Zool. Soc. London for 1853, 21(259): 182-186 (May 16, 1855). Boss, K. J. 1982. Mollusca, in S. P. Parker, ed., Synopsis and Classification of Living Organisms, 1 McGraw-Hill Book Co., New York, pp. 945-1166, illus. Emerson, W. K. 1981. Two new Indo-Pacific species of Morum (Gastropoda: Tonnacea). The Nautilus 95(3): 101-105, 7 figs, 1985. Remarks on some western Pacific species of Morum (Gastropoda: Tonnacea)., in J. M. Lindsay, ed.. Stratigraphy, palaeontology, malacology papers in honour of Dr. Nell Ludbrook. Spec. Publ. So. Aust. Dept. Mines and Energy 5, pp. 51-56, 2 pis. Hughes, R. N. 1986. Anatomy of the foregut of Morum Roding, 1798 (Gastropoda: Tonnacea) and the taxonomic misplacement of the genus. The Veliger 29(1), in press. Kaicher, S. D. 1983. Card Catalogue of World-Wide Shells, Pack #36, Cassidae & Oocorythidae, card #3730. Kosuge, Sadao 1981. Descriptions of two new species of the genus Morum with remarks on the Recent species from Philippines. Bull. Inst. Malac. Tokyo 1(7):101-104, pi. 33. Kuroda, Tokubei and Tadashige Habe, in T. Habe. 1961. Colored Illustrations of the Shells of Japan, 2, Hoikusha, Osaka, Appendix, p. 1-41. Morch, 0. A. L. 1852. Catalogus conchyliorum . . . de Yoldi., Regis Daniae. Copenhagen. Fac. 1. 170 p. Raven, J. G. M. 1985. Homonymy in the families Harpidae Hawle & Corda, 1847 (Trilobita) and Harpidae Bronn, 1849 (Mollusca, Gastropoda). Z.N.(S.) 2331. Bull. Zool. Nom. 42(l):79-80. Rehder, H. A, 1973. The family Harpidae of the world. Indo- Pacific Mollusca 3(16):207-274, pis. 183-247. Sowerby, G. B. I. 1824. The Genera, of Recent and Fossil Shells 1, Oniscia, pi. 233. Thiele, Johannes. 1929. Handbuch der Systematischen Weichtierkunde. Jena. Bd 1, teil 1. pp. 1-376, 470 text figs. A NEW DEEP-WATER SPECIES OF LEPIDOPLEURUS (POLYPLACOPHORA) FROM THE VENEZUELA BASIN. Antonio J. Ferreira^ Research Associate, Department of Invertebrate Zoology California Academy of Sciences San Francisco, CA 94118 ABSTRACT A new species o/Lepidopleurus /rom the depths of the Venezuela Basin, Carib- bean Sea, is described and compared with L. scrippsianus Ferreira. 1980, from the Eastern Pacific. From October to December 1981, the Naval Ocean Research and Development Activity (NORDA), NSTL Station, MS, conducted in- 'Published posthumously. Died 1986. tense physical and biological investigations of deep-sea sediments in the Venezuela Basin from aboard USNS Bartlett (cruise 1301-82). Among the benthic macrofauna were 6 specimens of chitons: 3 specimens, ca. 13, 11, and 9 mm long Vol. 100(3) July 30, 1986 THE NAUTILUS 99 3 4 5 FIGS. 1-5, Lepidopleurus bartletti Ferreira, spec nov.: Holotype (USNM 859000), side view. 2, view of posterior valve. 3, first and second valve. 4, paratype, posterior valve. 5, paratype, under view, valves i, iv and viii. (sta. 40: 15°08'N, 69° 12^, depth 3867 to 4009 m, 28 Oct. 1981); 1 specimen, ca. 15 mm long (sta. 63: 13°45'N, 67°45'W, depth 5046 m, 8 Nov. 1981); 1 specimen, ca. 10 mm long (sta. 65: 13°45'N, 67°45'W, depth 5046 m, 9 Nov. 1981); and 1 specimen, 1.3 mm long (sta. 88: 13°30'N, 64°45'W, depth 3516 to 3550 m, 25-26 Nov. 1981). All chitons were found on pieces of wood brought up in trawls. Study of this material, generously made available through Michael D. Richardson and Paula M. Mikkelsen, revealed a new species here allocated to the genus Lepido- pleurus Risso, 1826. Order Neoloricata Berhenhayn, 1955 Family Lepidopleuridae Pilsbry, 1892 Genus Lepidopleurus Risso, 1826 Lepidopleurus bartletti Ferreira, spec. nov. Fibres 1-6 Diagnosis: Small (up to 15 mm long) white chitons. Valves somewhat rugose; posterior edges moderately angled; surface covered with microgranular cuticle over sculptureless, chalky tegmentum. End valves remarkably flat; mucro slightly anterior; lateral areas not prominent, with some concentric growth rugae continuing into central areas. Gills posterior. Girdle thick; upper surface carpeted with small spiculoid elements with occasional, thin, straight, long spicules interspersed; undersurface reduced to fine cuticle devoid of scales or spicules. Radula median teeth wider in back than in front, with tricuspid major lateral teeth and rakelike spatulate teeth. Type material: Holotype (USNM 859000) and paratypes (CAS 059845; IRCZM 061:085; LACM 2123). Other material: Specimen 1.3 x 0.9 mm, from sta. 88, tentatively identified as L. bartletti (CAS 059846) but not given paratypic status in view of its small size and ill-defined, juvenile characters. Type locality: Venezuela Basin, Caribbean Sea, 13°45'N, 67 "45^, at depth of 5046 m. (Station 63, roughly 150 mi. N.E. of Bonaire, Netherlands Antilles). 100 THE NAUTILUS July 30, 1986 Vol. 100(3) Description: Holotype (from USNV Bartlett sta. 63), preserved in alcohol, curled, ca. 15 mm long, 10 mm wide (figs. 1-3). Valves subcarinate, moderately beaked, with angled posterior edges. Shell surface covered with microgranular cuticle, easily scraped off to show chalky white, sculptureless tegmentum beneath. Anterior valve remarkably flat. Lateral areas of inter- mediate valves not prominent except for greater valve thickness and concentric growth rugae often continuing unto sculptureless cen- tral areas. Posterior valve extremely flat; macro slightly anterior, poorly defined. Gills posterior, about 20 plumes per side, extending 40% of length of foot. Girdle creamy white, thick, with conspicuous straight, thin, glassy spicules inter- spersed on background of much smaller, thin- ner, closely packed spiculoid elements. Paratypes (from USNV Bartlett sta. 40 and sta. 63.) very similar to holotype. Paratype from sta. 40, 13 mm long, disarticulated: Cuticle easi- ly removable from valves uncovering chalky, sculptureless tegmental surface beneath (fig. 4). Articulamentum white with no insertion teeth or slits (fig. 5). Valve i, 5.2 mm wide, 3.2 mm long, 1.2 mm high at posterior edge. Valve v, 6.0 mm wide, 2.0 mm long in midline. Valve viii, 5.1 mm wide, 3.7 mm long, 1.0 mm high at anterior edge, slightly convex in premucro area, slightly concave in postmucro area. Sutural laminae short, triangular; sinus very wide; on valve viii, relative width of sinus (width of sutural sinus/ width of sutural lamina) is 1.8. Girdle dorsal sur- face densely carpeted with blunt-ended, cigar- shaped, spiculoid elements up to 100 x 25 nm (up to 120 X 35 ^im at sutures), often longi- tudinally striated (fig. 6-A), interspersed straight, vaguely striated, glassy spicules up to 300 X 28 ^m (fig. 6-B), interspersed; undersur- face reduced to fine cuticle without scales, spicules or other elements; girdle bridges emp- ty. Radula 6 mm long, comprising 55 rows of mature teeth; median teeth (fig. 6-A) 80 ^im long, 30 ^m wide at anterior blade, enlarging posteriorly to 60 ^m; first lateral teeth about 80 ^)0 PLATE 1. A-D. Litharca lithodomus (Sowerby). Manta, Ecuador. A, Exterior, right valve. B, Interior, left valve. C, Ventral view, both valves showing small byssal gape (shell length 86.5 mm). D, Dorsal view, both valves showing truncated posterior to the left. A-D, California Academy Sciences #055190. E-F, Barhatia iCucuUaearca) reeveana (Orbigny). Perlas, Panama. E, Left valve, exterior. F, Left valve, interior (shell length 79.7 mm). E-F. Florida State Museum #23624. Litharca lithodomus, but the byssal gape is nar- row and elongate, and the byssus is flexible so that the animal is able to move downward in its borehole (Olsson, 1961). As is shown in Frizzell's figure 5 (1946), the valves are wedge-shaped with the most convex portion near the posterior end. The anterior end is narrow. This is an unusual shape for a rock-borer. Thomas (1974, 1976) has assumed that the boring is accomplished by chemical means in Litharca. but this may not be so. Both Frizzell (1946) and Olsson (1961) have observed that Litharca bores into massive sandstones in Ecuador. Are these sandstones calcareous or cemented with calcium carbonate? Field and laboratory observations on these sandstones would be essential to come to a final conclusion as to the mode of boring. Just because Litharca has a weak ligament and a pitted shell exterior does not necessarily rule out the possibility that it may bore mechanically. Furthermore, of the remaining rock-boring species of pelecypods. only some species of mytilids bore by chemical means whereas other species of rock-boring mytilids are mechanical borers (Yonge, 1951). Perhaps, as Thomas implies (1976), both mechanical and chemical means of boring may be employed by L itharca. The fact that at least one arcid species has developed a rock-boring habit is apparently not as incongruous as it would at first seem. The two most unusual features found in Litharca lithodomus that are generally not seen in other rock-boring species are the posterior displace- ment of the beaks and the hatchet-shaped con- vexity of the valves with the most convex region near the posterior end. Most rock-borers are cylindrical in shape, and the beaks are either near the center of the valves or near the anterior end. The byssus is not unique because rock-boring mytilids and tridacnids have this organ developed in the mature rock-boring stage (Yonge, 1951). Compared to some other rock-boring pelecypods, Litharca is relatively 108 THE NAUTILUS July 30, 1986 Vol. 100(3) unspecialized. Rock-boring pelecypods are most diverse in the tropical marine regions. They do not occur in the abyssal and hadal regions of the oceans nor in fresh water. There are at least eight families that have been purported to have one or more rock-boring species and they are: Arcidae, Mytilidae, Tridacnidae, Petricolidae, Hiatelli- dae, Myidae, Gastrochaenidae, and Pholadidae. The Mytilidae and Pholadidae comprise most of the rock-boring species, but there are probably no more than 100 living species of rock-boring pelecypods. Rock-boring species of pelecypods are of average to large size. The largest rock- borers are three species of Tridacna, but they are the smallest-sized species in the family and none attains a maximum size of more than about 400 mm. Furthermore, the species that bores the deepest into the rock is Tridacna (Chametrachea) crocea Lamarck, and it is the smallest of the three species, attaining a max- imum size of only 150 mm (Rosewater, 1965). It also appears that species of pelecypods that attain a size of less than 20 mm are not rock borers, and this may be caused by a lack of bor- ing capabilities in small-sized species. It was assumed by Olsson (1961) and Thomas (1978) that the ancestor of Litharca was a species of Area. An hypothesis ascribed by Dechaseaux (1969, p. N765) to Douville is as follows: "The ancestor was a species living where its earliest representatives are found and having some of its distinctive characters." If one uses this logical principle, then a likely ancestor to Litharca lithodomus could be Barbatia (Cuculla^arca) reeveana (Orbigny, 1856) (plate 1). According to Keen (1971), this is a common species found throughout the present range of Litharca lithodomus. The ribbing and periostra- cum of Barbatia (Cucullaearca) reeveana resem- ble Litharca. The size and convexity of the valves in the two species are similar. There would have to be a reduction of the byssal gape because it is much larger in Barbatia (Cucul- laearca) reeveana than it is in Litharca. The beaks of Barbatia (Cucullaearca) reeveana are commonly subcentral (not near the anterior end as in Area), and it would require not a great modification to have the beaks thrust toward the posterior as in Litharca lithodomus. Bar- batia (CucullcLearca) reeveana has a habit of at- taching between rocks (Keen, 1971) so that a shift to a rock boring habit as in Litharca lithodomus would be a likely change in adapta- tion. This is also borne out by Yonge's observa- tion (1958) that some rock-boring species have evolved from a byssate nestling species. Whether Litharca lithodomus will be a short- lived experiment within the Arcidae or will con- tinue to evolve and produce more rock-boring arcids is impossible to assess. As Frizzell pointed out (1946), the lack of a fossil record of Litharca lithodomus may be caused by the small number of rock-boring habitats in the fossil record and the relatively thin shells of this species inhibiting preservation. However, the morphologic changes for the origin of Litharca lithodomus must have been rapid, and it is quite possible that the appearance of this species was less than one million years ago. Frizzell (1946) erected a separate subfamily, the Litharcinae, for Litharca lithodomus, and Yokes (1980) has followed Frizzell in this classification. We agree with Frizzell and Yokes because taxonomy should show the differences as well as the likenesses between species, and Litharca lithodomus is so unique in morphology and habit that it merits being placed in a separate subfamily within the Arcidae. Acknowledgment We are indebted to Dr. Peter U. Rodda of the California Academy of Sciences for granting us a loan of specimens of Litharca lithodomus for study. LITERATURE CITED Dechaseaux, C. 1969. Hippuritoida-Hippuritacea-Origin and extinction, p. N765. In Treatise on Invertebrate Paleontol- ogy. (R. C. Moore, ed.). Part N, Vol. 2, Mollusca 6, Bivalvia. Univ. Kansas Press, Lawrence. Frizzell, D. L. 1946. A study of two arcid peleeypod species from western South America. Jour. PaLeo. 20:38-51. Keen, A. M. 1971. Sea Shelh of Tropical West America. 2nd ed. Stanford Univ. Press, California. 1064 p. Nicol. D. 1984. Changes in the hinge teeth of arcaceans from Devonian to Recent. Tulane Studies Geol. & Paleo. 18:70-72. Nicol, D., and D. S. Jones. 1984. Review of Postiigata, a Late Cretaceous peleeypod. Tulane Studies Geol. & Paleo. 18:67-69. Oliver, P. (!. 1981. The functional morphology and evolution of Recent Limopsidae (Bivalvia, Arcoida). Malacologia. 21:61-93. Olsson, A. A. 1961 . MoUusks of the Tropical Eastern Pacific: Vol. 100(3) July 30, 1986 THE NAUTILUS 109 Panamic-Pacific Pelecypoda. Paleontological Research Institution. Ithaca, New York. 574 p. Rosewater, J. 1965. The family Tridacnidae in the Indo- Pacific. Indo-Pacific Mollusca l:347-.396. Stanley, S. M. 1972. Fuctional morphologj' and evolution of byssally attached bivalve mollusks. Jour. Paleo. 46:165-202. Thomas, R. D. K. 1976. Constraints of ligamental growth, form and function on evolution in the Arcidae (Mollusca: Bivalvia). Paleobiology 2:64-83. 1978. Shell form and the ecological range of living and extinct Arcoida. Paleobiolog)'. 4:181-194. Yokes, H. E. 1980. Genera of the Bivalvia: A Systematic and Bibliographic Catalogue (Revised and Updated). Paleon- tological Research Institution. Ithaca, New York. 307 p. Yonge, C. M. 1951. Marine boring organisms. Research 4:162-167. 1958. Observations on Petricola carditoides (Conrad). Proc. Malaeological Soc. London 33:25-31. THE RADULA OF ACRORBIS PETRICOLA (PULMONATA: PLANORBIDAE) W. Lobato Paraense Departamento de Malacologia Institute Oswaldo Cruz Caixa Postal 926 20000 Rio de Janeiro, RJ-Brasil ABSTRACT Observation of the radular teeth of Acrorbis petricola Odhner, 1937, and A. odhneri Scott. 1960 with the scanning electron microscope showed no difference between the two nominal species, which also are indistinguishable in shell and anatomic characteristics. A. odhneri, therefore, is a junior synonym, of A. petricola. The radular teeth o/ Acrorbis are similar to those o/Gyraulus, but taxo- nomically important differences in qualitative characters of the genital system show that the two geyiera are less closely related than suggested by their radular characteristics. Acrorbis petricola is an atypical planorbid snail described by Odhner (1937) from speci- mens from Nova Teutonia (27°16'S, 52°20'W), a village in the Brazilian state of Santa Catarina, sent to the Swedish Ricksmuseum by the natural history collector, Herr Fritz Plaumann. Decisive characteristics for the erection of the new germs Acrorbis were the Helix-\ike shape of the shell, and especially the squarish radular teeth with short hooks (see Fig. 1). Pilsbry (1938) pointed to the relationship be- tween Acrorbis and Drepanotrema (misprinted Drepanostoma) by the similarity of the penial complex and the presence of a flagellum on the penial sheath, but distinguished the two genera by the teeth characteristics as shown by Odhner (1937). Baker (1945:123) commented on the alleged differences in the radular teeth: "The radula [of Acrorbis] appears very different from that of Drepanotrema because of the apparent absence of interstitial cusps between the larger cusps. These might have been present but overlooked because of the small size of the radula teeth. The general shape of the mesocone in Odhner's figure 4 is suggestive of the same cusp in Drepanotrema, but the ectocone and entocone are much shorter." . . . "Acrorbis. therefore, appears to be another one of the peculiar groups of South American Planorbidae, related to Drepanotrema, but distinct by both shell and anatomy." Examining specimens from Nova Teutonia, Hubendick (1955) observed that "though not very detailed, Odhner's description and figure of the radula in Acrorbis is not incorrect". His drawings of a central, a lateral and a marginal teeth, reproduced in Fig. 2, may be compared no THE NAUTILUS July 30, 1986 Vol. 100(3) &&®SFQ a^' .''^^v^ ^=^:^ ^^^^ ^Tryv^ lOjjm FIGS. 1-4. 1, Radular teeth of Acrorbis petricola reproduced from papers by Odhner, 1937; 2, Hubendick, 1955; 3, Scott, 1960, under the name Acrorbis odhneri; 4, Paraense and Deslandes, 1959. (c, central; i, in- termediate; 1, lateral; m, marginal. Scale bar refers to Fig. 4). with Odhner's (Fig. 1). A redescription of A. petricola, based on specimens collected at my request by Herr Plaumann at the same breeding-place from which the sample studied by Odhner had been taken, was published by Paraense and Deslandes (1959). The teeth were represented as shown in Fig. 4. Accessory cusps can be seen on either side of the main cusps in the central teeth, and also between the two main cusps in the fifth figured central tooth. Spaces wider than usual appear between the bases of the cusps in several teeth; such spaces, as later realized, were really occupied by small cusps which could not be clearly discerned owing to their minuteness added to light diffraction effect. After discuss- ing the anatomic affinities between Acrorbis and Drepanotrema, Paraense and Deslandes (1959) concluded that the shape of the shell was the only character that might warrant separa- tion of the two genera. Stating, however, that a revision of the genus Drepanotrema would show so wide interspecific variation in shell configura- tion that any generic diagnosis based only on the shell characters of the type species would be meaningless, they treated A. petricola. as a member of Drepanotrema. Subsequently, Paraense (1975) adopted the genus Acrorbis on account of its Helix-shaped shell not sculptured with longitudinal rows of minute dots (puncta- tion), and of its very long spiral radula project- ing far beyond the buccal mass. Under the name Acrorbis odhneri, Scott (1960) described a snail from Argentina anatom- ically identical with A. petricola, but differing in shell characteristics: larger size (largest specimen 4.25 mm in diameter, as against 2 mm in Odhner's specimens), rapid whorl growth, proportionally wider aperture, umbilicus par- tially covered by the basal reflection of the lip, and absence of spiral lines on the shell surface. Scott (1960) states that Odhner so precisely represented half a teeth row that she could recognize in it the radula of her own specimens (Fig. 3). (Translated): "There are no interstitial teeth between the cusps of the laterals, and the marginals are markedly short showing no oblique reflection". Considering the lack of a long oblique reflection in the marginals suffi- cient to separate Acrorbis from the Planorbidae, Scott (1960) created the family Acrorbidae to in- clude A. petricola and A. odhneri. On 17 May 1973 I collected 23 specimens of yl. odhneri at its type-locality, Salto Encantado (27°03'S, 54°50'W), Departamento Libertador Oeneral San Martin, Province of Misiones, Argentina. One may easily reach Salto Encan- tado by Road 14 (Ruta 14) as far as km 212, Vol. 100(3) July 30, 1986 THE NAUTILUS 111 where it intersects Ruta 220 (about 14 km east of Aristobulo del Valle, Departamento Cainguas), and then northward by the last- mentioned road for about 3 km. Salto Encantado is a waterfall which drops vertically and sprays the adjacent cliff, favoring a thick growth of moss where the snail is found. Such peculiar habitat is similar to that of A. petricola, which lives among moss on rocky banks of streams kept wet by trickling water from the surrounding ground and above the stream level. The collected specimens were indistinguish- able, in shell and anatomy, from A. petricola from Nova Teutonia, as described by Paraense and Deslandes (1959). The largest shell was 3.2 mm in diameter. As in the specimens from Nova Teutonia in our collection, the umbilicus was to a variable extent covered by the basal reflection of the lip and the shell surface was more or less perceptibly sculptured with thin spiral lines. Other differences mentioned by Scott (1960) as diagnostic of A. petricola (shell size, rate of whorl increase, proportion of apertural dia- meter) can be ascribed to intraspecific or inter- populational variation. As to the radula, no significant difference was observed between the two forms, so that only teeth of a specimen from Salto Encantado are shown in Figs. 5-10. As observed with the scan- ning electron microscope, the central tooth (Figs. 5, 9) has two dagger-like cusps, of which the left one is longer than the right; there is one (sometimes two) minor spine-shaped cusp high on either lateral side of the major cusps, and another one between the latter. The laterals (Fig. 6) have three dagger-like major cusps, of which the mesocone is much longer, and show one smaller spine-shaped cusp high on the lateral side of the ectocone and entocone and FIGS. 5-lU. Scanning electron micrographs of radular teeth of Acrorbis petricola from Salto Encantado, Misiones province, Argentina. 5, central (arrow), a little slanted to the right; 6, laterals; 7, intermediates; 8, marginals; 9, central, a little slanted to the right; 10, marginals. Figs. 5-8, x 29,000; 9, x 25,000; 10, x 20,000. 112 THE NAUTILUS July 30, 1986 Vol. 100(3) between the major cusps. Small cusps between and outside the three major cusps are added from the intermediates (Fig. 3) to the extreme marginals (Figs. 4, 6), while the ectocone, the entocone and particularly the mesocone remain easily distinguishable. Both major and minor cusps grow smaller toward the edges of the radular ribbon. The marginals have short wide reflections high up on the base of attachment. The extreme marginals usually show 13-14 cusps, including the three main ones. The radular teeth of Acrorbis are similar to those of Gyraulus, recently studied by Meier- Brook (1983) and Burch and Jeong (1984), but the absence in Acrorbis of taxonomically impor- tant characters such as a penial stylet and a separate prostatic duct, and the presence of flagella on the penial sheath, show that the two genera are less closely related than suggested by their radular characteristics. The observations described above point to the identity of A. petricola and A. odhneri, lending no support to the removal of the genus from the family Planorbidae. Acknowledgments The author is grateful to Petrobras-CENPES (Centro de Pesquisas e Desenvolvimento) for use of the facilities of its SEM Laboratory, to Carlos R. Cordeiro and Flavio J. Versiani for technical assistance, and to Prof. Walter Berto- lazzo, Universidade Federal do Rio de Janeiro, who introduced the author to the staff of the SEM Laboratory. LITERATURE CITED Baker, F. C. 1945. The molluscan family Planorbidae. Univ. Illinois Press, Urbana. xxxvi + 530 p. Burch, J. B. and K. H. Jeong. 1984. The radular teeth of selected Planorbidae. Malacol. Rev. 17:67-84. Hubendick, B. 1955. Phylogeny in the Planorbidae. Trans. Zool. Soc. London 28:453-542. Meier-Brook, C. 1983. Taxonomic studies on Gyraulus (Gastropoda: Planorbidae). Malacologia 24:1-113. Odhner, N. H. 1937. Acrorbis petricola n. gen. n. sp., eine merkliche Siisswasserschnecke aus Brasilien. Ark. Zool. 29B:l-8. Paraense, W. L. 1975. Estado atual da sistematica dos planorbideos brasileiros. Arq. Mus. Nac. Rio de Janeiro, 55:105-128. Paraense, W. L. and N. Deslandes. 1959. The Brazilian species oi Drepanotrema. VII. D. petricola (Odhner, 1937). Rev. Brasil Biol. 19:319-329. Pilsbry, H. A. 1938. Acrorbis petricola. Review. The Nautilus 51:107. Scott, M. I. H. 1960. Nueva familia de pulmonado basoma- toforo (Mollusca). Neotropica 6:65-69. NATICA (GLYPHEPITHEMA) TEDBAYERI, A REPLACEMENT NAME Harald A. Rehder National Museum of Natural History Smithsonian Institution Washington, DC 20560 In a recent issue of The Nautilus (Rehder, 1986: Nautilus, 100(1):38) I proposed the name Natica (Glyphepithema) bayeri for Glyphe- pithema Jloridana Rehder, 1943, not Natica Jloridana Dall, 1892. Now Alan R. Kabat at the Museum of Com- parative Zoology has kindly called my attention to the fact that the taxon Natica bayeri is pre- occupied by Natica (Naticina) bayeri Koper- berg, 1931 (Jaarboek von het Mijnwezen in Nederlandsch Oost-Indie, 59: 139, pi. 3, fig. 48) from the Pliocene of Timor, Indonesia. I propose now to name the species found, from southeastern Florida to Brasil, Natica (Glyphepithema) tedbayeri Rehder. Vol. 100(3) July 30, 1986 THE NAUTILUS 113 BOOK REVIEWS Seashells of Western Australia by Fred E. Wells and Clayton W. Bryce. 1986. Western Austra- lian Museum, Perth. 207 pp., 74 colored pis. Paperback. $17.95 Australian (approx. U.S. $12.50). This is an excellent, well-illustrated guide to the more common and readily found species of marine mollusks of Western Australia. The western third of Australia is particularly rich in molluscan species because of the meeting of the tropical Indo-Pacific province in the north with the colder temperate waters of South Australia and Tasmania. Of the approximate 2000 species from this region, this accurate guide beautifully illustrates 671 species, including many photo- graphs of living specimens of opisthobranchs and cephalopods. Each family is defined and, when available, useful bibliographic references are included. Scientific name, author and date, size, geographical distribution, relative abun- dance within Western Australia, and an ex- cellent photograph are given for each of these common and conspicuous species. A short in- troduction on protecting mollusks, cleaning and maintaining a collection of shells, together with a glossary and index round out this fine little book.-/?. T. Abbott. The Littorinid Molluscs of Mangrove Forests in the Indo-Pacific Region, by David G. Reid. 24 April 1986. 228 pp., 98 figs., 1 color plate. Publ. no. 978, British Museum (Natural History). Cloth, $35.00 (Approximately U.S. $52.00). This is one of the most complete and com- petently executed monographs of a well-known group of common mangrove-dwelling gastro- pods. The taxonomy of this otherwise perplex- ing genus, Littoraria. has been well worked out. Despite the recognition of 27 Indo-Pacific species and subspecies in a group heretofore usually treated as three or four good species, the author has proposed only two Indo-Pacific new species, one new subspecies and one new sub- genus, (Paliistorina). His detailed presentation of anatomical, egg-capsule, radular and distribu- tional characteristics will probably lead to a universal acceptance of his views. Very for- tunately a large number of earlier names solely based on shell characters were available for these new biologically recognized species. Ex- cellent drawings of penes, pallial oviducts, sperm nurse cells, and distributional maps ac- company each of the 27 Indo-Pacific taxa. Worldwide, 37 taxa are recognized in five sub- genera of the genus Littoraria Griffith and Pidgeon, 1834. The first 70 pages of this work have an excellent discussion of morphological characters, reproduction, habitats and biogeo- graphy.-/?. T. Abbott. It's Easy to Say Crepidula! by Jean M. Cate and Selma Raskin. 1986. 155 pp. Pretty Penny Press. P.O. Box 3890, Santa Monica, CA 90403. Paperback, $19.95, plus $1.50 postage. This is a useful phoenetic guide to the pronun- ciation of the scientific names of about 3,000 common molluscan taxa. The authors usually give two pronunciations, one evidently used in American Latin classrooms, the other used cus- tomarily among most American malacologists. Readers may take their choice. However, the customary pronunciation of the -ae ending, used in species named for a woman, is not given. The name myrakeeyiae, for instance, is usually pro- nounced: MY' ruh KEEN' ee, but the authors give only MY' ruh KEEN' ay and MY' ru KEEN' eye. They are correct in suggesting that names for males, ending in -i, may be pronounced "eye", as in laurenti: law RENT' eye. There is also an eleven-page glossary of malacological terms.-/?. T. Abbott. Seashell Treasures of the Caribbean by Lesley Sutty. Edited by R. Tucker Abbott. E. P. Dutton, New York. 1986. 128 pp., 139 color plates. $19.95. (with autographed, numbered bookplate $21.95 from American Malacolo- gists, P.O. Box 1192, Burlington, MA 01803). The author, an accomplished naturalist, diver and photographer, has chosen 100 species of Caribbean mollusks "to illustrate and comment upon so that the reader may share my enthu- siasm and joys in discovering the shells of these beautiful tropical islands." She has attempted to inspire others to study mollusks by relating her experiences in seeking these elusive and fascin- ating animals in the Lesser Antilles. Her lively and engaging narrative, coupled with her stun- ning photographs, are certain to provide any shell lover with many hours of pleasurable 114 THE NAUTILUS July 30, 1986 Vol. 100 (3) reading, and should stimulate interest in seek- ing and observing marine shells. A preface by the editor and the author's in- troduction begin this volume, followed by chapters entitled, "What is a Rare Shell?", "Strange and Beautiful Shapes", "Treasures of Sea and Reef, and "Science in Shells." Rare shells discussed and illustrated include Cypraea surinamensis, Pterynotus phyllopterus and Conies granulatus, to name just a few. Unusual mollusks, such as Umbraculum umbraculum, Glossodoris clenchi and Xenophora conchylio- phorci become familiar through the author's animated writing style. The bulk of the species are treated in the "Treasures" chapter where we meet the rediscovery of Hexaplex straussi and such desirable species as Lyria archeri, Conus cedonulli and Chlamys multisquamata. An ab- breviated history of Caribbean malacology, followed by a brief bibliography, index and glossary of terms, completes the text. There is a great deal of merit in this beautiful- ly illustrated book, and it is important again to stress that the author has fully achieved her aim of sharing with others her collecting activities. This book is not intended to be an identification guide. Despite meticulous editing, last minute publisher's errors do creep in, such as the transposition of the captions for plates 22 and 23, and the indiscriminate use of parentheses around every author and date. Nonetheless, it is hoped that this well-produced and lavishly il- lustrated work will be enjoyed and appreciated by nature lovers and conchologists, and will serve to introduce living mollusks to many new enthusiasts -Walter E. Sage. North Atlantic Nudibranchs (MoUusca) Seen By Henning Lemche by Hanne Just and Malcolm Edmunds. 1985. 170 pp., 69 pis. in color. Paperback. Supplement 2 of Ophelia, Inter- national Journal of Marine Biology, Denmark. 500 kroner (approx. U.S. $60.00) The excellent 69 color plates depicting the late Dr. Henning Lemche's personal paintings of 76 of the known 217 species of opisthobranchs of the North Atlantic will prove useful to students of nudibranchs. The descriptions of the species and the information on feeding and reproduc- tion, together with up-dated taxonomic notes by the author-editors add considerable usefulness to the book. A geographical checklist of the 217 species by Elizabeth Platts is very helpful. Most of Lemche's illustrated species are in the genera Doto, Onchidoris, Flabellina. Eubran- chus, Cuthona and Aeolidiella. 26 of the 76 species are given no trivial names and await fur- ther taxonomic consideration. Perhaps someday this unfinished book will be carried to conclusion by one or both of the authors. -i?. T. Abbott. INFORMATION FOR SUBSCRIBERS The annual sviscription rate for The Nautilus is $15.00 for individuals (foreign $18.00) and $20.00 for institutions (domestic or foreign). Subscriptions may begin in January. Send check or money order made out to "American Mala- cologists" to the Business Manager, P.O. Box 2255, Melbourne, Florida 32902-2255, 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. CONTRIBUTORS Mamcscripts: Authors are requested to follow the recommendations of the Style Manual for Biological Journals, which may be purchased from the American Institute of Biological Sci- ences, 1401 Wilson Boulevard, Arlington, Va. 22209. Manuscripts should be typewritten and doublespaced; original and one copy are re- quired, to facilitate reviews. Tables, numbered in arable, should be on separate pages, with the title at the top. Legends to photographs should be typed on separate sheets. Explanatory terms and symbols within a drawing should be neatly printed, or they may be pencilled in on a translu- cent overlay, so that the printer may set them in 8 pt. type. There is a charge of 50 cents per word for this extra service. All authors or their institutions will be charged 50 cents per line of tabular material and taxonomic keys. The pub- lishers reserve the right, seldom exercised, to charge $45 per printed page. An abstract should accompany each paper. Reprints are available at cost to authors. When proof is returned to authors, information about ordering reprints will be given. They are obtained from Economy Printing Co., Inc., R.D. 3, Box 169, Easton, Maryland 21601-9430. MOLLUSK VOUCHER SPECIMENS It is becoming increasingly important for future research purposes that an identified sam- pling of species mentioned in publications be deposited in a permanent, accessible museum specializing in mollusks. This is particularly true of mollusks used in physiological, medical, parasitological, ecological, and experimental projects. Several museums of natural history have ex- tensive modern facilities and equipment for the housing and curating of voucher specimens. Material should be accompanied by the identifi- cation, locality data and its bibliographic reference. There is no charge for this perma- nent curating service, and catalog numbers, if desired, will be sent to authors prior to publica- tion. WANTED - OLD SHELL BOOKS Will pay good prices for libraries, second- Phone (1-305-725-2260) or write: R. Tucker hand books and reprints on mollusks, shells Abbott, American Malacologists, Inc., P.O. and conchology. Back numbers of The Box 2255, Melbourne, FL 32902. Free ap- Nautilu^, vols. 40-71 wanted, $1.50 each. praisals. americanmalacologists, inc. PUBLISHERS OF DISTINCTIVE BOOKS ON MOLLUSKS THE NAUTILUS (Quarterly) MONOGRAPHS OF MARINE MOLLUSCA STANDARD CATALOG OF SHELLS INDEXES TO THE NAUTILUS {Geographical, vols 1-90; Scientific Names, vols 61-90) REGISTER OF AMERICAN MALACOLOGISTS OCTOBER 31, 1986 THE NAUTILUS ISSN 0028-1344 Vol. 100 No. 4 A quarterly devoted to malacology and the interests of conchologists UBRAnr NOV 4 1986 -- ^SlHoie, Mass. Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker. Editor-in-Chief: R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Mr. Samuel L. B. Fuller 1053 Mapleton Avenue Suffield, CT 06078 Dr. M. G. Harasewych 363 Crescendo Way Silver Spring, MD 20901 Mr. Richard L Johnson Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, OH 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill c/o Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, FL 33149 Dr. G. Alan Solem Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, FL 33149 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, FL 32902-2255 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 2255 Melbourne, FL 32902-2255 Second Class Postage paid at Melbourne, Florida and other post offices The Nautilus (USPS 374-980) ISSN 0028-1344 A quarterly magazine devoted to malacology. Copyright ^^1986 by American Malacologists, Inc. OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbourne, FL 32901) Mail: Box 2255, Melbourne, FL 32902-2255 POSTMASTER: Send address changes to above. Subscription Price: $15.00 (see inside back cover) $17.00 (foreign); institutions $20.00 Marine BioioQical bboralorv LIBRARY I NOV 4 19B6 Woods Hole, Mass. THE NAUTILUS Volume 100, number 4 - October 31, 1986 ISSN 0028-1344 CONTENTS " Louise Russert-Kraemer and Elmer G. Berry "Things Truly Excellent. . .": Henry Van Der Schalie, 1907-1986 116 R. Tucker Abbott Cauthdt'us multangulus New Subspecies Grandanus from Northwest Florida (Buccinidae) 120 Jon-Arne Sneli and Oystein Stokland On the Taxonomical Status of Tritonium viridulum Fabricius, 1780 (Gastropoda: Cancellariidae) 121 Douglas G. Smith Pisidium henslowanum (Sheppard) in the Connecticut River, Massachusetts (Bivalvia: Pisidiidae) 124 C. M. Hawkins and R. B. Angus Preliminary Observations of Predation on Ocean Quahaugs, Arctictt islandica, by Atlantic Wolffish, Annrhichas lupu^; 126 Jose D. Gomez. Mercedes Vargas and Emile A. Malek Freshwater Mollusks of the Dominican Republic 130 Paul W. Parmalee and Walter E. Klippel A Prehistoric Aboriginal Freshwater Mussel Assemblage from the Duck River in Middle Tennessee 134 David Nicol The Fate of Pelecypod Families, Subfamilies, and Tribes During and After the Cretaceous Period 140 Emile A. Malek Freshwater and Terrestrial Snails of Saint Lucia, West Indies 143 William K. Emerson and Walter E. Sage III On the Rediscovery of Teniniiiehin minihilin (Clench and Aguayo, 1941), and Its Relationship to Other Calliotectine Volutes 147 To Joe Rosewater — from R. Tucker Abbott Reminiscences 152 All future subscriptions and manuscripts should be sent to Tin Nautilus. P.O. Box 3430, Silver Spring, MD 20901, U.S.A. 116 THE NAUTILUS October 31, 1986 Vol. 100(4) "THINGS TRULY EXCELLENT. . .": HENRY VAN DER SCHALIE, 1907-1986 Louise Russert-Kraemer and Elmer G. Berrv Henry van der Schalie (1907-1986) Henry van der Schalie, extraordinary scholar, teacher and friend to so many of us, died Tues- day afternoon, 15 April, 1986, in his home near the Raisin River in Manchester, Michigan, with his beloved wife and colleague, Annette Rudolphi van der Schalie, at his side. Born in 1907 in Amsterdam, the Netherlands, Professor van der Schalie completed high school in Pater- son, New Jersey, and then earned an A.B. degree from Calvin College in Minnesota in 1929, an M.S. degree from the University of Michigan in 1931, and a Ph.D. from the Univer- sity of Michigan in 1934. At the University of Michigan, he served from 1929 to 1934 as Museum Assistant, from 1934 to 1944 as Assis- tant Curator of Mollusks, and as Curator of Mollusks from 1944 to 1977. He also held an ap- pointment as Instructor of Zoology at the University of Michigan from 1934 to 1937, and served as Exchange Professor at the University of Puerto Rico from 1940 to 1944. Returning to the University of Michigan, he became Assistant Professor of Zoology in 1944, Associate Pro- fessor in 1950, and Professor of Zoology in 1957, a position he held until his retirement in 1977. Highly respected as an outstanding interna- tional authority on mollusks, Professor van der Schalie served on numerous committees and as consultant to government agencies in Egypt, Sudan and Japan. He served as a consultant to the World Health Organization, as well. His work for these organizations centered on the mollusk hosts of human pathogens, such as Schistosoma. A well-known authority not only on disease-bearing snails around the world, but on land and freshwater mollusks of Puerto Rico, on the value of mussel distribution in tracing stream confluence, on the biology of land and freshwater mollusks, on medical malacology, on paleoecology, and other subjects as well. Pro- fessor van der Schalie published well over 110 manuscripts on his research. More complete details on his professional activities can be found in American Malacologists and its sequel, the Register of American Malacologists (1986). In the Bulletin of the American Malacological Union for 1980, Professor van der Schalie pro- vided an illuminating review of "Fifty years of Malacology at the University of Michigan (1929- 1975)." Herein he summarized the outstanding features of the "Goodrich Period (1929-1944)", the "van der Schalie Period (1944-1977)" and the "Highlights of the Mollusk Collections" during those years at the Mollusk Division of the Museum of Zoology at the University of Michigan. At a festschrift organized by his students, for Professor van der Schalie in Ann Arbor in April, 1978, on the occasion of his retirement in 1977, many tributes were presented to "Van" by Vol. 100(4) October 31, 1986 THE NAUTILUS 117 dozens of his students, colleagues and friends. Of particular poignance was the careful com- ment offered by one of Van's oldest friends and colleagues. Professor Elmer G. Berry. Several paragraphs from Dr. Berry's unpublished reminiscence on that occasion, follow: "Henry van der Schalie was born in Amster- dam, January 8, 1907. His father died when Henry was but an infant and his mother, in 1909, emigrated from Holland to Paterson, New Jersey, with Henry and his two older brothers. Circumstances necessitated Mrs. van der Schalie to find employment to provide for herself and the three boys, John, Herman and Henry. She was very gifted in knitting, crocheting and sewing in general and found employment in a cotton factory where, among other items, skirts were manufactured. Most likely, all three boys worked in this plant after school and during the summer months. Henry was assigned to the packing department, par- ticularly the wrapping and tying of cartons of shirts for shipment. Anyone who has observed Henry wrapping and tying a carton can vouch for his adroitness from the experience gained when he was a mere lad. "Somewhere around 1925 Henry left Paterson for Grand Rapids, Michigan, where he enrolled as a student in Calvin College. Although he majored in Zoology he also completed the pre- medical requirements because of his indecision regarding his future profession. The head of the Department at Calvin College was Dr. van Heisman, a recognized parasitologist. Henry had been influenced by (Dr. Heisman) and began to concentrate in the field of parasitology. His splendid scholastic record during his under- graduate years merited commendable recom- mendations from his professors. Although four universities offered him assistantships, he chose the University of Michigan and began his work under Dr. George R. LaRue, an outstanding parasitologist and Chairman of the Department of Zoology. (After coming to Ann Arbor, Henry accepted an assistantship in the Division of Mollusks in the new Museum of Zoology. With some reluc- tance, Dr. Berry reports. Dr. La Rue allowed Henry to transfer.) "Calvin Goodrich was the Curator of Mollusks at that time. He had formerly been the editor of a newspaper in Ohio, but had taken an interest in collecting shells as a hobby. He was ac- quainted with Dr. Bryant Walker, a highly suc- cessful practicing lawyer in Detroit. Walker had made an outstanding reputation as a mala- cologist and had been appointed Honorary Curator of Mollusks at the Museum of Zoology. Goodrich's association with Dr. Walker brought him in contact with A. G. Ruthven, Director of the University Museums (later President of the University of Michigan) and with F. M. Gaige, Director of the Museum of Zoology. It is reported that Goodrich asked Ruthven if he could have a small corner in the new Museums to "play with his shell collection." He was given a room, but the relationship between him and the fine Curator of Mollusks was not the best, which subsequently resulted in the resignation of Miss Mina Winslow. With this termination, Mr. Goodrich was appointed Curator. "Goodrich was a highly sensitive individual and unquestionably realized that being ap- pointed Curator of Mollusks at the University of Michigan was a unique position with a (status) equal to that of Harvard or the Philadelphia Academy of Natural Sciences. He was not academically trained as a scientist (and that) caused him to develop an inferiority complex. He had a flair for writing, especially historical or narrative experiences such as a collecting trip, and was a much better editor of his newspaper than a curator of mollusks. Elmer Berry continues: "Mr. Goodrich replied to my application for an assistantship (at the Museum in 1933) with a frank statement that the Museum's budget had already been trim- med. He advised me that the only job likely to be open would consist of feeding animals and clean- ing their quarters. With the severe cut of ap- propriations, even this job might be eliminated because of lack of funds. (In) his letter written to me on February 2, 1933, he wrote, 'We have been rather fortunate in the students who have worked in this division -H.B. Baker, whose work you are familiar with; W. J. Clench who is now clearing up his long studies of the Physidae; Henry van der Schalie promising to be an authority on the Unionidae; and Alan Archer, who already knows land shells very well and has undertaken to straighten out the Viviparidae of North America. It would be a pleasure to add 118 THE NAUTILUS October 31, 1986 Vol. 100(4) your name to this small, but excellent, company, and my hope is that it can be brought about.' "I shall always remember my introduction to Ann Arbor and the Museum of Zoology. It was terribly hot and humid, (July, 1933) but I was impressed with the beautiful rotunda of the Museum's building and the graceful marble stairs leading to the second floor. That morning I met Henry van der Schalie. He was very friendly and about the first thing he asked was where I was staying. When I told him, the Allenel Hotel, he replied, 'You can't afford to stay there very long!' Henry had been raised to be thrifty as well as energetic. Each month a portion of his small salary went to his mother and the remainder paid for clothing, tuition, books, room and board. He learned quickly that my salary of $60 per month would necessitate stringent budgeting because the cost of living in Ann Arbor, even in 1933, was very high. He in- vited me to have lunch with him, providing I could get along on a peanut butter and jelly sandwich, which I was happy to accept. In order to cut expenses Henry and three other zoolo- gists were sharing a two-room apartment just a half-block east of the Museum Building on North University Street. I don't know what persuasive powers Henry exerted on his three roommates, but by two o'clock on my first day in Ann Arbor, I was invited to become the fifth member of this fraternity. Henry's Dutch ancestry and home- training instilled in him a delight in getting things clean. Each of us had certain duties to perform in the apartment and Henry was assigned to cleaning the kitchen and, on alter- nate Saturdays, the bathroom. On reciprocal Saturdays the bathroom was to be cleaned by Nap Curtis who lived in the west wing of the 2nd floor. When Henry finished his assignment, the bathroom sparkled with cleanliness. Nap, in con- trast, considered it unnecessary to spend much time in cleaning. As soon as Henry discovered (Nap's) perfunctory job, a furor developed and eventually provoked a breach of friendship be- tween the two. "Henry received his I'h.U. in 1934 and at the beginning of the new fiscal year he was ap- [)ointed Assistant Curator of Mollusks. The American Malacological Union meetings were scheduled to be held at Stanford University dur- ing the summer. Coodrich, who was one of the founders of the Union, gave his endorsement (for Henry to attend, and present a paper on his dissertation). I was invited to drive west with him in his Model A Ford. It was Henry's first trip west so we made plans to see as much as we could in the limited time. We included Yellow- stone, the Tetons, Great Salt Lake and planned to do some collecting in the western canyons, Pyramid Lake -a relic lake in Nevada, Lake Tahoe, as well as some side trips in California. We had a delightful trip and met many mala- cologists for the first time, the Junius Hender- sons, the Oldroyds, and saw a few old friends, Stillman Berry, Allyn Smith, Leo Hertlein and others. Best of all Henry and I got to know each other very well and found each other's company very compatible. "Late in 1935 Henry became an active mem- ber of Phi Sigma. He enjoyed the meetings and found the members very congenial. Particularly, he was attracted to a botanist by the name of Annette Rudolphi. Both of them enjoyed ice skating and taking walks through the arbore- tum. By the spring of 1936 it was obvious to me that this attraction had developed into a serious romance. Walking to the Museum and back again, or while preparing dinner or washing dishes, the major conversation was about his girl friend. By midsummer Henry informed me that I had better find another place to live because they were going to be married and would occupy the apartment we were sharing. The date was set for Octotier 3rd and 1 was honored in being chosen as his best man at their wedding. "Goodrich retired in 1944 and Henry was ap- pointed Curator of Mollusks. He inherited a divi- sion rich in historical importance, perhaps the finest collection of land and freshwater mollusks in the country, and a top notch library. Unfor- tunately the yearly budget had i)een reduced to poverty level. Henry didn't remain discouraged for very long, however, and his frugality, cou- pled with his exploring abilities that might pro- vide increments to purchase microsco[)es, microtomes, establishing aquarium rooms, etc., not only restored the budget to its original status, but surpassed it many times. The new addition to the Museums Building was con- structed and this allowed the Mollusk Division to expand in scientific areas uniciue in malacology. Vol. 100(4) October 31, 1986 THE NAUTILUS 119 The Division's liigh status subsequently at- tracted splendid investigators to receive their advanced training in malacology at the Univer- sity of Michigan under Henry's direction. "I well recall two phrases Henry frequently used, particularly when it involved a person with whom he disagreed. He would say, "Well let me tell you, I've got news for him!" This remark would be followed with what he was going to do, and he wasn't fooling! The magnificent achieve- ments accomplished by the Mollusk Division were due to Henry's tremendous effort and unbelievable drive. When deeply disappointed by a granting agency's reply to a grant request which Henry had painstakingly made, and which had looked promising, that stated, 'We regret to inform you. . .' Henry, having given vent to his feelings, would end the discussion with a grin and a shrug of his shoulders, saying, 'Oh well, we're having fun!' " He meant it! An inspiring teacher and fierce advocate for rigorous study and understanding of mollusks, Henry van der Schalie expected a lot from us. He expected us to be rational, careful, committed, and to derive joy from our dedication, as he did from his. As one of his last students, I have treasured the great legacy of his intellect and his spirit. I never knew "Van" to be half-hearted about anything. He was em- phatic, enthusiastic about whatever he was do- ing, whether it was digging into a new research project, encouraging a student, pursuing his beloved mollusks in their improbable habitats, or sharing his parenting energies with his in- comparable wife and colleague, Annette. He regularly doted on all the accomplishments of his children, each of whom he and Annette always regarded with boundless, affectionate concern and pride. Van had a strong environmental ethic long before the raised consciousness of the present era. He evinced articulate, righteous indigna- tion whenever he perceived that human error, indifference or greed had resulted in the destruction of a precious habitat, or failed to fund research, or allowed neglect of irreplace- able museum collections. I remember his re- counting to me his participation in a panel discussion, along with other biologists and lawyers, on the University of Michigan campus, in the 1970's, on aspects of environmental law. Van related to me his consternation with the at- titude evinced by his colleagues, who had caved in to sophistry and had adopted a resigned, expedient demeanor in the discussion. Van ex- postulated, "My God! When I think of what (natural resources) we have squandered!" Van insisted, rightly I think, that better laws to pro- tect the environment could not be made in a society where knowledgeable experts didn't care. He believed passionately in the coupling of the fine old values of knowledge and respon- sibility. Van's capacity for moral outrage was good for us all. It continues to inspire and sus- tain us as we pursue urgent, sometimes un- popular, environmental causes of our own. In recent years, a visit to Manchester, Michigan, to see Van and Annette for good mollusk talk, in their palatial spread along the Raisin River, was a revitalizing joy. One ambled with them about their acres, past the huge, burgeoning vegetable gardens Van and Annette and their son, Arnold, were cultivating in delighted competition with each other. One followed them through the verdant, leafy marshland next to the river, where Van would pridefully point out the most recent wonders of Annette's botanical expertise, and then to the Raisin River itself, where, thanks to van der Schalie effort, a number of species of Lampsilis still maintain reproducing populations. On such jaunts I always got a lump in my throat. I knew again that this wonderful man, Henry van der Schalie, lived a life of indefatig- able personal integrity. His career spanned almost five decades of vital research at the University of Michigan and around the world. More, to the end, he cared deeply about mollusks as mollusks, about human beings as human beings. In a glowing, over-arching way Van cared about the lovely planet that we share. Such jaunts invariably left me with the poignant reminder that I shall always think of Van as a grand model. I shall always think of Van with admiration and love. 120 THE NAUTILUS October 31, 1986 Vol. 100(4) CANTHARUS MULTANGULUS NEW SUBSPECIES GRANDANUS FROM NORTHWEST FLORIDA (BUCCINIDAE) R. Tucker Abbott American Malacologists, Inc. Melbourne, Florida 32902-2255 During the last few years, members of the Gulf Coast Shell Club in Panama City, north- west Florida, have been very active in surveying the marine mollusks of their area. In fact, they have published several checklists and in 1983 issued their illustrated Seashells of Bay County and the Gulf Coast. In 1982, Robert Granda, and later other members, collected and sent me an unusual form of Cantharus which I now take pleasure in naming after its original discoverer. Family Buccinidae Rafinesque, 1815 Genus Canthnrn.'^ Roding, 1798 Cantharus tnultangulus (Philippi, 1848) new subspecies grandanus Descrij)tiom.-?>\\Q\\ small, up to 28 mm in length, fusiform, light-weight, finely but coarse- ly sculptured, and with 6 or 7 whorls. Nuclear whorls IV2, smooth, rounded, rapidly descend- ing, translucent tan to clear. First 4 or 5 postnuclear whorls slightly shouldered, with 10 or 11 strong, rounded, axial ribs crossed by FIG. 1. Canlhariix muUanifuiu.s subspecies ynmdanux Abbott, holotype, UNNM no. 8.59096, 28.7 x 14.4 mm. FIG. 2. Cantharus multangulus multangulus (F^hilippi, 1848) from off Shell Island, Panama City, N.W. Florida, 29.1 x 15.6 mm. Both collected by Robert Granda, 1982. about 9 or 10 very small, irregularly-sized, square-topped, spiral threads. On the penulti- mate and last whorl, which is smoothly rounded, the axial ribs are obsolete. Body whorl and short siphonal canal with about 30 to 40 fine spiral threads of uneven size. Columella with two weak, oblique plicae at the base. Outer lip simple and minutely jagged. Inside of last whorl glossy tan and with about a dozen, smooth spiral threads. Color of outer shell tannish brown, red- dish or whitish yellow with numerous fine flecks, occasional narrow spiral bands and sparse axial flames of chocolate-brown. Oper- culum chitinous, translucent tan, elongate-oval and filling most of the aperture. Soft parts and radulae not examined. Type locality -2 to 4 feet of water, on sand and weed bottom. Islet, V4 mi east of Black Isle, Saint Joseph Bay, off Port St. Joe, Gulf County, northwest Florida. Robert Granda, collector, March 13, 1982. Types -The holotype is deposited in U. S. Na- tional Museum as no. 859096. Parat>']oes from the type locality have been deposited in the Academy of Natural Sciences of Philadelphia, the American Museum of Natural History, N.Y., the Museum of Comparative Zoology at Har- vard, and the Florida State Museum, University of Gainesville, Florida. Records -This moderately rare subspecies has been collected in several places in St. Joe Bay. Linda Brunner has also collected this subspecies in St. Andrews Bay in nearby Bay County. Measurements (mm.)- length width Holotype 28.7 14.4 F'aratype 26.8 13.1 Paratype 26.7 14.0 i'aralype 24.0 12.1 Paratype 21.1 11.0 Paratype 21.0 11.0 Remarks-The subspecies grandanus of the northeast Gulf of Mexico differs from the nominate Cantharus multangulus (Philippi, Vol. 100(4) October 31, 1986 THE NAUTILUS 121 1848) in having a lighter-weight shell, lacking the strong shoulder nodules on the last two whorls, in having more numerous spiral threads, and in being a little more elongate in propor- tions. The variations in colors and patterns are very similar. It is possible that these colonies in the "Panhandle" region of Florida represent an ecological, rather than a genetic, form. Mr. Granda obtained two "clutches" of small, horny egg-capsules which he found on pieces of carapace from the horseshoe crab, Limulus. The urn-shaped capsules, about 5x8 mm, closely resembled those so well illustrated by D'Asaro in his account of the capsules of CantharuH multangulus from the same region (1986, p. 86, figs. A-D). Very similar capsules of the nominate species from Sanibel Island were il- lustrated by Perry and Schwengel, 1955, pi. 50, fig. 340. Cantharus cancellarius (Conrad, 1846) from the same region differs in lieing more ovoid, having a shorter spire and in having much stronger and fewer spiral threads. The similar muricid, Cnlotrophon ostrearum {Conrad, 1846), has stronger shoulder nodes and a mauve to rosy-purple aperture. Fossil C. multangulus from the old St. Petersburg pits have fewer axial nodes per whorl. This grou]) of species ap- pears to t>e largely confined to southeast United States, the Bahamas, the north coast of Cuba and Yucatan, Mexico. LITERATURE CITED D'Asaro. Charles N. 1986. Egg Capsules of Eleven Marine Prosobranchs from Northwest Florida. BtiU. Murine Sri. 39(1): 76-91, 4 figs. Gulf Coast Shell Club (Bob Cranda and -Jim Brunner, editors). 1983. SeaskeLls of Bay Cininlii mtd the Gi(lfC(t(it;t. 26 pp., 96 photos, 1 map. Perry, Louise M. and Jeanne S. Schwengel 19.5.5. Murine Shells of the Westeiyt Coast of FUiridti. Paleont. Research Inst.. Ithaca. 318 pp, 55 pis. Robertson, Robert. 1957. A Study of Cantharus multaiigu- his (Philippi) with Notes on Cantharus and Pseudo- neptunea (Gastropoda:Buccinidae). Notulae Naturae. Philadelphia, no. 300, pp. 1-10, 19 figs. ON THE TAXONOMICAL STATUS OF TRITONIUM VIRIDULUM FABRICIUS, 1780 (GASTROPODA: CANCELLARIIDAE) Jon-Arne Sneli and Oystein Stokland Trondhjem Biologiske Stasjon 7000 Trondheim, Norway ABSTRACT The taxonoviic and nomenclatorial problems associated with the species Admete viridula (Fahricius. 1780) are discussed. The fact that the type specimens of both Admete viridula and A. crisps. Moller, 1842 are misingfrom the Zoological Museum in Copenhagen complicates the situation considerably as three species are involved, Tritonium viridulum Fabricius, 1780, Defrancia viridula Moller, 1842. and D. exarata Moller. 1842. Fabricius's name is to be regarded as a nomen dubium and that Admete couthouyi (Jay, 1839) should be used as the correct name for that species. Mailer's D. viridula and D. exarata are both good species; but D. viridula should probably be named Oenopota decussata (Couthouy, 1839) and D. exarata, Propebela exarata (Moller, 1842). The original description of Tritonium viridulum, which was published by Fabricius in 1780 without an illustration, has commonly been regarded as the original description of Admete viridula auctt. The description fits the species fairly well, and the taxonomic situation would appear quite simple. However, Ball (1886: 298) after having ex- amined the holotype in the Zoological Museum in Copenhagen, claimed that the species be- longed to the group Bela auctt. He also found that Defrancia viridula Moller, 1842 (Fig. 1) 122 THE NAUTILUS October 31, 1986 Vol. 100(4) 0H:- 5 6 ^7 FIGS. 1-7. 1, Pro-pehela exarata (Moller, 1842); one of the larger specimens from the syntype-collection of Moller (Zool. nius., Copenhagen). 2, Propebela exarntri (Moller) as drawn by G. O. Sars. 1878. 3 and 4, A specimen of Oenuputn deccusata (Couthouy, 1839) = Bda viriduta. from the collection of Moller (Zool. mus, Copenhagen). On the label is also written: "B. viridula (et var. inflata) = B. deccusata Couthouy var. ventricosa". 5, Admete viriduta uctt. as drawn by G. 0. Sars, 1878. 6 and 7, Holotype of Cancellaria buccinoides Couthouy, 1838 (Reg. no. 279394 in Mus. Comp. Zool, Cambridge, Mass.). The species is Admete couthouyi Jay, 1839. was founded on the same specimen, and con- cluded that it was identical with Defrancia ex- arata Moller, 1842 (Fig. 2). Ball writes that Morch had discovered these facts before him and Posselt (1898: 168) confirms this by referr- ing to a handwritten catalogue by Morch. Con- cerning the nomenclatorial situation, Posselt claims that the correct name o^ Admete viridula auctt. (Fig. 3) should be Admete couthouyi Jay, 1839, because the older Cancellaria buccinoides^ (Fig. 4) of Couthouy, 1838 was described in another genus. All three species involved in this problem show a great deal of variability, and Fabricius description fits both Moller's Defrancia viridula and Admete viridula auctt. as well. Concerning Defrancia exarata, Posselt (1898: 168) states that Admete crispa Moller 1842, which conmion- ly is regarded as conspecific with A. viridula auctt., resembles D. exarata with respect to sculpture. This situation could have easily been cleared up if the type specimens in question, which were all from Greenland, had been available, but un- fortunately the type specimens of both Tritonium riridulum and Admete crispa appear to have disappeared early in this century (Jorgen Knudsen, pers. commn.). Only one sam- ple in the collection of the Zoological Museum in Copenhagen could possibly be Fabricius type, but this sample contains three specimens whereas the original description states that Fabricius had only one specimen available. In the Zoological Museum in Copenhagen there are type lots of both Mollers species Defrancia ex- arata (Fig. 2) and D. viridula (Fig. 1). The type Vol. 100(4) October 31, 1986 THE NAUTILUS 123 lots of the last species contains seven syntypes and thereby shows that Moller did not base his description on Fabricius specimen. None of Mollers type specimen fits very well with Fabricius description. However, Knipowitsch (1901) illustrates a specimen oi D. exarata from Svalbard which rather closely resembles Admete viridula auctt. Considering this, and that the form caWed Admete critipa has a sculpture like that of D. exarata, we assume that some morphological overlap between the species is present. Although fitting the common forms of Admete viridula auctt. better than those of Holler's Dejrancia exarata, Fabricius description covers the overlapping forms, with the exception that we never have observed D. exarata without prominent ribs on the last whorl as mentioned in the description. How- ever, considering the large intraspecific varia- tion within most species in this group it is no wonder that Dall (1886) reduced them to synonomy. Moller's sample of Defrancia viridula fits Fabricius description fairly well except that the ribs, which are said to be straight ("costae longitudinales") in Fabricius description, are somewhat curved. As Moller's Defrancia viridula, which probably is conspecific with Pleurot.oma decussata. Couthouy, 1839, has a large intraspecific variation like all species in this group, Fabricius description fits both species fairly well, and one cannot apply it to one rather than the other. Concerning Morch's earlier opinions as first published by Dall (1886), there is a sample in the Zoological Museum, University of Bergen (no. 28208) which should be mentioned. This con- tains two dry specimens of Moller's viridula from Greenland which are identified to "Bela viridula M. Sars" by Morch. Michael Sars never described any species with this name, but the sample could indicate that Morch had Moller's species in mind when he claimed to have dis- covered that this and Fabricius species was founded on the same specimen. However, this is in contradiction to the presence of the seven syntypes of Defrancia viridula Moller in the Zoological Museum of Copenhagen. Some additional information about the prob- lems within the genus Admete may be found in Troschel's (1866-1893) work on gastropod radulae. Troschel investigated preserved material of both Defrancia viridula and the genus Admete in the Zoological Museum in Copenhagen with respect to radular teeth. He found the radulae in D. viridula were like those of other species within Beta auctt., while in Admete he found two kinds of teeth. He also found two shell forms which he called Adm,ete viridula (Fabricius) and A. crispa Moller accor- ding to figures in Middendorffs (1849) work on Russian mollusks. The shell forms corresponded with the radular forms, which may indicate that Troschel was correct in dividing the traditional Admete viridula into two species. The nomenclatorial consequences of these tax- onomic circumstances may now be summarized. Moller's Defrancia exarata presents no prob- lems, and in our opinion it is best placed within Propebela Iredale, 1918. Fabricius Tritionium viridulum seems difficult to identify with any species, and in our opinion it should, since the holotype is lost, be regarded as a nomen dubium. One could use the existence of the sample iden- tified by Morch to attach the name to Moller's viridula, having also the facts presented by Dall in mind. However, since both Morch and Dall seems to have considered that Fabricius original specimen belonged to Propebela exarata (Moller, 1842), this argument seems somewhat dubious. The fact that Moller founded his Defrancia viridula on a sample of seven syntypes still pre- sent in the Zoological Museum in Copenhagen also contradicts the view of Morch and Dall. In our opinion, Moller's Defrancia viridula is best placed in Oenopota Morch, 1852, probably as a synonym of Oenopota decussata (Couthouy, 1839). Concerning ylrf?nete viridula auctt., the oldest name for this species is Cancellaria buccinoides Couthouy, 1838. This name is, however, a primary homonym of Cancellaria buccinoides W. Wood, 1828 and has to be rejected as there are no strong reasons why it should be referred to the International Commission on Zoological Nomenclature for eventual preservation. The next available name is Admete couthouyi Jay, 1839, which meets all demands as a correct name for the species. The type of this is also Couthouy 's type of buccinoides since Jay's name was proposed as a nomen novum. An eventual separation of Mollers Adynete crispa from A. 124 THE NAUTILUS October 31, 1986 Vol. 100(4) couthouyi as proposed by Troschel (1866-1893) will not be discussed in detail, but as the type specimen of A. crispn is lost either a neotype should be selected or a new name should be pro- posed and Mdller's name, A. crispa, regarded as a nornen dubium. As MoUer never described the radula of his species the second possibility is probably the best. Mr. Georg Crawford kindly corrected our English text. LITERATURE CITED Couthouy, J. P. 1838. Descriptions of new species of Mollusca and shells and remarks on several polypi found in Massachusetts Bay. Boston Jour. Nat. Hist. 2:53-111. 1839. Monograph on the family Osteodesmacea of Deshayes, with remarks on two species o( Patelloidea, and descriptions of new species of marine shells, a species of Anculotu.'i. and one of Eolis. Boston Jour. Hist. 2:129-189. Dall, W. H. 1886. Supplementary notes on some species of mollusks of the Bering Sea and vicinity. Proc. U. S. natn. M«,s. 9:297-309. Fahricius, 0. 1780. Faumi Groenliindini. Johannes Gottloli Rolhe, Kobenhavn. 452 pp. Iredale, T. 1918. Molluscan nomenclatural prulilems and solutions. -No. 1. Proc. Malac. Soc. Land. 13:28-40. Jay, J. C. 1839. A catalogue of shells in the collection of John C. Jay, M.D., with descriptions. Ed. 3. 125 pp. Knipowitsch, N. 1901. Ueber die in den Jahren 1899-1900 im Gebiete von Spitzbergen gesammelten recenten Mollus- ken und Brachiopoden. Ezheg. zool. Muz. 6:435-558. Middendorff, A. T. v. 1849. Beitrage zu einer Malacozoologia Rossica. Abt. 2 & 3. Mem. Acad. Imper. Sci. St.-Petersb.. ser. 6. Sci. nat. 6:1-187. Moller, H. P. C. 1842. Index Molluscorum Groenlandiae. Naturhist. Tids.'^kr. ser. 1. 4:76-97. Posselt, H. J. (A. S. Jensen ex.) 1898. Grondlands brachiopoder og bloddyr. Meddr. Gronland 23:1-298. Sars, G. O. 1878. Mollusca regionis arcticae norvegiae. Bidrag til kundskaben oni Norges nrkliske fiiuna. A. W. Brogger, Christiania (Oslo). 446 pp. Trochel. F. H. 1866-1893 (cont. by J. Thiele in 1891). Das Gebiss der Schnecken zur Begmndung einer naturlichen Classification. Vol. 2. Nicolaische Verlags-Buchhandl., Berlin. Wood, W. 1828. Index Testnceoi.Ed. 2. with the plates and suppL, London. 188 |)p. PISIDIUM HENSLOWANUM (SHEPPARD) IN THE CONNECTICUT RIVER, MASSACHUSETTS (BIVALVIA: PISIDIIDAE) Douglas G. Smith Museum of Zoology University of Massachusetts Amherst, Massachusetts 01003-0027 ABSTRACT The pisidiid dam Pisidium henslowanum ha^ been coyisidered by several authors as an introdnced species in North America despite its discovery in several parts of North America well away from industrial centers. The species has been recently found in the Connecticut River in Massachusetts which is the southeastern most record yet in North America. It is suggested that this .species is native but locally distributed. The Connecticut River record seems also to repre- sent the softest water in which the species has been reported in North America. Since its first discovery in North America by Sterki (1899), the pisidiid clam, Pisidium henslowanum (Sheppard, 182.'j), was known on the continent from only a small number of sites mostly clustered in the Great Lakes and St. Lawrence River drainages (Heard, 1961, 1962; Ilerrington, 1962). The species is well known in the palearctic region (Woodward, 1913; Zhadin, 1957; Ellis, 1978). Herrington (1962, 1965) and Heard (1962) determined that P. henslowanum was introduced into North America. Subsequent to Herrington's (1962) and La Roccjue's (1967) reviews of the North American distribution of P. henslou'anum. the species was reported from a number of localities in central and western Canada by Harris (1973), who also documented Vol. 100(4) October 31, 1986 THE NAUTILUS 125 the occurrence of P. henslowanum in 7000 year old deposits. Harris (1973) used this information to question Herrington's (1962) and Heard's (1962) conclusions that the species was not native to North America. Harris (1973) also cited Herrington's (1962) listing of a much earlier fossil date (Pliocene-Pleistocene) for P. supinum, then considered by Herrington a synonym of P. henslowanum. However, subse- quent study led Herrington (1965) to accept P. supinum as a distinct species, thus weakening Harris' (1973) argument. Nonetheless, Harris' (1973) data compelled a reassessment of the in- troduced status of P. henslowanum, yet subse- quent studies (Burch, 1975; Clarke, 1981; Mackie, 1981) continued to follow Herrington (1962, 1965) and Heard (1962). During the summer of 1985, P. henslowanmn was collected in the Connecticut River at two locations in Hampden County, Massachusetts. The first collection was made in the Holyoke in- dustrial canal complex (in direct communication with the river) in Holyoke and the second in the river in Longmeadow. The Holyoke collections were first made on 1 July and repeated on 27 October. Prior to each collection the canals had been drained for routine maintenance. A total of 98 specimens were collected, of which a series of shells have been deposited into the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ). The remain- ing alcohol preserved material has been placed into the Museum of Zoology, University of Massachusetts at Amherst. The Longmeadow collection was made on 16 September and com- prised a single adult specimen. Previous to these collections the only New England records for P. henslowanum were Lake Champlain in Vermont (St. Lawrence River system) and Aroostook County, Maine (Johnson, 1915). Although the Vermont record is generally accepted in subse- quent literature, the Maine record has not been repeated. The Maine record was listed by Lermond (1909) as well and was credited to Nylander as collector and Sterki (presumably) as the authority. A search of the pisidiid collec- tions at the MCZ revealed no specimens of P. henslowanum from either locality. The Connecticut River is clearly outside the Great Lakes and St. Lawrence River drainages and the populations in the Connecticut River could not have been established by recent migra- tion. Additionally, the section of the Connec- ticut River in Massachusetts is well above the commercially navigable portion of the river and it seems unlikely that the population in the Holyoke canals has been artificially founded by "stowaways" from Europe. As argued by Harris (1973), this species may not be introduced in North America. It might, however, be native but very localized and thereby easily overlooked. The Holyoke canal collections were made in mud and silt sediments normally under 4.5 m of water. The single Longmeadow specimen was collected in .5 m of water. The species is com- mon in the Holyoke canals, particularly along the base of the retaining walls, and is associated with the pisidiid species Musculium securis and Sphaerium. striatinum and juveniles of the unionid mussels Elliptio complanata and Ano- donta implicata. Chemical data for the canals (Anon., 1978, 1980) includes the following para- meters (during July, August and September): pH = 7.0-7.7, total alkalinity = 25-35 mg/1 (CaC03), dissolved oxygen = 5.2-10.4 mg/1, nitrate = 0.0-0.5 mg/1, chloride = 7-35 mg/1. The chloride values in part represent runoff of residues of municipal operations. Chlorides in the Longmeadow portion of the river average about 10 mg/1 (Anon., 1978). These values com- pare well with data given by Harris (1973) and Okland and Kuiper (1980) except for the alkalin- ity values. In Europe, Ellis (1978) has indicated that P. henslowanum prefers calcium rich waters and Harris (1973) concludes that the presence of dissolved salts are an important characteristic of this species' habitat. Overall, the water of the Connecticut River is relatively soft and has average levels of chloride salts (Wetzel, 1975). Thus P. henslowanum can ap- parently exist in somewhat soft water. Specimens collected on 1 July were for the most part larviparous (11 of 13 specimens or 84% of the sample), and the 16 September specimen from Longmeadow was larviparous also. Of the 27 October sample, however, only 2% (2 of 85) were larviparous though many con- tained egg masses in the gills. These seasonal data compare well with Holopainen and Jonasson's (1983) findings for the presence of brooded larvae in Danish populations. 126 THE NAUTILUS October 31, 1986 Vol. 100(4) Acknowledgments I thank Alan Richmond and Jenifer Bush for assistance in the field. I also thank D. Kenneth Boss for allowing access to collections at the MCZ, Harvard University. LITERATURE CITED Anonymous. 1978. The Connecticut River water quality data- 1978. Massachusetts Department Environmental Quality Engineering. Division Water Pollution Control. Westborough, Massachusetts. 147 pp. . 1980. Connecticut River Water Quality Data- 1980. Massachusetts Department Environmental Quality Engineering. Division Water Pollution Control. West- borough, Massachusetts. 60 pp. Burch, J. B. 1975. Freshwater Sphaeriacean Clams (Mol- lusca: Pelecypoda) of North America. Malacological Pub- lications, Hamburg, Michigan. 96 pp. Clarke, A. H. 1981. The Freshivaler Molluscs of Canada. National Museums of Canada, Ottawa. 446 pp. Ellis, A. E. 1978. British Freshwater Bivalve Mollusca. Synopsis of the British Fauna (New Series) No. 11. Academic Press, London. 109 pp. Harris, S. A. 1973. Pisidium henslowanum (Sheppard) in western Canada. The Nautilus 87:86-87. Heard, W. H. 1961. Pisidium henslouianum (Sheppard) in Lake Michigan. The Nautilu.^ 74:12.3. 1962. Distribution of Sphaeriidae (Pelecypoda) in Michigan, U.S.A. Malacologia 1:139-161. Herrington, H. B. 1962. A revision of the Sphaeriidae of North America (Mollusca: Pelecypoda). Miscellaneous Publications Museum of Zoology. Univ. Michigan. No. 118. 174 pp. 1965. Corrections of sphaeriid nomenclature. The Nautilus 79:42-45. Holopainen, L J. and P. M. Jonasson. 1983. Long-term popu- lation dynamics and production of Pisidium (Bivalvia) in the profundal of Lake Esrom, Denmark. Oikos 41:99-117. Johnson, C. W. 1915. Fauna of New England 13. List of the Mollusca. Occasional Papers Boston Society Natural History 7(13):1-231. Lermond, N. W. 1909. Shells of Maine. Report of the Com- missioner of Agriculture of Maine, 1909. Pp. 217-262. Mackie, G. L. 1981. Nearctic freshwater Sphaeriacea (Bivalvia). Bulletin Amfrican Malacological Union for 1981: 49-52. Okland, K. A. and J. G. J. Kuiper. 1980. Small mussels (Sphaeriidae) in freshwater in Norway-distribution, ecology', and relation to acidification. SNSF project, IR 61/80. Oslo-As, Norway. 85 pp. Sterki, V. 1899. Pisidia new to our country, and new species. The Nautilus 13:9-12. Wetzel, R. G. 1975. Limnology. W. B. Saunders Co., Phila- delphia. 743 pp. Woodward, B. B. 1913. Catalogue of the British species of Pisidium (Recent and fossil) in the collections of the British Museum (Natural History), with notes on those of western Europe. British Museum (Natural History), Lon- don. 144 pp. Zhadin, V. I. 1952. MoUusks of fresh and brackish waters of the U.S.S.R. Zoological Institute Academy Sciences U.S.S.R. No. 46. (1965 Israel Program Scientific Transla- tion. 365 pp.) PRELIMINARY OBSERVATIONS OF PREDATION ON OCEAN QUAHAUGS, ARCTICA ISLANDICA, BY ATLANTIC WOLFFISH, ANARHICHAS LUPUS C. M. Hawkins and R. B. Angus Fisheries Research Branch, Scotia-Fundy Region Halifax Fisheries Research Laboratory Halifax, Nova Scotia B3J 2S7 ABSTRACT Populations of the ocean quahaug, Arctica islandica, may be siisceptable to in- tensive grazing pressure by the Atlantic wolffish. Anarhichas lupus. In stomach analysis of four wolffish from an inshore Nova Scotia area, 85-99% of the wet weight contents were Arctica islandica. The number of quahaugs in the stomach contents ranged from 3-11. These individuals were 35-86 mm. in shell length and were 4-60 years old. The ocean quahaug (or quahog), Arctica is- landica (Linnaeus, 1767), is an underexploited commercial mollusk found in coastal waters of the south shore of Nova Scotia, Canada, and on the Scotian Shelf (Rowell and Chaisson, 1983). These bivalves are slow* growing (Murawski et Vol. 100(4) October 31, 1986 THE NAUTILUS 127 al, 1982) reaching a minimum market size of about 51 mm (2 inches) in 13 years, and long- lived (225 years, Ropes and Murawaski, 1983). Since they are commercially important in the northeast of the United States (Murawski et a/., 1982) and are of potential commercial impor- tance in the south shore area of Nova Scotia it is important for the fishery biologist and manager to assess the impact of natural predation on stocks in relation to stock assessment and potential yields. Quantative data on the diet of the common Atlantic wolffish is sparce as are data on natural predation of ocean quahaugs. Early works (Verrill 1871, Gill 1911 and others) are primarily qualitative but indicate that shell- fish play an important role in wolffish nutrition. Bowman et al. (1976) provide more quantitative data. They examined the stomach contents of 127 wolffish from offshore waters of western Nova Scotia. Approximately 42% of all stomachs were empty, but the remainder on average contained about 28% (by wet weight) bivalves. However, no further taxonomic break- down was made. Although there is evidence of offshore predation of Ardica islandica by cod {Gadus morhua) (Arntz 1974, 1978) there is a lack of both qualitative and quantitative data relating to predator induced natural mortality ofArctica islandica in coastal waters. From our investigation in the coastal waters of Nova Scotia, Canada, qualitative and quantitative data make the common Atlantic wolffish, Anarhichas lupus, a prime suspect as an impor- tant predator oi Ardica islandica. During the first of a three year study begun in 1982 to study maturation oi Ardica islandica at Port Mouton, Nova Scotia (64 5rN, 43 56'W) one of us (RBA) noted the arrival of wolffish to the study site in the spring and their departure in late summer. The time of this annual migra- tion corresponded to wolffish bycatches in net and traps reported by local fishermen operating in other areas near our study site. The fisher- men also noted that the wolffish guts were full of shell material. This information prompted us to catch wolffish at our study site to determine whether or not they were feeding on quahaugs and, if so, their impact on known quahaug stocks. The study site, about 4000 m^ of flat sandy bottom, had been surveyed 30 times with SCUBA prior to sampling for wolffish. The area was found to be generally void of any macro- epibenthos commonly eaten by wolffish, i.e. crabs, gastropods, and sea urchins (Bigelow and Schroeder, 1953). Wolffish inhabiting this area were collected by a diver who speared the fish and returned to the support craft where fresh weight {+/- 0.1 kg) overall length (+/- 0.5 cm) and sex were recorded. Whole intact stomachs were then removed and fixed in 10% formalin in seawater. In the laboratory the total weight of the stomach contents (excluding liquids) was first determined. Subsequently, stomach contents were sorted and recognizable materials were pooled prior to weighing ( + /- 0.1 g wet weight). The number and size of quahaugs eaten by wolffish was then determined. Recognizable umbo portions of valves were separated from stomach contents through careful examination of shell fragments. The number of paired umbones was considered representative of the number of quahaugs eaten. The original size of the live quahaugs eaten was determined by matching the fragmented umbo pieces with other valve fragments to assemble whole or par- tial valves. Size was attained by superimposing on the whole intact valves of a known size. Suc- cess in assembling the valves contributed to con- fidence in determining the numbers and sizes of the quahaugs consumed. From previous re- search on the biology of Ardica islandica by Rowell and Chaisson (1983), relationships were developed for animal size (overall length in cm) to age and whole fresh weight to equivalent meat weight. Estimates of the total fresh weight as well as equivalent meat weight con- sumed by each fish was then determined from the number and size composition of quahaugs in gut contents. The general, solitary, non-schooling nature of wolffish (Bigelow and Schroeder, 1953) is sup- ported by our findings that few wolffish were caught at our study site. We encountered wolf- fish in only four of twelve SCUBA surveys over three months of study (June 1983; May, July 1984). In addition, we noted that once a wolffish was removed another occupied the former resi- dence position by our return. This may indicate that wolffish occupy and possibly defend large territories which may exceed the 4000 m^ area 128 THE NAUTILUS October 31, 1986 Vol. 100(4) of our study site. Only five woiffish were caught for stomach content analysis (Table 1). The largest individual measured 101.6 cm in overall length and weighed 12.0 kg, the smallest was 72.6 cm and 4.4 kg wet weight. Four fish contained easily identifiable quahaug shells (Arctica islandica), comprising 85-99% of the total gut content wet weight (Table 1). The remaining foods (1-15%) were trace amounts of periwinkles {Littorina sp.) sea urchins (Stronglyocentrotus droe- hachiensis) and unidentifiable crustacean exo- skeleton. The fifth fish was packed, to a point where the stomach was distended, with finely crushed tests of the sanddollar Echinarchinus parma. After assembling shell fragments for qualita- tive analysis, we found from 3-11 quahaugs in the gut contents of a single fish (Table 1). The size of ingested quahaugs ranged from 35-86 mm in shell length, representing quahaugs ap- proximately 4-60 years old. Qualitative estimates of total fresh weights of quahaugs in- gested per fish ranged from 255.5-737.7 g; meat weights were from 81.1-155.3 g, respectively. Meat weights expressed as a percentage of total fish weight (Table 1) indicated that only 0.7- 2.2% of fish live weight is a food ration in the form of quahaug meat. It is realized that the sample size is small, but the fact that ocean quahaugs comprised over 85% (by wet weight) of the diet of these woiffish is of interest. This is supported by the insignifi- cant amounts of macro-invertebrate fauna in the stomaches of fish with quahaugs. In con- trast, the sanddollar {E. parma) constituted 100% of the diet of one fish. This finding sup- ported our reconnaissance of the study site which revealed a general lack of macro- epibenthos. These results further suggest a possible selection preference for quahaugs by woiffish, although other items of marginal food value are also eaten. The slow growth oi Arctica islandica (Murawaski et al. 1982) makes it available to woiffish for 50-60 years after the clam settles on the bottom. This is a long period of potential exposure to predation by woiffish, l)ut the duration of the life span infers low natural mortality. This in part retlects low levels of woiffish al)undance, their ability to locate and capture prey as well as the clams ability to avoid CO > o Z S >■, 13 c 3 c/: < ^ ^ %-s. 3 !i C 0) 3 ^ O 4t 8S 5!_ x: 4) • — O C 4J ~- e %~ fN < CO r- . \ . . -N 2 •- O f^ tn r~ rf o in ■- r- 1 >- in CD CO en r- i£) v£i in r- rs) •? o o *T o O^ li) •- I*) o f*l O^ 'T CD r^ ^ o vD CD r- o> 1-1 ^ O I CD Z r- o Z o •- •- o o •- (N m «» in Ul Vol. 100(4) October 31, 1986 THE NAUTILUS 129 predation. Under the assumption that a minimum daily ration of only 5% body fresh weight is required for maintenance (pers. comm. J. Castell, DFO, Halifax, N.S.) the consumption of quahaugs by wolffish in this study was considerably lower than this minimum value and ranged from 0.7 to 2.2% (Table 1). This is probably a reflection of low availability of ocean quahaugs in the study area. Rowell and Chaisson (1983) estimated that the mean density of ocean quahaugs in the im- mediate study area was 3.8 m^ Over a much larger area of commercial potential in the same region (12 km^) they estimated a mean density of 5.4 m\ In the study area, the results indicated that wolffish probably prey on quahaugs equal to the number in one m^ each day. Consequent- ly, even in low abundance wolffish may exert considerable impact on the commercial densities of quahaugs. It's impact on quahaug beds in areas suitable for commercial exploitation may be higher but more research is needed to sup- port such a hypothesis. Acknowledgments We wish to thank P. Woo, I&MP, DFO; D. Chaisson, Bio-Atlantech; and J. Ropes, U. S. Dept. Commer., NOAA, NMFS, Woods Hole, Mass., for their assistance in sample collection, analysis and manuscript review. LITERATURE CITED Arntz, W. E. 1974. A contribution to the feeding ecology of juvenile cod (Gadus morhua L.) in the western Baltic. Rapp. P.V. Reun. Cons. int. Explor. Mer. 166:13-19. 1978. The food of adult cod {Gadun morhua L.) in the western Baltic. Meeresfarsch. 26(1977-78):60-69. Bigelow, H. B. and W. C. Schroeder. 1953. Fishes of the Gulf of Maine. Fish Bull. U.S. 53, 577 pp. Bowman, R. E., R. 0. Maurer Jr. and J. Murphy. 1976. Stomach contents of twenty-nine fish species from the five regions in the Northeast Atlantic. Data report Northeast Fisheries Centre, Woods Hole. Mass. Lab. Ref. No. 76-10, 37 pp. Gill, T., 1911. Notes on the structure and habits of the wolf- fishes. Proceedings U.S. National Museum 39(1782): 157-187. Murawski, S. A., J. W. Ropes, and F. M. Serchuk, 1982. Growth of the ocean quahaug, Arctica islandica. in the Middle Atlantic Bight. Fish Bm.//.80:21-34. Ropes, J. W. and S. A. Murawski. 1983. Maximum shell length and longevity in ocean quahaugs, A rctica islandica. Linne. ICES/CM. 1983/K:32, Shellfish Comm. 8pp. Rowell, T. W. and D. R. Chaisson. 1983. Distribution and abundance of the ocean quahaug {Arctica islandica). and Stimpson's Surf Clam {Spisula polymyma) resource on the Scotian Shelf. Can. Ind. Rep. Fish. Aquatic Sci. 142:v + 75 pp. Verrill, A. E. 1871. On the food habits of some of our marine fishes. Amer. Nat 5:397-400. ATTENTION: RESEARCHERS AND SMELL COLLECTORS. ANNOUNCING MALACOLOGY DATA NET ECOSEARCH SERIES A new and unique journal specializing in rapid publication of worthy papers on marine, freshwater, and terrestrial moUusks. Special attributes are: > 6 numbers per year (150-175 pages); > Peer review and publication within 8 weeks; > Inexpensive: $14.50/year, $18.00 foreign; > Exchange notices (up to 30 words) printed free, once a year, for subscribers; Most journals take a year to publish manuscripts. That is too long. By facilitating rapid publication. Data Net provides unusual opportunities for timely interactions between amateurs who are willing to contribute specimens and experience with professionals who are at the cutting edge of science. Please mail in your order today. Make your check payable to ECOSEARCH, Inc. This new venture needs your support. Thank you. Arthur II. Clarke, Editor (former AMU President and Associate Curator, Smithsonian Institution) ECOSEARCH, IMC. Etokigital Sufveyi ^nd Rese<*(ch 325 E. Bayview, Portland, Tx. 78374 130 THE NAUTILUS October 31, 1986 Vol. 100(4) FRESHWATER MOLLUSKS OF THE DOMINICAN REPUBLICi Jose D. Gomez, Mercedes Vargas and Emile A. Malek Universidad Autonoma de Department of Tropical Medicine Santo Domingo Tulane Medical Center Dominican Republic New Orleans, LA 70112 ABSTRACT A survey of the freshwater mollusks of the Dominican Republic revealed the presence of 27 species, 2U of them, gastropods and 3 bivalves. There were 10 species of the family Planorbidae and of these k were species o/Biomphalaria, namely, B. glabrata, the intermediate host of Schistosoma mansoni, B. havanensis, B. helophila, and B. straminea. The role of the latter 3 species in transmission of schistosomiasis in the country is under investigation. Other families found were Lymnaeidae, Physidae, Ampullariidae and Thiaridae with 2 species each, and Ancylidae. Hydrobiidae and Pleuroceridae with 1 species each. There were 3 species of the Neritidae. There have been only a few reports on the freshwater mollusks of the Dominican Republic. Probably the first was by Crosse (1891), who listed the land and freshwater snails of the island of Santo Domingo (Hispaniola), but the nomenclature he used naturally does not agree with that used today. Other writers have reported the occurrence of the snail Bi.07nph.a- laria glabrata (Say, 1818) in either a few or several habitats (Ponce Pinedo, 1947; Olivier et al., 1952; Etges and Maldonado, 1969; Schneider et al., 1985), and still others have listed a few freshwater mollusks associated with B. glabrata (Gomez, 1973; Vargas and Gomez, 1976; Schneider et al., 1985). The lymnaeid in- termediate hosts of Fasciola hepatica were reported upon by some workers (Alvarez and Cordeiro, 1977; Gomez et al., 1986). In this paper we include the results of surveys for the freshwater mollusks of the country car- ried out for the last five years by the Institute of Research in Bilharzia of the Autonomous Uni- versity of Santo Domingo and by the junior author. Materials and Methods Hydrographic maps were consulted to locate •Contribution from the Institute de Investigaciones en Bilharzia, U.A.S.D. Dominican Republic and the World Health Organization Collaborating Center for Applied Medical Malacology, Tulane Univer.sity. freshwater bodies and detailed sectional regional maps were prepared to locate the various habitats. Identification of the specimens was based on information from the Pan American Health Organization (PAHO) Guide (1968), Malek (1985), and various literature on mollusks from the Great and Lesser Antilles. Included in the information obtained for each body of water was the associated fauna and flora as well as the rainfall cycle and other physical and chemical factors influencing the mollusks; these data will be reported in a separate paper. Results Twenty-four species of freshwater gastropods were collected during this survey; these be- longed to nine families (Table 1). Three species of bivalves were also found. Among the gastropod families there were 10 species of Planorbidae, of which Biomphalaria helophila (Orbigny, 1835) was the most common (31 habitats), B. glabrata. intermediate host of Srhisto.'^oma mansoni (26 habitats), followed by B. hnvanensis (Pfeiffer, 1839) (15 habitats), and B. straminea (Dunker, 1848) (1 habitat). B. helophila and B. hnvanensis are potential hosts for S. mansoni. and B. straminea is a known transmitter of this schistosome in South America. Other families encountered were Lymnaeidae, Physidae, Ampullariidae and Thiaridae with 2 Vol. 100(4) October 31, 1986 THE NAUTILUS 131 TABLE 1. Families and species of gastropods and their loca- tions in the Dominican Republic. Planorbidae Biomphala ria glnhrnta Hato Mayor; Santo Domingo; La Vega; San Cristobal; Bayaguana; Haina; El Seibo; Sabana de la Mar; Guerra; San Jose de los Llanos; San Pedro de Macoris; San Fco de Macoris; Arenoso; Higuey; Miches; Nisibon; Constanza; Jarabacoa; Cotui; San Rafael del Yuma; La Romana; Ramon Santana; Santiago; Quisqueya; Nagua; El Valle. B. havanensis Santo Domingo; Haina; Bani; Palenque; S. J. de la Maguana; Las Matas de Farfan; La Romana; Villa Vas- quez; S. P. de Macoris; La Vega; Valverde; Puerto Plata; Esperanza; Navarrete; Villa Gonzalez. B. helophila Santo Domingo; Haina; Valverde; Las Matas de Farfan; S. J. de Maguana; Barahona; Cabral; Las Marias; Neyba; Batey 4 and 5; Guerra; La Vega; Sabana Rey; Seibo; Hato Mayor; LA, Yabiquin River; LA, Yugery River; LA, Yonu River; Miches; Sta. Lucia Stream, Seibo; Lebron Stream. Seibo; Pana Pana River; S.F.M., Juda River; S.F.M. El Gran Estero; S.F.M. Arenoso; Bhona. Los Patos; Bhona, Quarantitre River; La Vega. .Juma River; Ducantamiento River; Neyba; La Cabirma, Neyba. B. straminea Los Llanos, SPM. Helisomti trivolvis Valverde Mao (canals in rice fields). H. foveale Santo Domingo, Distrito Nacional; concrete-lined foun- tains, Centro de los Heroes; concrete-lined fountains. Natural History Museum; Cabral Pond, Barahona. Drepaiiotrema tucidum Haina; S. J. Maguana; Cabral; Villa Vazquez; La Vega; Puerto Plata; Mao; San Raf. del Yuma; Guerra; S.F.M., Casa de Altos; S.F.M.. La Estancia; S.F.M, El Aguacate; S.F.M.. Arenoso. D. cimex Constanza; Haina; Gabon Pond, Haina; La Vega, El Ran- chito; La Vega, La Cabulla; La Vega, Sabana Rey; Val- verde, Guayacanes; Mao; Yerba de Guinea. Mao; Puerto Plata; Villa Vazquez; Las Lagimas, Santiago; Moca; Guerra, D.N.; S.F.M., Arenoso; D.N., Botanical Gardens. D. anatinurn D.N., Botanical Gardens; Haina; Km. 9 Aut, Duarte D.N.; Villa Mella; Mao; S.J. de la Maguana; Cabral; Villa Vazquez; Gabon Pond, Haina; Ranchito, L.V,; Tabagiia; Puerto Plata; Dajabon; Moca; San R. del Yuma; Guerra; Nisao; S.F.M., casa de Alto; San Fco. de Macoris; S.F.M. Arenoso; S.F.M., El Gran Estero; Bonao; San Pedro de Macoris. D. aeruginotiiim La Vega, El Ranchito. Physidae Physa cubensis Constanza; La Vega, Gima; La Vega, Sabana Rey; Guayacanes, Mao; Tabagua; Cerro Gordo, Mao; Los Pinos, Mao; Haina. P. morrnorata D.N., Botanical Gardens; Haina; Hda. La El.strclla; Bani; Sabana Grande de Palen(|ue; Guayacanes. Mao; Las Matas de Farfan; Neyba, Batey 4 and .5; Biran Stream, Barahona; Azua; Pedregal, Bani; Jarabacoa; Guayubin; Villa Vaz- quez; La Vega; Mao; Tabagua; Puerto Plata; Moca River; Juan Lopito, Moca; Hato Mayor; Martin Avila, LA; Anguilla River, El Seibo; Yonu River, LA; Miches; Santa Lucia Stream; El Seibo; Lebron Stream, Seibo; Guerra; Don Gregorio Pond, Nisao; Nisao; San Fco. Macoris, Casa de Alto; SFM, La Estancia; SFM, El Aguacate; SFM, Arenoso; Juma; Bonao. Lymnaeidae Fossaria cubensis D.N.. Botanical Gardens; Constanza; Haina; KM. 9 Duarte Highway, D.N.; Sabana Grande de Palenque; Valverde; Santiago; San J. de la Maguana; Barahona; Las Matas de Farfan; Azua; Bani; Jarabacoa; La Vega; Esperanza, Mao; Villa Vazquez; Hato Mayor; Miches; Seibo; San Fco de Macoris; Bonao. Pseudosuccinea coin mella Quita Sueno Pond, Haina; D.N.. Botanical Gardens. Ancylidae Ferrissia irrorata Mao River, Mao; D.N., Botanical Gardens; Cabral; Seibo; San Cristobal; San Juan de la Maguana. Thiaridae Thiara granifera and T. tuberculntn D.N., Botanical Gardens; La Vega; Santiago; San J. de La Maguana; Paraiso, Bhona; Cabral; Descubierta; Las Barias; Jimani; Neyba; Pedernales; Haina; Bani; Azua; Jarabacoa; Guayubin; Villa Vasquez; Sabana de la Mar; Miches; Nisibon; Seibo; Higuey; La Vega; San Pedro de Macoris; San Fco. de Macoris; Guerra; Hato Mayor; Moca; Dajabon; Ramon Santana; Puerto Plata; Nagua; Castillo: Mao; Samana; Sanchez; Bayaguana; Nigua. Ampullariidae Marisa comuarietis D.N., Botanical Gardens; Guerra; Bayaguana; San Pedro de Macoris; La Vega; Jarabacoa; Nigua; Hato Mayor; Nisibon; Valverde; Nagua. Ampullaria glnuca Los Llanos; D.N., Botanical Gardens; Hato Mayor. Hydrobiidae Pyrgophorus parvulus D.N., Botanical Gardens. Pleuroceridae Goniobasis sp. Nigua; D.N.. Botanical Gardens; Nisibon. Neritidae Neritina sp.; N. pmictulata: N. virgiriea Puerto Plata; Haina; Santo Domingo; Nisibon. species each, and Ancylidae, Hydrobiidae, and Pleuroceridae with 1 species each. Three species belonging to the Neritidae were encountered, and 3 species of bivalves: Pisidium puncti- forum. Eupera cubensis. and Anodonfa sp. The first 2 species are sphaeriids, family Sphaeri- idae, and the third belongs to the family Unionidae. 132 THE NAUTILUS October 31, 1986 Vol. 100(4) More than one mollusk shared the habitat with at least one other moliusi<. Thiara spp., however, did not occur together with B. glabrata, but Marisa cornuarietis (Linnaeus, 1758) did inhabit the same waterbody as B. glabrata. The flora and fauna in each habitat did not limit the distribution of mollusks. On the other hand, aquatic vegetation favored the oc- currence of most of the species. However, Thiara spp. always favored habitats with little or no vegetation. Discussion Until 1972 B. glabrata occurred in a total of 1 1 habitats in the country (Vargas and Gomez, 1976), whereas we found it in 26 habitats in the present survey, indicating an apparent continu- ing dissemination. In addition to natural factors which favor spreading, the movement and intro- duction of ornamental plants in parks and other locations have apparently contributed to the dispersal of this medically important species. B. haimnensis and B. helophila also show wide distribution, possibly because of their ability to adapt to habitats in the dry region of the coun- try. B. glabrata has not yet invaded areas in the south. B. havanensis and B. helophila are poten- tial hosts of 5. mansoni, and specimens of B. havanensis from Haiti have been infected ex- perimentally (Michelson, 1976). Specimens of B. helophila from Puerto Rico have also been in- fected experimentally (Richards, 1961, 1963). While this paper was in preparation a study by Sodeman et al. (1985) was published in which they collected 6 specimens only of B. straminea. in the river Iguamo. We have had the opportuni- ty to collect close to 1000 specimens of B. straminea from Los Llanos, a different locality but in the same general area of San Pedro de Macoris. Our specimens showed the characteris- tic features of this species (PAHO, 1968; Malek, 1985), that is, those of the shell, of the penial complex, and corrugations on the vaginal sur- face, but these corrugations were not distinct in all specimens dissected. The previous known geographical range of B. straminea was Mar- tinique, Costa Rica, and South America (Brazil, Venezuela, Guiana). We are at present testing the susceptibility of our Dominican specimens to infection with S. mansoni. It seems that Helisoma trivolvis (Say, 1817) was introduced and became established in the northeast section of the country, and now oc- curs in large numbers. However, Helisoma foveale (Menke, 1830) is mainly encountered in small artificial habitats, such as small pools and fountains, indicating its possible introduction with tropical ornamental fish. B. glabrata now does not occur in the northeast, and competition may have occurred between Helisoma trivolvis and B. glabrata, as was demonstrated in the laboratory (Malek and Malek, 1978). There were 4 species of the genus Drepano- trema; the first was D. lucidum (Pfeiffer, 1839) and we believe that it is not D. hoffmani F. C. Baker, 1940. This is because, first, our specimens were never more than 7.5 mm in diameter, and second, it seems that/), hoffmani was described by F. C. Baker based on large specimens of D. lucidum. The lymnaeid Fossaria cuhensis (Pfeiffer, 1939) has been known to occur on the island since the time of Crosse, and is a transmitter of the liver fluke, Fasciola hepatica. Another lym- naeid, Pseudosuccinea columella (Say, 1817), was recently reported to occur in the Dominican Republic (Gomez et al., 1986). This latter species is an experimental host for i^. hepatica in Puerto Rico (Leon-Dancel, 1970), and is a natural host in Brazil (Ueta, 1980). Thiara granifera (Lamarck, 1822) and T. tuberculata (Muller, 1774) have in recent years been introduced into the Dominican Republic as well as other Caribbean islands. Their popula- tions share the same habitats and they can be confused morphologically, and thus are often reported in the Caribbean literature as one species, T. granifera. We have examined specimens from Puerto Rico, Guadeloupe, and Martinique and they all represent mixed popula- tions but predominantly T. tuberculata. The junior author expressed these views in a short note (McCuUough and Malek, 1984). In a recent publication (Starmuhlner, 1985), Thiara tuber- culata was reported from Guadeloupe, Dominica and Martinique. Starmuhlner made an error in two of his figures; plate 8, Figures 6, 8 and 9 are T. tuberculata, but Figure 7 and especially Figure 10 are T. granifera. The two species can be differentiated as follows: in T. granifera the shell is characterized by distinct and raised axial ribs reticulating with spiral threads. Whorls of Vol. 100(4) October 31, 1986 THE NAUTILUS 133 the spire are rather flat-sided; body whorl is wide and its height is more than half of total height of shell. In T. tuberculata the shell usual- ly has low axial ribs and distinct and raised spiral cords especially on body whorl; sometimes whorls are nearly smooth with only incised spiral threads. Whorls are evenly rounded; body whorl is less in width than in T. granifera and its height is less than half of shell height; the shell is more slender than that of T. granifera. and is uniformly turreted. The position of the brood pouch is similar in both species, as well as the mantle edge bearing finger-shaped fringes in both. Alvarez and Mena Sanchez (1973) re- ported that T. granifera had been introduced in- to the eastern part of the Dominican Republic and at present it has spread to other parts of the country (Vargas et al., 1982). There may now be biological control of B. glabrata by competition with T. tuberculata and T. granifera. Some habitats previously occupied by B. glabrata now harbor only the two melaniid species. According to Ferguson (1978), T. granifera competes suc- cessfully with B. glabrata in Puerto Rico. In St. Lucia, four field trials demonstrated that B. glabr-ata was eliminated from marshes and streams 6-22 months after the introduction of T. granifera (Prentice, 1983). Marisa corriuarietis. an operculate ampul- lariid, was introduced into the Dominican Republic by the personnel of the Health Depart- ment in charge of the control of schistosomiasis as a competitor of B. glabrata. because of the encouraging result in nearby Puerto Rico (see review by Ferguson, 1978). In our experience the two snails now occur together in some habi- tats in the Dominican Republic, and if M. cor- nuarietis did succeed in biological control in Puerto Rico this has taken place only in certain situations, for example, small ponds and reser- voirs. The bivalves of the Dominican Republic have not received much attention. Anodonta sp. (family Unionidae) was recently introduced into the country and is now present in the fish {Tilapia and carp) ponds of the Ministry of Agriculture in Nigua, close to the capital, Santo Domingo. The main species of freshwater mollusks of the Dominican Republic are also present on cer- tain other Antillean islands. In Haiti, which shares the island of Hispaniola with the Dominican Republic, Robart et al. (1976) reported 16 species; in Puerto Rico, van der Schalie (1948) reported 14 species; In Guadeloupe, Pointier (1974) listed 21 species, and in Martinique, Guyard and Pointier (1979) listed 19 species. In this paper we report 27 species to be present in the Dominican Republic, but some of these are of recent introduction. LITERATURE CITED Alvarez, J. M. V., and Cordero, E. T. C. 1977. Fascioliasis bovina en la Republica Dominicana. Publ. Universidad Autonoma S. Domingo. Vol. 224. Col. Ciencia y Tech. No. 5:1-189. Alvarez, J. M. V., and Mena Sanchez, V. 1973. Hallazgo en Dajabon, Republica Dominicana del caracol de agua dulce "Thiara" spp. de importancia medica. Revista de la Faciiltad de Ciencias Agroitomicat: y Veterinarias 2:.53-.54. Crosse, H. 1891. Faune malacologique terrestre et fluviatile de rile de Saint Domingue. Jour. Conrhyliol. 39:69-211. Etges, F. J., and Maldonado, J. R. 1969. The present status of bilharziasis in the Dominican Republic. Malacologia 9:40-41. Ferguson, F. F. 1978. The role of biological agents in the control of schistosome-bearing animals. Atlanta, Georgia, U.S.A., U.S. Department of Health, Education and Welfare, Center for Disease Control, 107 pp. Gomez, J. 1983. Distribucion geografica del Biomphalaria glabrata (huesped intermediario del Schistosoma mansoni) en Republica Dominicana. Thesis for Licenciado en Biologia, Universidad Autonoma de Santo Domingo. Gomez, J., Vargas, M. and Malek, E. A. 1986. P-'seudosiic- cinea columella (Lymnaeidae) in the Dominican Republic and transmission of fascioliasis in the Caribbean region. The Nautilus 100:66-68. Guyard, A. and Pointier, J. P. 1979. Faune malacologique dulcaquicole et vecteurs de la schistosome intestinale en Martinique (Antilles Francaises). Ann. Parasitol. (Paris) 54:193-205. Leon-Dancel, D. 1970. Life history of Lymnaea columella (Say) and its e.xperimental infection with Fasciola hepatica (L.). Jour. Agric. Umv. Puerto Rico. 54:297-305. Malek, E. A. 1985. Snail hosts of Schistosomiasis and Other Snail-transmitted Diseases in Tropical America: A Manual. Pan American Health Organization, Washington, D. C, Scientific Publication No. 478, 1-325. Malek, E. A., and Malek, R. R. 1978. Potential biological control of schistosomiasis intermediate hosts by helisome snails. The Nautilus 92:15-18. McCullough, F. S., and Malek, E. A. 1984. Notes on the mol- luscan intermediate hosts oi Paragonimus spp. and their possible role in biological control. Ann. Trap. Med. Para- sitol. 78:339-340. Michelson, E. H. 1976. A potential intermediate host of Schistosoma mansoni from Haiti. Jour. Parasitol. 62:648-649. Olivier, L., Vaughn, C. M. and Hendricks, J. B. 1952. Schis- 134 THE NAUTILUS October 31, 1986 Vol. 100 (4) tosomiasis in an endemic area in the Dominican Republic. ArMT. Jour. Trop. Med. Hyg. 1:680-687. Pan American Health Organization. 1968. A Guide for the Identification of the Snail Intermediate Hosts of Schisto- somiasis in the Americas. PAHO Scientific Publication No. 168, 1-122. Pointier, J. P. 1974. Faune malacologique dulcaquicole de rile de la Guadeloupe (Antilles Francaises). Bull. Mus. Hisl. Natur.. Paris, 3rd Ser., No. 235, Zool. 159:905-933. Ponce Pinedo, A. M. 1947. Schistosomiasis mansoni in the Republic of Santo Domingo with a report of six cases studied. Puerto Rico Jour. Publ. Hlth. Trap. Med. 22:308-324. Prentice, M. A. 1983. Displacement of Biomphalarin gin- brata by the snail Thiarn granifera in field habitats in St. Lucia, West Indies. .Ann. Trop. Med. Parasitol. 77:51-59. Richards, C. S. 1961. Another potential intermediate host snail for Schistosoma mansoni in Puerto Rico. Jour. Parasitol. 47:64. 1963. Infectivity of Schistosoma mansoni for Puerto Rican mollusks, including a new potential inter- mediate host. Ainer. Jour. Trop. Med. Hyg. 12:26-33. Robart, G., Mandahl-Barth, G. and Ripert, C. 1976. Inven- taire, repartition geographique et ecologie des mollusques dulcaquicoles d'Haiti (Caraibes). Hnliotis. 8:159-171. Schneider, C. R., Hiatt, R. A., Malek, E. A. and Ruiz-Tiben, E. 1985. Assessment of schistosomiasis in the Dominican Republic. Public Hlth. Rejjorts 100:524-530. Sodeman, W. A., Rodrick, G. E., Paraense, W. L. and Vargas de Gomez, M. 1985. Biomphalaria straminea and other planorbids in the Dominican Republic. Mem. Inst. Oswaldo Cruz, Rio de Janeiro 80:453-456. Starmuhlner, F. 1985. Erstfunde von drei Arten von Suss- wasser Gastropoden auf den Inseln Guadeloupe, Dominica und Martinique (Kleine Antiilen). Heldin 1:55-58. Ueta, M. T. 1980. Ocorrencia de infeccao natural Ae Fasciola hepatica Linnaeus, 1758 em Lymnaea columella Say, 1817 no Vale de Paraiba, SP, Brasil. Re^i. Saude Publica 14:230-233. Van der Schalie, H. 1948. The land and fresh water mollusks of Puerto Rico. Misc. Publ. Mus. Zool. Univ. Michigan, No. 70. Vargas Castro, M., and Gomez, J. 1978. Algunos aspectos de la distribueion del Biomphalaria glabrata en la Republica Dominicanas. Ciencia, Organo de la Direccion de Investi- gaciones Cientificas de la U.A.S.D., 3:87-94. Vargas, M., Gomez, J. D. and Guerrera, N. 1982. Distribu- eion en la isla de Santo Domingo de Thiara granifera (Lamarck), Mollusca: Melanoide. Rev. Dam. Microbiol. Parasitol., 1:14-18. A PREHISTORIC ABORIGINAL FRESHWATER MUSSEL ASSEMBLAGE FROM THE DUCK RIVER IN MIDDLE TENNESSEE Paul W. Parmalee and Walter E. Klippel Department of Anthropology University of Tennessee Knoxville, TN 37996 ABSTRACT A tofiil of 761 valves of freshwater mussels, representing a minimum of 28 species, was identified from a prehistoric ahoriginal cave site located along the Duck River, Maury County, Tennessee. Two thirds of the mussels are Cumber- landian forms and. as a group and based on known habitat requirements, suggest that this stretch of the Duck River for some period between ca. 7.000 and 1,000 BP was more shallow than at present with numerous riffles and swift current. Recovery of numerous specimens of Pegias fabula from aboriginal sites located along a ca. 150 km stretch of the Duck River irfects an extensive prehistoric population of this mussel in Middle Tennessee. This prehistoric assemblage is discussed in relation to Duck River m.ussel faunas reported over the last 60 years. Several small rockshelters situated in the Duck River bluffs were tested for aboriginal occupation during archaeological survey work in 1978 in the proposed Tennessee Valley Author- ity Columbia Reservoir area. One of these, Cheek Bend Cave (40MU261), is located approx- imately 13 km ESE of the city of Columbia, Maury County (Fig. 1), and testing (three 1 x 2 m excavation units) showed it to contain stratified deposits of bone and shell to a depth of approxi- Vol. 100(4) October 31, 1986 THE NAUTILUS 135 TENNESSEE 10 20 JOk FIG. 1. The Duck River in Middle Tennessee with the loca- tion of Cheek Bend Cave and other aboriginal sites contain- ing naiads discussed here. mately 4.5 m. On the basis of the vertebrate species composition within and among strata, differing fill zones, and certain other stratigraphic features, the deposit reflects two distinct and well-defined major episodes of fill. The top 2 m represent the Holocene stage and contain remains of extant modern species. In contrast, the faunal assemblage in the bottom 2 m, although it includes some species that still inhabit the cave area, contains many such as the prairie chicken, 13-lined ground squirrel and pocket gopher that are now extirpated and reflect a prairie habitat (Parmalee and Klippel, 1981) and/or a boreal environment, species such as red-backed and yellow-cheeked voles, red squirrel, northern flying squirrel, and arctic and water shrews (Klippel and Parmalee, 1982). Matrix from two additional one-meter-square columns was removed during 1982 and 1983. Only the vertebrate remains from the Pleisto- cene strata have undergone preliminary iden- tification, but all freshwater mussel valves from the Holocene strata were removed and incor- porated with the 1978-1979 specimens for this study. Although terrestrial gastropods occurred in all strata, aquatic gastropods, freshwater mussel shells and prehistoric human artifacts were encountered only in the Holocene strata. Mussels were commonly used as a food resource by prehistoric Indians in eastern North America (Parmalee and Klippel, 1974) and in all proba- bility most if not all of the naiads present in this cave deposit were gathered by aboriginal groups occupying the site. The majority of valves were well-preserved with some in the top two strata still retaining remnants of the periostracum, but also a large number of shells were broken with only portions of the hinge line remaining. Because of this lat- ter condition, an additional 555 valve fragments could not be identified (Table 1). All species determinations were made using comparative reference specimens in the collections of the Section of Zooarchaeology, Department of Anthropology, University of Tennessee, Knox- ville; all mollusk and vertebrate remains recovered from Cheek Bend Cave are housed in the Department of Anthropology. The Duck River and Molluscan Studies Interest by American malacologists in the varied and abundant freshwater molluscan fauna once present in most major river systems in eastern North America began well over a cen- tury ago. These beginning efforts centered around taxonomy, and emphasis was placed on describing and naming species new to science. By the 1850s the majority of "new" species had been described and the tendency then was to treat the total molluscan fauna of a specific region or river system. List of Shells Collected in Central Tennessee by Hinkley and Marsh (1885) is only one of many such references, but we cite it because it provides not only a list of species known from the Duck River (". . . the Duck River and adjacent country at Columbia, Maury Co.") at that time but also makes reference to the abundance of many species and the habitat in which they occurred. In addition, the following description by Hinkley and Marsh (1885:2) seems appropriate: "The Duck River, at Colum- bia, is a pretty and rather picturesque stream, the limestone bluffs along its banks having been sculptured by the action of rain and frost into various shapes, in places overhanging the water and forming quite a cavern underneath. The water is shallow, swift, and clear. . ." See Fig. 2. After collecting the Duck River in 1921, 1922, 136 THE NAUTILUS October 31, 1986 Vol. 100(4) FIG. 2. View of the Duck River, Maury County, winter 1978-79, approximately 100 m upstream from Cheek Bend Cave. and 1923, Ortmann (1924) published a compre- hensive treatment of the naiad fauna that in- cluded 63 species and forms which he confirmed and 7 that "should be credited to the Duck River" based on previous published reports. More recent studies by Isom and Yokley (1965) and van der Schalie (1973) have pointed up the fact that the Duck River naiad assemblage has now been reduced by nearly one-fourth to about 45 species or forms. Completion of the Columbia dam and reservoir will bring about the extirpa- tion of most species in the effected stretches of the river and will probably eliminate two species, now known to inhabit only this locale, from the entire river system. Ahlstedt (1981), in a recent survey of the molluscan fauna of the Duck River between the Normandy and Colum- bia dams, found a drastic decline in the numbers and abundance of species compared with earlier surveys in 1965 and 1973. The section of the Duck River above Columbia contains the last known populations in that river of such species as Quadrula intermedia and Lemiox rimosus. In 1982 TVA biologists transplanted ca. 4,000 in- dividuals of L. rimosus into three other rivers (and at one location in the upper Duck River above the Normandy reservoir) in hopes of re- establishing this endangered species in known habitats where it once occurred. Success or failure of this ongoing program has not yet been fully evaluated (S.A. Ahlstedt, pers. commun.). Cheek Bend Cave Mussel Assemblage Although the shell was, for the most part, ex- ceptionally well preserved, the disintegration of the periostracum in the majority of specimens and consequently loss of all color (including the nacre in most) and pattern made specific iden- tification of valves representing certain groups (e.g. Villosa. Epioblasrna) impossible or ques- tionable at best. For example, Ortmann (1924: 52) commented that "The males of D. f = Dysyioyriia = Epioblasm.aJ florentina walkeri, closely resemble those of the next species {cap- saeformis), but they differ chiefly in color." Shells of species as distinct as Lampsilis ovata and Actinonaias pectorosa, when fresh, may be impossible to distinguish when only the dorsal portion of the valve (hinge line with lateral and pseudocardinal teeth) is preserved -as in archaeological or cave contexts. Nevertheless, those specimens that could be identified provide an interesting record of the naiad species that inhabited the Duck River in Middle Tennessee in prehistoric times (on which there is no published data) and which, in turn, reflect river conditions at the time they were collected. Valves of Epioblasrna capsa^formis (some of which may be E. florentina) comprised nearly 20% of the identified naiads recovered in the excavation units; at the time Ortmann col- lected the Duck River at Columbia/Leftwich/ Lillard Mill/Willhoite it was "Rather abundant- New [previously unreported] for Duck River" (Ortmann 1924:53). Although still found in the Duck River above Columbia, it is uncommon and local in occurrence. We collected several dead mature specimens below Lillard Mill that were, unlike those from the cave, extremely large and thick-shelled. The spike, Elliptio dilatata, most of which were small, thin-shelled (for the species) and compressed -suggestive of a shallow small stream habitat - comprised about 12% of the cave naiad assemblage. Ptychobranchus subtentum is another species that was apparently common in the Duck River near the cave site in prehistoric times; 107 valves (14%) were identified from the deposit. It was evidently uncommon at Columbia/Leftwich/ Normandy when Ortmann (1924:40) made his 1921-1923 collections, reporting "Only a few specimens found." His further comment that it is "Most abundant in smaller streams above the range of Ptychobranchui^ fasciolare, and this seems to hold good also in Duck River" is of par- Vol. 100(4) October 31, 1986 THE NAUTILUS 137 ticular interest. This observation, coupled with the numerous valves of P. subtentum. and other species (e.g. Medionidus conradicus) or forms from the cave that are indicative of a head- waters and/or small stream habitat, suggests that that section of the Duck River flowing in close proximity to Cheek Bend Cave during the mid-Holocene was smaller and more shallow than it is today. Another species represented in the cave deposit (Stratum VII), namely Pegias fabula, is also indicative of a fast-flowing, shallow, small stream habitat. Archaeological surface collec- tions were obtained from a Middle to Late Archaic (ca. 7,000-4,000 BP) shell midden at the Ervin Site (40MU147) during 1979. This pre- historic site, situated ca. 29 km upstream from Cheek Bend Cave in Maury County (Fig. 1), pro- duced 15 specimens of P. fabula out of 1339 identified valves (Hofman, nd). During the 1982-1984 excavations at the Hays site (40ML139), ca 40 km upstream from Cheek Bend Cave in Marshall County, huge quantities of aquatic gastropods and freshwater mussel valves were removed from this Middle to Late Archaic (ca. 7,000-4,000 BP) shell midden. In- cluded among the valves of the 32 species iden- tified from this site thus far (Turner, nd) were 104 specimens of P. fabula. Even farther upstream, ca. 134 km above Cheek Bend Cave, Robison (nd) identified 43 valves of P. fabula (from a sample of 2,545 shells) from the Shofner site (40BD55), Bedford County, a Middle Woodland village dating ca. AD 600-500. Archaeological sites on tributaries of the Duck River have also produced P. fabula; one specimen was recovered from Woodland and one from Middle Archaic strata (266 valves iden- tified) at the McCollum Rockshelter (40MU390) along Fountain Creek in Maury County. Two additional specimens (135 valves identified) were recovered from Goatcliff Rockshelter (40MU436) along the same Duck River tributary in what appears to be Woodland context (O'Hare, nd). Excavation of these rockshelters took place during 1980-1981 . A single specimen of P. fabula in the United States Na- tional Museum collections ("Duck River," USNM Lot No. 86229, catalogued Feb. 1888: Paul Greenhall, pers. comm., April 1986) appears to be the only documented historic record of this naiad for the Duck River (Clarke, 1981). In a recent paper Starnes and Starnes (1980) report a viable population of P. fabula from the Little South Fork Cumberland River, Kentucky, a small fast-flowing stream ca. 20-25 m wide with an average water depth of about 20 cm at low river stages. Of additional interest is the fact that the associated mussel assemblage in- cludes P. subtenturn, P. fasciolare, E. dilatata, M. conradicus. Villosa iris, Villosa taeniata, Villosa vanuxemensis and Lampsilis fasciola. Although valves of Alasmidonta viridis, Toxo- lasyna cylindrellus and Lemiox rimosus were not numerous in the Cheek Bend Cave deposit, the presence of these species is also indicative of a small stream habitat or stable stretches of shallow riffles in somewhat larger rivers like the Duck. Stansbery (1976) comments that "Pegias fabula appears to be a rare Cumberlandian species characteristic of stream conditions near, but not quite in, the uppermost headwaters." The three-ridge, Amblema plicata, and purple warty-back, Cyclonaias tuberculata, are two of the more common species inhabiting the Duck River today. The 27 valves of A. plicata com- prised only 3.5% of the total sample but were of interest in that they exhibited characteristics (small and compressed) of those inhabiting a headwaters or small stream habitat. Only two valves of the latter species were recovered in the cave. Lexingtonia dolabelloides, a species fairly well represented in the Cheek Bend Cave naiad assemblage (ca. 6% of the identified valves), still occurs in the Duck River, one of the few remaining rivers still supporting viable populations of this mussel. Quad^-ula inter- media is another species of interest in that it was present in the middle stretches of the Duck River in prehistoric times (2 valves from Cheek Bend Cave) and continued to survive until pre- sent (Ahlstedt, 1981) but at very low population levels. It was apparently never a common shell in the Duck River as evidenced by the paucity of specimens recovered at Cheek Bend Cave and at the Hayes site where only five shells of Q. in- termedia were identified out of a sample of 3,870 valves (Turner.nd). Anomalous Valves During the identification process it was necessary to compare certain archaeological 138 THE NAUTILUS October 31, 1986 Vol. 100 (4) Table 1. Freshwater mussels Identified from Cheek Bend Cave. Maury County. Tenness. Percent of each species within Strata and for Total Valves listed In parentheses { Amblema pllcata (Say. 1817) Three-Rld^e Fusconala barnestana (Lea. 1838) Tennessee Plgtoe Lexington la dolabelloldes (Lea, 1840) Slabslded Pearlyoussel F. bameslana and/or L. dolabelloldes Quadrula CvllQ(^l.ca (Say, 1817) Rabbitsfoot VII 13 (4.19) 5 (1.61) 10 (3.23) 2 (.65) 7 (12.28) 2 (.79) 4 (3.13) 2 (3.5) 4 (1.57) 2 (1.56) A (7.02) 23 (9.06) 1 (.39) 10 (7.81) Total Valves 27 (3.55) 13 (1.71) 47 (6.18) 3 (.39) Quadrula InCenpedla (Conrad, 1836) Cumberland Monkeyface Cyclonalas tuberculata (Raflnesque, 1820) Purple Wartyback Elliptlo dllatata (Raflnesque. 1820) Spike Hemlstena lata (Raflnesque. 1820) Cracking Pearlymussel Pleurobema ovlforme (Conrad. 1834) Tennessee Clubshell Alasmldonta virldla (Raflnesque, 1820) Slippershell Peglas Fabula (Lea. 1836) Llttle-wlnged Pearly Mussel Lasaigona costata (Rafineaque, 1820) Fluted Shell SCrophitus undulatu! Squawfoot Actlnonalas pectorosa (Conrad, 1834) Pheasant she 11 Toxolasma cyllndrellus (Lea. 1868) Pale Lllllput Toxolasma llvldus (Raflnesque, 1831) Purple Lllllput Eploblasma capsaeformls (Lea. 1834) and /or E. fjgrentlna (Lea. 1857) Oyster HusseT i/or Tan Rlffleshell Eploblasma brevldens (Lea, 1834) Cumberland Combshell Lanpsllls fasclola (Raflnesque, 18 Wavy-rayed Lampmussel Lampsllls ovata (Say. 1817) Pocket book Lemiox rimosus (Raflnesque. 1831) Blrdwlng Pearlymussel Medlonldus conradlcus (Lea, 1834) Cumberland Moccasin Potamllus alaCus (Say, 1817) Pink Heelsplitter 1 (8.33) (.65) 10 (3.23) 4 (1.29) 1 (.32) 2 (.65) (.97) 1 (8.33) 17 (5.48) 1 (8.33) 5 (1.61) 4 (33.33) 49 (15.81) 1 (.32) (3.51) 1 (1.75) 10 (17.54) 1 (.39) 1 (.78) 2 (.26) 1 (.78) 2 (.26) 43 (16.93) 9 (7.03) 90 (11.83) 1 (.39) 1 (.78) 2 (.26) 1 (.39) 4 (.53) 1 (.39) 4 (3.13) 8 (1.05) 1 (.78) 11 (1.45) 2 (1.56) 6 (.79) 1 (.13) 4 (1.57) 8 (6.25) 15 (1.97) 1 (.39) 5 (.66) 1 (.39) (.13) 40 (15.75) 16 (12.50) 150 (19.71) 1 (.39) 1 (.13) 10 (3.94) 1 (.78) 14 (1.84) (4.69) 23 (17.97) 13 (1.71) 100 (13.14) Vlllosa Iris (L«a, 1830) Rainbow Vlllosa taenlata (Conrad. 1834) Painted Creekshell Vlllosa vanuxemensls (Lea, 1838) Mountain Creekshell Vlllosa spp. (Eploblasma?) Ptychobranchus faaciolare (Raflnesqui Kldneyshell Ptychobranchus sub ten cum (Say, Fluted Kldneyshell 17 (5.48) (1.29) 10 (3.12) 2 (3.51) 3 (1.18) 2 (1.56) 11 (1.45) 1 (1.75) 35 (13.76) 20 (15.63) 73 (9.59) 2 (3.51) 1 (.39) 1 (.78) 8 (1.05) 1 (1.75) 8 (3,15) 4 (3.13) 24 (3.15) 1 (.39) 1 (.13) 4 (7.02) 58 (22.83) 12 (9.38) 107 (14.06) Totals (Identified valves) Totals (Unidentified valves) (99.97) 17 310 (99.90) 57 254 (99.99) (99.96) 128 761 (100.01) (100.00) Totals (All Valves) Vol. 100(4) October 31, 1986 THE NAUTILUS 139 specimens, particularly incomplete valves and those of Villoma spp. and Epioblnsma spp., with fresh material using a binocular scope. Under magnification, a variety of anomalous struc- tures or conditions were noted that, in addition to making specific determinations difficult or impossible, pose some interesting questions. These anomalies consisted primarily of deep pits along, under and/or between the lateral and pseudocardinal teeth and the development of supernumerary "teeth" or projections between or adjacent to the pseudocardinal teeth (Fig. 3). Less than 3% of the valves exhibited one or more of these anomalies; the majority occurred in small, thin-shelled individuals. Possibly these conditions were the result of some trauma or perhaps parasites, but whatever the cause(s). F^IG. 3. Examples of anomalous naiad valves from Cheek Bend Cave that exhibit pitting and nacre deformities. such abnormalities have not been observed in other archaeological naiad material from pre- historic sites along the Duck River. General Discussion Ortmann (1924:19), in attempting to evaluate the zoogeographical affinities of the Duck River naiad assemblage, made the following signifi- cant observations: "Duck River being located between Cumberland River in Kentucky and Tennessee, and Tennessee River in northern Alabama, might be expected to contain a similar fauna to these two systems, a fauna which is known to contain peculiar elements, not found outside of this region in the Central basin (Mississippi and Ohio drainages), which might be called 'Cumberlandian' types (from the 'Cumberland subregion')." On the basis of his studies of the Duck River mussel species and those of other rivers of the southeastern region, he concluded (Ortmann 1924:61) that "It ap- pears that the Cumberlandian fauna is the original fauna of Duck River, while the interior- basin-fauna is a later invasion of the river, com- ing up from the lower parts; yet important elements of the interior fauna have not yet reached Duck River. In the lower Duck, at Cen- treville, and probably farther down, the interior fauna alone is present, and this part does not belong to the Cumberland region. Duck River originally was more directly connected with the Cumberland and Tennessee, and, at that time, it was a rather small river." Approximately 65% of the species represented in the Cheek Bend naiad assemblage are Cumberlandian as defined by Ortmann, and they reflect a headwaters or small stream habitat. Several other species (e.g. E. dUatata, A. plicnta) present in the cave with Mississippian or Interior Basin affinities exhibit small stream shell characters, that is, com- pressed, thin-shelled small valves. Therefore, the naiad assemblage recovered from the Cheek Bend Cave excavations and the occurrence of certain species such as P. fahula from this cave and other aboriginal sites in Maury, Marshall and Bedford counties suggest that the Duck River in Middle Tennessee was swift and shallow with riffles and a probable substrate of sand and small gravel for extended periods be- tween 7,000 and 1,000 BP. 140 THE NAUTILUS October 31. 1986 Vol. 100(4) Acknowledgments We would like to express our appreciation to Terry Faulkner for preparing Fig. 1 and to Betty W. Creech for typing the manuscript. Special thanks are extended to W. Miles Wright for photographing the specimens in Fig. 3 and the preparation of Figs. 2 and 3. We gratefully acknowledge the Tennessee Valley Authority under whose auspices (contracts No. TVA TV- 49244A and TVA TV-53013A) the excavation of Cheek Bend Cave, plus the analysis of the faunal materials recovered, were made possible. LITERATURE CITED Ahlstedt, S. A. 1981. The molluscan fauna of the Duck River between Normandy and Columbia dams in central Tennes- see. Bull. Amer. Malacol. Union for 1980. pp. 60-62. Clarke, A. H., Jr. 1981. The Tribe Alasmidontini (Unionidae: Anodontinae), Part I: Pegias. Alastnidonta. and Ar«rf«i.s. Smithsonian Contributions to Zoology No. 326. 101 pp. Hinkley, A. A. and P. Marsh. 1885. Lis? of shells collected in central Tennessee. Published privately. Aledo, Illinois. 10 pp. Hofman. J. L. nd. Ervin; A mid Holocene shell midden on the Duck River in the Nashville Basin of Tennessee. Manu- script on file, Department of Anthropology, University of Tennessee, Knoxville, TN. Isom, Billy G. and Paul Yokley, Jr. 1968. The mussel fauna of Duck River in Tennessee, 1965. Airie?'ican Midland Naturalist 80(l);34-42. Klippel, W. E. and P. W. Parmalee. 1982. Diachronic varia- tion in insectivores from Cheek Bend Cave and environ- mental change in the Midsouth. Paleobiology 8(4):447-458. O'Hare, C. R. nd. Goatcliff and McCoUum rock.shelters: analysis of cultural and non-cultural processes operating on the formation of two faunal samples. MA thesis in preparation. Department of Anthropology. University of Tennessee, Knoxville, TN. Ortmann, A. E. 1924. The naiad-fauna of Duck River in Ten- nessee. American Midland Naturalist 9(2): 18-62. Parmalee, P. W. and W. E. Klippel. 1974. Freshwater mussels as a prehistoric food resource. American Anti- quity 39(3):421-434. 1981. A late Pleistocene population of the pocket gopher, Geomys cf. bursarus, in the Nashville Basin, Ten- nessee. Jour. Mammalogy 62(4):831-835. Robison, N. D. nd. An analysis and interpretation of the faunal remains from eight late Middle Woodland Owl Hollow phase sites in Coffee, Franklin and Bedford coun- ties, Tennessee. Ph.D dissertation in preparation, Depart- ment of Anthropology, University of Tennessee, Knox- ville, TN. Stansbery, D. H. 1976. Status of endangered fluviatile mollusks in central North America: Pegias fabula (Lea, 1838). Report prepared for the U.S. Department of the In- terior, Fish & Wildlife Service and the Bureau of Sport Fisheries & Wildlife, Washington, D.C. 8 pp. Turner, W. B. 1984. A preliminary analysis of prehistoric fresh-water bivalve species composition changes from ca. 7500 to 2500 B.P.: a view from the central Duck River drainage. Manuscript on file. Department of Anthro- pology. University of Tennessee, Knoxville, TN. van der Schalie, H. 1973. The mollusks of the Duck River drainage in central Tennessee. Sterkiana 52:45-56. THE FATE OF PELECYPOD FAMILIES, SUBFAMILIES, AND TRIBES DURING AND AFTER THE CRETACEOUS PERIOD David Nicol Box 14376, University Station, Gainesville, FL 32604 ABSTRACT Of the lU surveyed supergeneric groups of pelecypods that occur in the Cretaceous, 38, or 33%, are extinct. In the Early Cretaceous one group became ex- tinct every 9,300,000 years. During the Late, but not latest. Cretaceous, one group became extinct every 2,333,000 years. In the Maastrichtian Stage (latest Cretaceous), the extinction rate increased to one group every 470.000 years. Only seven groups became extinct during the Cenozoic: four during the Eocene and three during the Miocene. There was only one extinction of a supergeneric group every 9,500,000 years during the entire Cenozoic, which is comparable to the ex- tinction rate during the Early Cretaceous. The rate of extinction of supergeneric groups of pelecypods increased rapidly in the Late Cretaceous and reached a climax at the end of that geologic period. The number of families, subfamilies, tribes and groups of pelecypods that undoubtedly had one or more genera living during the Cretaceous Period, as reviewed in the Treatise on Inverte- Vol. 100(4) October 31, 1986 THE NAUTILUS 141 brate Paleontology (1969, 1971), is 124. In each of these supergeneric categories the genera and subgenera were tallied in each geologic Period and Epoch. In a few instances, more current data were used when it was available (Keen's, 1980, latest review of the Cardiidae), and a few of the geologic ranges of other supergeneric groups have been brought up to date. Ten of the 124 supergeneric groups were excluded because of insufficient geologic data, as, for example, the Solemyidae, Manzanellidae, Dimyidae, and Sportellidae. The supergeneric groups that lived during the Cretaceous are, commonly, not ancient line- ages. Only the Nuculanidae and Malletiidae can be traced back to the Ordovician, and they are as diverse today as they have ever been. Fifty three supergeneric groups (47%) appear in the Cretaceous. Some of these are short-lived groups but many others first appeared in the Cretaceous and are presently represented by many genera and species. By the end of the Cretaceous, most of the basic adaptive radiation of the pelecypods had occurred. Of the 114 supergeneric categories considered herein, 90% first appeared in the Mesozoic. Stanley's graph (1968, p. 215) depicts the great Mesozoic radiation. The 114 supergeneric groups can be placed in seven categories based on the distribution of their genera and subgenera from the Cretaceous to the Recent. 1. Many supergeneric groups were relatively unaffected by the mass extinctions at the end of the Cretaceous and remained as diverse in the Cenozoic or increased in diversity during that Era. In this first category there are 46 super- generic groups or 40% of the total of 114. Although some genera became extinct during the Cretaceous, they were quickly replaced by new genera arising in the Cenozoic. Almost 62% of these supergeneric groups began in the Cretaceous and 87% began after the Paleozoic. Those groups first appearing in the Cretaceous are commonly represented by one or two genera. In other words, they are comparatively young supergeneric groups. The protobranch families Nuculidae, Malletiidae, and Nuculani- dae are exceptional in that they range well back into the Paleozoic and were not affected greatly by extinction at the end of the Mesozoic. 2. Some supergeneric groups were affected by Cretaceous extinction but later recovered and became most diverse in the Neogene. There are 15 supergeneric groups that have this distribution in diversity, which comprise about 13% of the total. In this category only six or 40% began in the Cretaceous but 93% began in the Mesozoic. In general, the families are longer ranging than those that were little affected by the Cretaceous extinctions. Some notable ex- amples of supergeneric groups in this category of diversity are the Pitarinae, Tellininae, Car- diinae, Protocardiinae, and Mactrinae. 3. Some supergeneric groups were affected by Cretaceous extinction but continued on to the Recent with a reduction in diversity. In this category are included 15 supergeneric groups or approximately 13% of the total. These super- generic groups are still longer ranging than those in the first two categories and only four, or 27%, began in the Cretaceous. However, 13, or 87%, had their inception in the Mesozoic. There are some notable groups that were adversely affected by the Cretaceous wave of extinction, including the Trigoniidae, which has survived to the present only in the Australian region. The Pholadomyidae is another group that is nearly extinct after its great diversity in the Mesozoic. Other relict groups at present are the Grammatodontidae, Cucullaeidae, Fim- briidae, and Arcticidae. All of these families were much more diverse and widespread during the Cretaceous. Of the 114 supergeneric groups studied, 38, or 33% are extinct. They can be grouped into four categories. 4. A few supergeneric groups became extinct during the Early Cretaceous (before the Cenomanian). There were only five families or about 4.4% of the supergeneric groups that became extinct during this span of time. Eighty percent of this small group are confined to the Mesozoic, which includes the Neomiodontidae, a small family that lived in brackish and fresh water. 5. There are nine supergeneric groups of pelecypods that became extinct in the Late Cretaceous but before the Maastrichtian. This category constitutes 7.9% of the total number of supergeneric groups. Four of these are small and short-lived families and subfamilies that 14:^ THE NAUTILUS October 31, 1986 Vol. 100(4) were confined to the Cretaceous Period. One of these short-lived families, the Trigonioididae, lived in fresh and brackish water. I have placed the Mactromyidae in this category because Bathycorbis Iredale is most likely an aberrant venerid and Dockery (1982, p. 72) has proved that Cordiula Meyer was based on a young specimen of Nemocardiurn. The Mactromyidae is found primarily in the Jurassic and Cretaceous. 6. Seventeen supergeneric groups, about 15% of the total, became extinct at the end of the Cretaceous during the Maastrichtian Stage. Some of the supergeneric groups that became extinct at this time were large-sized, diverse, and common in strata of Cretaceous age. These include the rudist families Hippuritidae, Radiolitidae, Caprinidae, Monopleuridae, and Requiniidae. It also includes the Inoceramidae and Exogyrini. Seven of these seventeen groups (41%) were confined to the Cretaceous Period, and 14 (82%) were found only in the Mesozoic. The great extinction at the end of the Cretaceous did not greatly affect most of the older supergeneric groups. 7. Seven supergeneric groups (6.1%) survived the Cretaceous but became extinct in the Ter- tiary-four in the Eocene and three in the Miocene. Except for the Bakevelliidae and Eriphylinae, they consist of small groups (no more than four genera), and all hut the Bakevel- liidae and Eriphylinae originated in the Cretaceous Period. It is known that extinction rates increased near the end of the Eocene Epoch, as in the planktonic Foraminiferida, and for this reason it is not surprising that a few supergeneric groups of pelecypods disappeared at this time. Two of the three extinctions of supergeneric groups that occurred in the Miocene were the Gryphaeostreinae and the Flemingostreinae (Ostracea) and may have been caused by oncoming glaciation at the end of the Miocene (Nicol, 1984). Fresh-water pelecypods were somewhat af- fected by the Cretaceous extinctions. Besides the two previously mentioned short-lived groups that became extinct in the Cretaceous, the Cor- biculidae and Unioninae lost some genera and subgenera at the end of the Mesozoic. In Table 1 I have summarized the rate of ex- tinction of the supergeneric groups of pele- TABLE 1. Extinction rates of families, subfamilies, and tribes of pelecypods during the Cretaceous and Cenozoic. Stratigraphic time units Number of Extinction and time in years extinctions rate in years 1. Early Cretaceous (Berriasian-Albian) i.0, 500,000 5 9,300,000 2. Late but not latest Cretaceous (Cenomanian- Campanian) 21,000,000 9 2,333,000 3. Latest Cretaceous (Maastrichtian) 8,000,000 17 470,000 4. Late Cretaceous (Cenomanian-Maastrichtian) 29,000,000 20 1,115,000 5. Total Cretaceous 75,500,000 31 2,1.35,000 0, Paleocene-Eocene 30,000,000 4 7,500,000 7. Oligocene-Recent 30,500,000 3 12,170,000 3. Total Cenozoic 00,500,000 7 9,500,000 9. Total of Cretaceous and Cenozoic 142,000,000 38 3,740,000 cypods in years throughout the Cretaceous and Cenozoic. The actual time spans of the various stratigraphic units were taken from Palmer, 1983. The extinction rate was low in the Early Cretaceous, but beginning with the Cenoma- nian, which is the earliest Stage of the Late Cretaceous, the extinction rate increased and continued into the Campanian Stage. This latter rate is a fourfold increase over the Early Cretaceous rate of extinction. The Maastrich- tian Stage at the end of the Cretaceous has the highest rate of extinction, being about five times faster than the previous figure in the rest of the Late Cretaceous, and it is about 20 times faster than the rate in the Early Cretaceous. Further- more, the extinction rate in all of the Late Cretaceous was more than eight times more rapid than it was in the Early Cretaceous. These extinction rates do not appear to be an artifact of the geologic record. Cretaceous strata, par- ticularly that of a shallow-water marine environ- ment, are exceedingly widespread and have Vol. 100(4) October 31, 1986 THE NAUTILUS 143 been thoroughly studied by invertebrate paleon- tologists. Evolution increased greatly in rate during the Late Cretaceous, a time span of 29,000,000 years. This rate of evolution in- creased and reached a clima.\ in the latest Cretaceous Maastrichtian Stage, which spans only 8,000,000 years. For example, three of the supergeneric groups that arose in the Late Cretaceous also became extinct during that part of the Cretaceous Period. Apparently new species, genera, and families arose with increas- ing rapidity during the Late Cretaceous as evinced by the rudists (Jones and Nicol, in press, and Nicol, in press). The Inoceramidae also evolved rapidly during the Late Cretaceous. One may ask, did the widespread warm shallow seas that covered the continents during the Late Cretaceous cause evolutionary rates to increase greatly? LITERATURE CITED Dockery, D. T., III. 1982. Lower Oligocene Bivalvia of the Vicksburg Group in Mississippi. A/).s-.s-(.s.s-(p/)) Bureau of Geology. Bull. 123, 261 p. Jones, D. S. and D. Nicol. Origination, survivorship, and ex- tinction of rudist taxa. Jour. Paleontology 60:107-115. Keen. A. M. 1980. The pelecypod family Cardiidae: A tax- ononiic summary. Tulnne Studief: in Geology and Paleon- tology 16:1-40. Moore, R. C. (ed.). 1969, 1971. Treatise on Invertebrate Paleontology. Part N, Mollusca 6, Bivalvia, Vols. 1-2 (1969). Vol. 3 (1971). Univ. Kansas Press, Lawrence, p. N1-N1224. Nicol, D. 1984. Critique on Stenzel's book on the Ostracea. The Nautilus 98:123-126. Some aspects of the evolution of the rudist pele- cypods. The Nautilus 100(2):69-71. Palmer, A. R. 1983. The decade of North American geology 1983 geologic time scale. Geology ll:.503-.504. Stanley, S. M. 1968. Post-Paleozoic adaptive radiation of in- faunal bivalve molluscs -A consequence of mantle fusion and siphon formation. Jour. Paleontology 42:214-229. FRESHWATER AND TERRESTRIAL SNAILS OF SAINT LUCIA, WEST INDIES^ Emile A. Malek Department of Tropical Medicine Tulane University Medical Center 1430 Tulane Avenue, New Orleans, LA 70112 ABSTRACT Surveys were made of the Jreshwater and terrestrial snails and slugs of Saint Lucia, one of the Windward Islands of the Lesser Antilles. Ten species of 7 families of freshwater snails, 9 species of 7 families of terrestrial snails, and 2 species of one family of slugs were encountered. The freshwater snail fauna inhabit all types of freshwater habitats available, i.e., rivers, br-ooks (ravines), stream embankments, flood plains and swamps adjacent to rivers, drainage ditches and ponds. Only one species of Biomphalaria, B. glabrata, was found, although other species of this genus occur on some other neighboring islands, alone or together with B. glabrata. The majority of the land srmil species were repre- sented in the capital city, Castries, and the nearby hills and coastal lands. Cer- tain species of land snails were associated with limestone deposits and others, although inhabiting the same areas, were also found in the igneous mountain highlands and in Jbrests. A scarcity of land operculates was observed; only one species was found. There is some old information on the mollus- 'Contribution from the World Health Organization Colla- borating Center for Applied Medical Malacology, Tulane University. can fauna of Saint Lucia. Bland (1861), E. A. Smith (1889, 1895) and E.A. Smith and Feilden (1891) listed land snails and a few freshwater snails of Saint Lucia and some neighboring islands. The author (1965), in an abstract form. 144 THE NAUTILUS October 31, 1986 Vol. 100 (4) enumerated and discussed briefly the fresh- water and land snails based on three trips to Saint Lucia for which reports to the Pan American Health Organization were prepared (Malek, 1962, 1963). The snails were later (Malek, 1980) illustrated to emphasize shell characteristics. Jordan (1985) in a brief appen- dix included only a short list of the freshwater and terrestrial snails on Saint Lucia, some of which were identified by the author. The present report was prepared in view of the significance of the molluscan fauna in research on snail-transmitted diseases on the island, especially schistosomiasis and fas- cioliasis, and the reports in recent years on freshwater snails in other Antillean islands such as Guadeloupe (Pointier, 1974), Haiti (Robart et al., 1976), Martinique (Guyard and Pointier, 1979), Guadeloupe, Dominica and Martinique (Starmuhlner, 1985) and the Dominican Republic (Gomez et al., 1986). The Island of Saint Lucia Saint Lucia, one of the Windward Islands located between Martinique and Saint Vincent, is a small island 27 by 14 miles and is about 233 square miles in area. The middle part is moun- tainous; there is a longitudinal main ridge, almost median, with other mountains running down to the coast on either side. Between the offshoots of the main mountain ridge are broad, flat valleys which spread out toward the sea. Small rivers, measuring only 7 to 12 feet in width in most parts, and only a few feet in depth, run from the mountain heights through the valleys. Most rivers form separate drainage systems that are subject to frequent flooding. Rainfall over most of the island averages 80 inches per year, but with only 40 inches on the southern and northern tips. The highest rainfall occurs from May through October, with peaks in August and September. Remnants of past volcanic activity are still noticeable in the sulphur area near Soufriere. The river beds have big rocks, seemingly of volcanic origin. Also in the geologic history certain areas were elevated, and during the Cretaceous period, deposition of limestone took place on several parts of the island, which support large numbers of the molluscan species. Results Ten species belonging to 7 families of freshwater snails, 9 species belonging to 7 families of terrestrial snails, and 2 species of one family of slugs were encountered in this survey (Table 1). Of the freshwater snails the Planorbi- dae was represented by 3 species, the Neritidae by 2 species and the Lymnaeidae, Physidae, An- TABLE 1 . Families and species of gastropods and their loca- tions in Saint Lucia. FRESHWATER Planorbidae Biomphala ria glnbrata Rivers: Castries, in the Entrepot area; Derniere, north of Dennery; Esperance, near Fond St. Jacques; Bance, near bridge south of school; Choc; Raveneau, at Delcer. Brooks (ravines): Ti Rocher; Trou Macis near Bance; above fall at Sulphur Springs. River and brook embankments: Ravine Poisson at Ravine Poisson; Augier River. Flood plains or swamps off rivers: Vieu.x Fort north of Joyeox; Canaries; Augier. Drainage ditches: Banana fields near Marquis and in Cul de Sac Valley. Ponds: At fall near Sulphur Springs; at Fond St. Jacques; at Derniere Fort. Concrete-lined channels: At Ferrand Estate. Drepanotrema depressissimum. Pond at entrance of Union Agricultural Station; pond in Bonne Terre area; pond in Ville Boutille; pond near Monchy school; drainage ditch in Marquis Estate; Ti Rocher brook; swamp by road between Vieux Fort and Labourie. D repanotrema I uctiiu m Pond near Monchy school; pond in Bonne Terre area; pond in Ville Boutille. Lymnaeidae Fossaria cubensis Ditches in banana fields, Cul de Sac Valley; along bank of river near LInion Agricultural Station; Ferrand Estate. Physidae Phusn ruheiisis Derniere River; embankment along Poisson brook; Mar- quis River; Bagatelle River in the Marchand area; pond in Bonne Terre area; pond at Ville Boutille; pond in Monchy area; brook joining Vieux Fort River north of Joyeax. Ancylidae Gundlachia radiata Pond near Sulphur Springs; Derniere River; Castries River in Marchand area; Choc River; ditches in banana fields, Marquis Estate. Hydrobiidae Pyrgopkonis parmtlus Brook, near Fond St. .Jacques; brook near Joyeux; brook in Cul de Sac Valley. Anipullaridae Atiipulldnii ghiiica Derniere River; Marquis River; Bonne Terre pond; pond at entrance of Union Agricultural Station. Neritidae Neritina punctulata and N. virginea Vol. 100(4) October 31, 1986 THE NAUTILUS 145 Castries River; Marquis River; Choc River TERRESTRIAL Helicinidae Helicimi fasriata Ferrand Estate, on banana plants and weeds along drain- age ditches. Suceineidae Omalonyx guadeloupensis Banks of pond at entrance of Union Agricultural Station; Cul de Sac Valley; along ditches in banana fields. Succinea approximans Along Poisson brook; along Choc River; near Vieu.x Fort; at Union Agricultural Station. Subulinidae Subulina octona Castries; Bois d'Orange; near Choc River; at Monchy; near Union Agricultural Station. Streptaxidae Guletla bicolor Castries and vicinity. Bulimulidae Protoglyptus aanctaeluciae Castries and vicinity. Bulimulus guadalupensis Castries and vicinity; several localities along the coast and foothills; on banana trees and shrubs in Ferrand Estate. Urocoptidae Brachypodella tatei Several localities along coast and foothills Camaenidae Pleurodonte orbiculata Castries; Choc River area; several localities between Castries and northern tip of island; Bois d'Orange; Cul de Sac Valley; Dennery. Veronicellidae Vaginulus occidental is Cul de Sac Valley on banana trees; Ferrand Estate on banana trees; hills overlooking Bois d'Orange; along Poisson Brook. Veroni.cella floridana Several localities in the northern half of island, sometimes with V. occidentalis cylidae, Ampullaridae and Hydrobiidae by one species each. Of the terrestrial snails the Suc- eineidae and Bulimulidae were represented by 2 species each, while the Helicinidae, Subulinidae, Streptaxidae, Urocoptidae and Camaenidae were represented by one species each. Two species of slugs found belonged to the family Veronicellidae. The planorbid Biomphalaria glabrata (Say, 1818), intermediate host of Schistosoma man- soni, was found in 20 localities. Specimens, which are up to about 10 mm in diameter, never attain the large size of those found in Puerto Rico, the Dominican Republic or South America. Accordingly, F. C. Baker (1945) showed figures of B. glabrata from Saint Lucia and commented that they were immature. Examination of shell features and anatomical details of biompha- larids from various drainage systems indicated that B. glabrata is the only species of the genus Biomphalaria that exists on Saint Lucia. It is known that species such as B. havanensis. B. helophila and B. straminea occur on some of the Lesser and Greater Antilles, alone or in addition to B. glabrata. The snail was not found in fast- flowing waters, but frequents and breeds on flooded river and ravine (brook) embankments, in swampy areas close to rivers, in side pools and backwaters of rivers and ravines protected by rocks and vegetation, in drainage ditches of banana fields, and in concrete-lined channels. Sizable colonies were always found upstream rather than near the mouth of a river. The planorbids Drepanotrema depressissi- mum. (Moricand, 1839) and D. lucidum (Pfeiffer, 1839) live in ponds rich in aquatic vegetation. The lymnaeid Fossaria cubensis (Pfeiffer, 1839) is the snail host of Fasciola hepatica which is common on the island. Wet mud outside the water, with or without vegetation, represents the typical habitat of this lymnaeid, although it was also found in narrow drainage ditches in banana fields, in shallow water, and in narrow concrete-lined channels. The Saint Lucian material of the hydrobiid Pyrgophorus parvulus (Guilding, 1828) exhibits variations common among individuals of this species. Some specimens have evenly rounded and smooth whorls, whereas other specimens have prominent spiral carinae situated a short distance between the sutures, and each carina carries a number of pronounced spines. The land snails and slugs found during the survey seemed to be widely distributed, especially in the island's northern two-thirds where the tropical vegetation and coastal deposits of limestone exist. Species such as Subulina octona (Bruguiere, 1789), Pleurodonte orbiculata (Ferussac, 1821), Bulimulus guadaloupensis Bruguiere, 1792 and Protoglyp- tus sanctaeluciae (E.A. Smith, 1889) occur in abundance in the most populated town, Castries. The several hills on which the town is located and the coastal strip harbor large col- onies of these snails. Banana plantations in several parts of the island support large popula- 146 THE NAUTILUS October 31, 1986 Vol. 100(4) tions of helicinids, bulimulids, and veronicellid slugs. Discussion The distribution of the freshwater snails on Saint Lucia seems to be related to characteris- tics of the habitat, such as the stream gradient, water velocity, rainfall, and salinity. For exam- ple, B. glabrata was not found in parts of streams with steep gradients or in streams close to the sea. The nearest colony was found at about 400 yards from the coast. The bionomics of S. glabrata on Saint Lucia were investigated by Sturrock (1973, 1974). Neritina spp., however, seem to withstand higher salinity than the other freshwater snails. The drepanotre- matids were found to be almost exclusively pond-inhabiting species, the exception being the occurrence of Drepanotrenia depressissimum once in a drainage ditch and once in a slow- flowing brook. The distribution of the land snails on this and other Antillean islands follows the usual correla- tion between the distribution of species and the geological formation of the area in which they occur. Species belonging to Brachypodella, Gulella, and Subulina are restricted to the limestone coastal land and adjacent hills. Pleurodonte orbiculata (Ferussac, 1821), Bulimulus guadaloupensis (Bruguiere, 1792), and Protoglyptus sanctaeluciae (E.A. Smith, 1889) also inhabit these areas, but they are addi- tionally found in the igneous mountains in the highlands. The main species of freshwater and terrestrial snails on Saint Lucia also occur on certain other Caribbean islands. The faunal relationship of Saint Lucia to other Caribbean islands and to the mainland in Central and South America deserves comment. Several theories have been postulated to explain the faunal and floral similarities among the islands of the West Indies on the one hand and among those of the islands and the fauna and flora of Central and South America on the other hand. Among the theories postulated is the past existence of land l)ridges (Ihering, 1931), or isthmian links (Schuchert, 1935) between the islands themselves and be- tween the islands and the mainland of Central and South America. Snails have played an im- portant part in helping to correlate the geo- logical history and the zoogeography of the islands. Certain species, however, such as the cosmopolitan Subulina spp. and others are of lit- tle value in providing evidence of previous land connections. The land operculate snails offer evidence of zonal grouping within the West Indies. Surveys on Saint Lucia revealed the presence of only one land operculate, Helicina fasciata Lamarck, 1818-1822. Thompson (1967) described another operculate from Saint Lucia, Lifhacaspis xanthoglauca (family Cyclophori- dae). In general, however, there is a scarcity of land operculate snails on the Lesser Antillean islands and South America as compared to a preponderance of land operculates on the Greater Antilles islands. Among the freshwater snails, the planorbids show relationships with the South American fauna. Drepanotrem.a depressissimum (Mori- cand, 1839) is typically South American. B. glabrata is found on some islands of both the Lesser and the Greater Antilles, and its range extends into South America. In the Antilles, B. glabrata occurs on Saint Lucia, Martinique, Guadeloupe, St. Kitts, Antigua, Dominica, Hispaniola, and Puerto Rico. Ampullaria glauca (Linnaeus, 1758) is also South American. The melaniid Thiara granifera (Lamarck, 1822) was introduced by Prentice (1983) in field trials in marshes and streams for experiments on biological control oiB. glabrata. Whether T. granifera still exists in these habitats or has spread to other habitats in Saint Lucia is not known. Jordan (1985) listed the sphaeriid l)ivalve Pisidium punctiferum (Guppy) as a member of the freshwater fauna of the island, but it was not encountered in my surveys. How- ever, his list did not include the terrestrial oper- culate Helicina fasciata Lamarck, nor the slug Veronicella floridana (Leidy) which were both common in the material that I collected. LITERATURE CITED Baker, F. C. 1945. TheMolluncan Family Planorhidar. I'ni- versity of Illinois Press, Urbana, Illinois. Gomez, J. D., Vargas, M., and Malek. E, A. 1986. Fresh- water mollusks of the Dominican Republic. The Niiittilus 100 (in press). (Uiyani, A. and Pointier, J. P. 1979. Faune malacolofjique diiicaquicole et vecteurs de la schistosome intestinale en Martinique (Antilles Francaises). Ann. Pnra.'^itol. (Paris): 54: 193-205. Vol. 100(4) October 31, 1986 THE NAUTILUS 147 Ihering, H. von. 1931. Land bridges across the Atlantic and Pacific Oceans during the Kainozoic era. Quart. J. Geol. Soc. London 87:376-391. Jordan, P. 1985. Schistosomiasis. The St. Lucia Project. Cambridge University Press, Cambridge. Malel<. E. A. 1962. Report on precontrol studies of bil- harziasis in St. Lucia. Unpublished Document, Pan American Health Organization, Washington, D.C.: 1-18. 1963. Report on a precontrol survey of bil- harziasis in a pilot study area of St. Lucia. Second visit. Unpublished Document, AMRO-1.55, Pan American Health Organization, Washington, D.C.: 1-9. . 1965. Freshwater and land snails of St. Lucia, the West Indies. Amer. Malacot. Union. Annual Report. p. 38. 1980. Snail-Tra7isntitted Parasitic Diseases. Vol. I. CRC Press, Boca Raton, Florida. Pointier, J. P. 1974. Faune malacologique dulcaquicole de I'ile de la Guadeloupe (Antilles Francaises). Bull. Mus. Hist. Natur.. Paris, 3rd Ser., No. 235, Zool., 159:905-933. Prentice, M. A. 1983. Displacement of Biomphalaria glabrata by the snail Thiara granifera in field habitats in St. Lucia, West Indies. Ann. Trap. Med. Parasitol. 77:51-59. Robart, G., Mandahl-Barth, G. and Ripert, C. 1976. Inven- taire, repartition geographique et ecologie des mollusques dulcaquicoles d'Haiti (Carabes). Hnliotts 8:159-171. Schuchert, C. 1935. Historical Geology of the Antillean- Caribbean Region. John Wiley & Sons, Inc., New York. Smith, E. A. 1889. On the Mollusca collected by Mr. G. A. Ramage in the Lesser Antiles. Report III. Ann. Mag. Natur. Hist. (6th Ser.) 3:400-405. 1895. Report on the land and freshwater shells collected by Mr. H. H. Smith at St. Vincent, Grenada and other neighboring islands. Proc. Malacol. Soc. London 1:300-322. Smith, E. A. and Feilden, H. W. 1891. A list of the land and freshwater shells of Barbados. Ann. Mag. Natur. Hist. (6th Ser.) 5:247-257. Starmuhlner, F. 1985. Erstfunde von drei Arten von Suss- wasser Gastropoden auf den Inseln Guadeloupe, Dominica und Martinique (Kleine Antillen). Heldia 1:55-58. Sturrock, R. F. 1973. Field studies on the transmission of Schistosoma mansoni and on the bionomics of its inter- mediate host, Biomphalaria glabrata, on St. Lucia, West Indies. Int. Jour. Parasitol. 3:175-194. . 1974. Ecological notes on habitats of the fresh- water snail Biomphalaria glabrata, intermediate host of Schistosoma mansoni, on St. Lucia, West Indies. Carib- bean Jour. Sci. 14:149-161. Thompson, F. G. 1967. A new cyclophorid land snail from the West Indies (Prosobranchia) and the discussion of a new subfamily. Proc. Biol. Soc. Washington 80:13-18. ON THE REDISCOVERY OF TERAMACHIA MIRABILIS (CLENCH AND AGUAYO, 1941), AND ITS RELATIONSHIP TO OTHER CALLIOTECTINE VOLUTES William K. Emerson and Walter E. Sage III Department of Invertebrates American Museum of Natural History New York, N. Y. 10024 ABSTRACT A second known specimen o/Teramachia mirabilis (Cleyich and Aguayo, 19Jfl) is here recorded from, deep water (UGS.U meters) on the Little Bahama Bank. Com- parisons of this dead-collected, crabbed speciynen are jnade with illustrations and descriptions of the holotype, which was live-taken in 521.3 meters off Matayizas, Cuba, in 1939. Largely on the similarity of shell characteristics, this New World species is retained in the genus Teramachia, as is Calliotectum fischeri Olsson, 1964, from the Pliocene of Ecuador. Examination of the typological specimens of Calliotectum vernicosum Dall, 1890, for which a lectotype is here designated, con- firms the distinction between Calliotectum Dall, 1890 and Teramachia Kuroda, 1931. Zoogeographically, Teramachia is confined mostly to the western Pacific and is known to date froyn the Neogene of Japan as ivell as Ecuador. Thus the widely 148 THE NAUTILUS October 31, 1986 Vol. 100(4) separated ynodem distributional pattern of the genus Teramachia (six species in the western Pacific, one in the western A tlantic) suggests that T. mirabilis is a sur- viving element of a presently disjunct Pacific fauna. Howellia mirabilis Clench and Aguayo, (1941, pp. 177-178, pi. 14, fig. 2; Weaver and duPont, 1970, p. 178, pi. 76 A, B) was described on the basis of a single, live-collected specimen dredged at Atlantis Station No. 3483, off Matanzas. Matanzas Province, Cuba (23°12'N, 81°23'W), in 521.3 meters. The soft parts unfor- tunately were not recovered in a condition that could be preserved, and the shell was cleaned and dried during the trip. Regrettably, the holotype (catalog #135291) cannot be located in the collection of the Museum of Comparative Zoology and is presumed lost (teste D. Backus). This taxon was proposed as the type species for the monotypic genus Howellia by Clench and Aguayo (1941, p. 177). Howellia was placed pro- visionally by Clench and Aguayo (op. cit.) in the family Fasciolariidae, in the absence of know- ledge of the radula and soft-part anatomy. Sub- sequently, Clench and Turner (1964, p. 177) assigned Howellia to the Volutidae, subfamily Calliotectinae, following the placement of Pilsbry and Olsson (1954, p. 19), based on shell characters. Clench and Turner (1964, p. 178), however, noted the resemblance in shell mor- phology of Howellia mirabilis to the western Pacific genus Teramachia Kuroda, 1931. Later Weaver and duPont (1970, p. 176) placed Howellia, 1941, in the synonymy of Teramachia, 1931, where it was retained by Emerson (1985, pp. 102, 103). Recently Harry G. Lee of Jacksonville, Florida kindly called our attention to a crabbed specimen of Teramachia mirabilis in the collec- tion of the Indian River Coastal Zone Museum (IRCZM) at Harbor Branch Oceanographic In- stitution, Inc., Fort Pierce, Florida. This specimen, apparently only the second known for this taxon, was found in a sediment sample col- lected on a Johnson-Sea-Link I submersible dive during R/V Johnson Cruise 159, on October 4, 1983. The bucket had been left on the bottom for 408 days to collect accumulating sediment near Black Rock, Little Bahama Bank (26°16.5'N, 77°38.5'W) in 465.4 meters, by C. M. Hoskin and J. K. Reed. The specimen presumably was carried into the 20-liter plastic bucket by the hermit crab occupying the shell. Although long-dead and now discolored from the muddy sediment, this specimen, IRCZM #065:02138 (figs. 5, 6), compares favorably with the description and illustration of the holotype. The present specimen is larger (125 vs. 93 mm in height) and has weaker axial sculpture on the body whorl. In outline and in the development of axial sculpture on the body whorl Teranfiachia mirabilis most closely resembles the western Pacific Teramachia johnsoni (Bartsch, 1942, p. 12, pi. 2, fig. 3; holotype illustrated in color by Weaver and duPont, 1970, pi. 750, H; cf. figs. 5, 6 with figs. 7, 8 herein). Teramachia johnsoni, however, has a tannish brown shell with a darker aperture, whereas the holotype of Tera- machia mirabilis is whitish and tinged with a faint brownish red. The aperture is light brown- ish red within. The discovery of the newly recognized speci- men of Teramachia mirabilis extends the known provenance of this species northwest- ward from the vicinity of Cay Sal Bank, off northern Cuba, to the Little Bahama Bank. Specimens can be expected to occur at suitable depths elsewhere on these and other regional banks. In the absence of anatomical data on the type species of Howellia it seems prudent to refer this New World species to the genus Tera- machia on the basis of shell characters held in common with the western Pacific species of this group of calliotectine volutes (Weaver and duPont, 1970, p. 177, Emerson, 1985, p. 103). A more precise systematic assessment oi Howellia must await knowledge of the soft parts and radular characters of Teraynachia mirabilis. Clench and Turner (1964, p. 178) briefly noted the close resemblance of the shell morphology of Howellia mirabilis to that of the Japanese species of Teramachia, and they stated, "Rela- tionships of this sort from widely separated areas are unusual, but they do exist". Shortly thereafter, Olsson (1964, p'. 129, pi. 23, fig. 4) described Calliotectum fischeri from late Pliocene rocks of the Esmeraldas formation of Ecuador. The holotype is based on a 52.8 mm Vol. 100(4) October 31, 1986 THE NAUTILUS 149 150 THE NAUTILUS October 31, 1986 Vol. 100(4) fragmental specimen, which was estimated to be "between six and eight inches in length" before the badly fractured specimen was extracted from the matrix. Olsson compared his specimen to Prodallia dalli Bartsch, 1942 from the Philip- pines and he referred Teramachia Kuroda, 1931 and Prodallia Bartsch, 1942 to the synonymy of Calliotectum Dall, 1890. We have examined the holotype and six fragmental specimens of Olsson's "Calliotectum" fischeri from near the type locality (Tulane Geol. Dept. localities 1397 [figs. 3, 4, herein] and 1402, and W. D. Pitt Coll. locality 9A-B3-915, all from the Esmeraldas for- mation). These specimens are from a deep-water facies and confirm the allocation of Olsson's taxon to the genus Teramachia. The only other known extinct species of Teramachia was described from the Mio-Pliocene [?Pliocene] Shimajiri formation of Okinawa, Japan as T. shinzatoensis (MacNeil, 1961, p. 96, pi. 9, fig. 1; Rehder, 1972, p. 9, figs. 5, 6). It was compared by the describer and Dr. Rehder with T. johnsoni (Bartsch, 1942). The placement by Olsson (1964, p. 128) of his Pliocene Ecuadorian fossil (Teramachia fischeri) in the monotypic genus Calliotectum Dall (1890, p. 304) requires comment. Dall (1890, p. 305, pi. 5, fig. 8) considered the type species of Calliotectum (C. vernicosmn Dall, 1890) to be pleurotomoid, but he noted the shell lacked an anal notch and a fasciole. Subsequent- ly, Pilsbry and Olsson (1954, p. 19, fig. 16, rachi- dian tooth) proposed the volutid subfamily Calliotectinae with Calliotectum the type genus, on the basis of shell and radular characters, the radular ribbon being uniserial with a tricuspid rachidian tooth. Pilsbry and Olsson (1954, op. cit.) also referred Teramachia Kuroda, 1931, with Prodallia Bartsch, 1942 as junior synonym, to Calliotectinae. Weaver and duPont (1970, pp. 175, 176) added Howellia Clench and Aguayo, 1941 to the synonymy of Teramachia, 1931. Dall (1890, p. 305) based Calliotectum ver- nicosum on five specimens dredged by the "Albatross", in 1888. Three of these were taken at station 2807, in 1485 meters from the type locality near the Galapagos Islands (00°24'00"S, 89°06'00"W). From this lot are Dall's figured specimen USNM #96555 (1890, pi. 5, fig. 8; Abbott and Dance, 1982, p. 224) here selected as lectotype (figs. 9, 10), and two paralectotypes: USNM #633904, and DMNH #10135 (Weaver and duPont, 1970, pi. 75, A, B) - both ex USNM #96555. The two remaining paralectotypes (USNM #97068) were dredged at station 2793, in 1355 meters off the coast of Ecuador (01°03'00"N, 18°15'00"W). These typological specimens are very thin- shelled, with inflated whorls, the largest specimen being the lectotype with a height of 47.5 mm (spire incomplete) and with 6V2 post- nuclear whorls, and the smallest paralectotype being 24.8 mm in height (spire incomplete) with 5 postnuclear whorls. Although superficially resembling the shells of Teramachia. specimens of Calliotectum. of the same stage of growth FIGS. 9, 10. Calliotectum reniira^um Dall. 1890, near the GalapaKos Islands, in 1485 meters, lectotype USNM #96555; X 1. FIGS. 1,2. T. dalli daydoni Poppe, 1986, off Port Hedland, northwestern Australia, in 450 meters, AMNH #221239. FIGS. 3, 4. Teramachia fifickeri (Olsson, 1964), Quebrada Camerones, Province of Esmeraldas, Ecuador, Tulane Geol. Dept. loc. 1397, Pliocene (early whorls and basal whorls missing). FIGS. 5, 6. T. mirabilis (Clench and Aguayo, 1941), Little Bahama Bank, in 465 meters, IRCZM #065:02138. FIGS. 7, 8. T. johnsoni (Bartsch, 1942), off Panglao, Bohol. Philippines, in 365 meters, AMNH #219986. FIGS. 1-8, approximately x 1. Vol. 100(4) October 31, 1986 THE NAUTILUS 151 have expanded, bulbous whorls and appear to represent a separate calliotectine lineage (cf. figs. 9, 10 with figs 1, 2 and figs. 3, 4). Therefore we reject Olsson's placement of Teramachia Kuroda, 1931 and Prodallia Bartsch, 1942 in the synonymy of Calliotectum Dall, 1890. The widely disjunct modern distributional pat- tern of Teramachia, six species in the western Pacific and one in the western Atlantic (Emer- son, 1985; Bouchet, 1986; Poppe, 1986), with ex- tinct species in the Mio-Pliocene of Okinawa and the Pliocene of Ecuador, presents some prob- lems in zoogeographic interpretation. The dis- tribution of these deep-water mollusks is poorly known and the genus may be of wider occur- rence than the available data suggest. Within the past few months, Teramachia dalli claydoni Poppe (1986) was described from the continen- tal slope of northwestern Australia and an ap- parently new species of Teramachia inhabiting deep water off New Caledonia was reported (Bouchet, 1986). Additional species may even- tually be found living in the eastern Pacific and elsewhere in the Atlantic Basin. The presence of Teramachia mirabilis living in the western Atlantic, however, may be an example of a Pacific faunal element which survived after being carried into the Caribbean region on the East-Pacific-Caribbean plate to its present posi- tion adjacent to Cuba (see Durham, 1985). Ac- cording to Sykes et al. (1982), the Caribbean plate has moved east-northeast about 1,400 km since late Eocene time. Perhaps the precursors of T. mirabilis were carried into the western Atlantic on this plate. Durham (1985) cites addi- tional examples of Pacific faunal elements that apparently moved with this plate into the Carib- bean area. Acknowledgments In addition to Dr. Harry Lee, we thank Paula Mikkelsen, Indian River Coastal Zone Museum, Harbor Branch Oceanographic Institution, Inc., Fort Pierce, Florida, William D. Pitt, Sacra- mento, California, Drs. Emily and Harold Yokes, Tulane University, and Drs. Richard S. Houbrick and Thomas D. Waller, National Museum of Natural History, Smithsonian In- stitution, for the loan of specimens in their care. We also thank Drs. Joel Cracraft, Leslie F. Marcus, Malcolm McKenna, and Richard H. Tedford for an exchange of views on the bio- geographical significance of Caribbean plate tectonics. We are indebted to our colleagues in the De- partment of Invertebrates, American Museum of Natural History for their contributions to this study: Stephen M. Butler for the photography and Stephanie Crooms for word-processing the manuscript. LITERATURE CITED Abbott, R. T. and Dance, S. P. 1982. Compendium, of Sea- shells. New York, 411 pp., illus. in color. Bartsch, P. 1942. Some deep-sea Philippine volutids. The Nautilus 56(1):9-13, pi. 2. Bouchet. P. 1986. Oceanographic campaigns in New Cale- donia. Rossiniana, Bull. L'Assoc. Conchyl. De Nouvelle- Caledonie 31;3-8, illus. Clench, W. J. and Aguayo, C. G. 1941. Notes and descrip- tions of new deep-water Mollusca obtained by the Harvard-Havana Expedition off the coast of Cuba. IV. Mem. Soc. Cubana Hist. Nat. 15(2):177-180, pi. 14, Clench, W. J. and Turner, R. D. 1964. The subfamilies Volu- tinae, Zidoninae, Odontocymbiolinae, and Calliotectinae in the western Atlantic. Johnsonia 4(43):129-180, pis. 80-114. Dall, W. H. 1890 [1889]. Scientific results of explorations by U. S. Fish Commission Steamer Albatross. VII. Prelimi- nary report on the collection of Mollusca and Brachiopoda obtained in 1887-'88. Proc. U. S. Nat. Mu.s. 12(737): 219-362. pis. 5-14. Durham, J. W. 1985. Movement of the Caribbean plate and its importance for biogeography in the Caribbean. Geology 13(2):123-125, 2 figs. Emerson, William K. 1985. Teramachia dupreyae new species, from off Western Australia (Gastropoda: Voluti- dae). The Nautilus 99(4):102-107, 8 figs. Kuroda, T. 1931. Two new species of Volutacea. Venus 3(1): 45-49, 3 figs. Olsson, A. A. 1964. Neogene mollusks from northwestern Ecuador. Paleont. Res. Inst., 256 pp. 36 pis, MacNeil, F. S. 1961. Tertiary and Quaternary gastropods of Okinawa. U. S. Geol. Surv. Prof. Paper 339 ("I960"): i-iv + 1-148, pis. 1-19 (distributed March 17, 1961). Pilsbry, H. A. and Olsson, A. A. 1954. Systems of the Volutidae. Bull. Amer. Paleont. 35(152):271-306, pis. 2.5-28. Poppe, G. 1986. A novel species and a novel subspecies of Volutidae from northwestern Australia. .4 pfx. Inform. Sci. Soc. Beige de Malac. l(l):27-36, pis. 1-3. Rehder. H. A. 1972. Some notes on the genus Teramachia Volutidae: Calliotectinae). The Veliger 15(1):7-10, figs. 1-7. Sykes, L. R., McCann, W. R., and Kafka, A. L. 1982. Motion of Caribbean plate during last 7 million years and implica- tions for earlier Cenozoic movements. Jour. Geophysical Res. 87(B13):106.56-10676. Weaver. C. S. and duPont, J. E. 1970. Living Volutes: a monograph of Recent Volutidae of the world. Delaware Mus. Nat. Hist. Mong. Ser., No. 1, xv + 375 pp., 79 col. pis., 44 figs., 13 maps. 152 THE NAUTILUS October 31, 1986 Vol. 100(4) REMINISCENCES [About a hundred years ago The Nautilus published many anecdotal accounts, but as the journal became more technical there was rarely an opportunity to add a personal touch. Because this 100th anniversary volume is dedicated to mala- cologists who have contributed in the past, we are including a few personal reminiscences by friends of William J. Clench and Joseph Rosewater. To the list of departed fellow-malacologists we regretably must add the name of that distinguished freshwater expert, Henry van der Schalie, whose touching obituary is included in this number of The Nautilus. -R.T. A., editor]. To Joe Rosewater— from R. Tucker Abbott [read at the 1985 memorial tribute at the Smithsonian Institution] The corridors of malacology may be darkened today with the passing of our colleague and friend, Joe Rosewater, but we and generations to follow will find many bright rooms off to the side that are filled with Joe's warm kindness and illuminating scientific labors. Calliostoma rosewateri- one of the most beau- tiful seashells of the Caribbean - was appropri- ately named after Joe by Drs. Clench and Turner twenty five years ago. Joe loved and understood the human qualities of his colleagues and a year later, in his famous biography of Calvin Goodrich, he quoted two letters written to his mentor, Bill Clench: "Thank you for naming the new species for me. In the days to come when the new genera- tion rolls up its sleeves to undo the work of this one, as this one is undoing that of [tax- onomic] workers who thought they had done a good, decent job, some one will note the name and speculate a bit about the guy it honors, and that will be in the nature of fame." And Joe, with a sense of humor, went on to quote another prophetic letter: "Your students [meaning Clench's students] are of a high order. I was delighted with them, and would steal them away if I knew just how to go about it. After all, you've more students than you should have, considering the rarity of boys interested in mollusks. I wish you would tell them that I consider it far more of a pleasure to have met them and I have delighted in their friendliness and intelli- gence, whatever the police may think of them." Just a month ago I received a letter from a now-retired, U. S. National Museum paleon- tologist living in Florida: "Joe really did a fine job of running the Divi- sion of Mollusks, and one could always write to him for help and expect a prompt reply. Joe was a careful and meticulous researcher as you well know." Joseph Rosewater was born in Claremont, New Hampshire, on September 18, 1928. He was of the same New England breed of mala- cologists as the Binneys, Dall, Morse, C. B. Adams, A. E. Verrill and W. J. Clench. Follow- ing his early college days at the University of New Hampshire, Joe began his advanced train- ing in malacology and museum curating at Harvard University. He was a favorite student of Clench's, and as Goodrich would say, was unknown to the police. His magnificent service to the Smithsonian In- stitution began in 1960. Joe's twenty-five years of malacology was a fulfillment of James Smithson's bequest for the "increase and diffu- sion of knowledge among all men." He served as President of the American Malacological Union in 1968, and as President of the National Capitol Shell Club in 1964. His contributions to the leading malacological journals of America are well-known to his colleagues. His travels in search of mollusks to the Marshall Islands, Indonesia, Australia and Panama were the envy of all devoted shell collectors. No more gentle soul, with such a quiet sense of humor, nor with a more natural ability to help both professional or amateur malacologists could we all have asked for. Let us hope that his successors will match his sterling qualities. Let's light up the halls of the National Museum once again. About Bill Clench -from Mary C. Kline [Mary Kline, now of Honolulu., and her late husband, George, were for many years ardent shell collectors who contributed many thousands of specimens, photographs and much biological Vol. 100(4) October 31, 1986 THE NAUTILUS 153 information on motlusks for- the journals. Johnsonia and Indo-Pacific Mollusca.] Bill Clench was one of my favorite "ex- tremists"-extremely kind, caring and loyal; an extremely good raconteur. By his enthusiasm for collecting shells and his insisting that the neophyte keep good field notes, he directed many beginners down the happy road to ex- tremely rewarding and meaningful adventures. Our friendship with Bill started about forty years ago when George and I first fell prey to the spell of the shell during a visit to Sanibel. On that first day of beach-walking George found a "different looking" shell. Local savants urged us to write to Dr. Clench at Harvard. The shell turned out to be an almost unheard of albino Fasciolaria distans. And so began an encouraging flood of corre- spondence in response to our new shelling trips to the Florida Keys and Bahamas. Bill was in those days launching his new journal, Johnsonia. We had many live-collected specimens, plenty of notes and colored pictures of animals. "Drop everything, come to the MCZ [Museum of Comparative Zoology] for a week- end and bring the slides." We went, we talked, we spent a long time on the steps of the museum bing innoculatead with Bill's enthusiasm. At his home, his wife Julia, was a marvelous hostess who had great patience and was a perfect house mother to all and sundry who interested Bill. Bill was many things but hardly the Ivory Tower type. For example: one day when we ar- rived at the museum lab he could hardly wait to show us his latest treasure. We knew that he and his associates had been working for a long time on a seemingly endless project -cleaning and rehabilitating the myriad drawers in the many huge specimen cabinets. As each drawer was finished a small blue star was affixed to the front, indicating that it had been completed. At last the goal had been reached and a party was given to celebrate. Bill was presented with a beautifully wrapped gift package. Inside was a suitably framed miniature of a real pair of Long- John bottoms with blue stars glued all over them! How this wonderful man loved those "Blue Starred Drawers!" He begged us to become interested in land shells as well as freshwater mollusks for, he said, "the human population will grow and grow and habitats will become neighborhoods -fresh- water shells will disappear for pollution is in- evitable but the sea will always be with us." He lived to see the land fills where once there were reefs. Many fashionable resorts now have replaced sleepy lagoons over much of the tropical world. What did I learn from Bill? -so many things - and here are just a few. 1 . A shell without data is just a pretty bauble. 2. Learn the scientific names, beach names are generally local. 3. Be sure to learn the dangers of the areas in which you are shelling -and avoid them. 4. Join the A.M.U. and your local Shell Club. 5. In a foreign country deal with the proper authorities and remember, in your own way you represent America. 6. Ask the museum curator how you can help -leave the scientific side to him. 7. Keep accurate notes and charts -be a good observer. 8. Don't take all shells from one col- ony-leave the habitat as you found it. 9. And lastly, but far from least, keep your sense of humor. Remembering Bill Clench — from Bunny Baker [Mrs. Horace B. Baker, for many years the business manager of The Nautilus, was raised in Boston, and has been an ardent shelter for over fifty years.] I first met Bill Clench in the fall of 1933 when I went to work at the Boston Children's Museum. The museum's exhibits were mostly on natural history topics, and though I was the financial secretary and only office worker, I thought I should know some natural history. My lifelong friend and childhood chum, who also worked in the museum, and I had collected shells along the Mass. coast, and so that seemed the logical place to start. I joined the Boston Malacological Club, and there met Bill who was to become a large part of my life. In the fall of 1934 my chum, her mother, sister and I went to Miami for our vacation. Her family returned home by boat after a few days, and we two girls drove across the Tamiami Trail to Sanibel Island to collect shells. Bill had told us so much about it that we wanted to collect there. After a week we drove back to the east coast and met Bill and two of his students who had been on a collecting trip to Cuba. We all drove 154 THE NAUTILUS October 31, 1986 Vol. 100(4) down to Homestead and into the Everglades where we camped out for five days and collected Liguus. Bill helped us identify the various color forms and taught us a lot about collecting and cataloguing our catch. It was a wonderful ex- perience and of course we all got really ac- quainted with each other. We got hundreds of shells, and those added to my Sanibel collection and others from around the world gave me quite a cabinetfull. They are now in Harvard's Museum of Comparative Zoology. Bill, his family and I saw much of each other; I became Secretary-Treasurer of the Boston Club and joined the A.M.U. It was at that meeting in Toronto in 1939 that I met my husband. Bill in- troduced us. In 1941 Bill drove his family and me to the meeting in Rockland, Maine. After a few days there I became engaged to H. B. Baker. We were married in December, 1941, and I moved to Philadelphia after a honeymoon on Sanibel. In subsequent years we saw Bill often and kept in close touch. He had many friends and ad- mirers. I am happy to have been one of them for over 50 years. We will never forget you, Bill. To Bill -Cecelia W. Abbott [Cecelia W. Abbott succeeded Mrs. Baker as Business Manager of The Nautilus, and she often accompanied Dr. Clench in his later Itfe on his collecting trips to Kentucky, Georgia and Florida and more recent A.M.U. meetings.] So much of Bill Clench's life was devoted to helping new students and encouraging amateurs to study and collect shells that I thought it only appropriate that a special trophy be created for shell shows that would emphasize his keen interest and love of land and fresh- water mollusks. The opportunity afforded itself when the FIG. 1. Three proud winners of the William J. Clench Award in Jacksonville, Florida, all of whom created outstanding ex- hibits on non-marine mollusks. (from the left: Archie Jones, Zida Kibler and Alfredo Romeu). Photo by William Kibler. Jacksonville Shell Club celebrated its 25th anni- versary in 1984. I lent a splendid sketch of Bill and with the efforts and talents of club member, Allan B. Walker, a handsome plaque was designed. Bill's likeness, surmounted by two of his favorite shells -Canthyri a spinosa (Lea) and Triop.ns albolabris (Say) graced the William J. Clench Award "to the outstanding exhibit of land or freshwater mollusks that shows overall excellence and best furthers interest in non- marine conchology." The award has now been given three times, the first one, in 1984 to Archie Jones of Miami, Florida, an old friend of Bill's and an expert on Liguus. Other winners, to date, have been Zida and William Kibler, in 1985, of Shallotte, North Carolina, for their worldwide land shell exhibit, and Dr. and Mrs. Alfredo Romeu of Jackson- ville, Florida, for their Land Shell of the World exhibit, 1986. Our hope has been realized that this trophy is inspiring others to produce outstanding, educational exhibits, and is reminding us all of the wonderful qualities of this kind and generous man after whom the Clench Award is named. 0 INFORMATION FOR SUBSCRIBERS The annual subscription rate for The Nautilus is $15.00 for individuals (foreign $18.00) and $20.00 for institutions (domestic or foreign). Subscriptions may begin in January. Send check or money order made out to "American Mala- cologists" to the Business Manager, P.O. Box 2255, Melbourne, Florida 32902-2255, 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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