trf^oS a ^ /) (/'> I INDO-PACIFIC MOLLUSCA Monographs of the Marine Mo/lusks of the Tropica / Western Pacific and Indian Oceans R. Tucker Abbott Volume I Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia Philadelphia, Pennsylvania, U.S.A. INDO-PACIFIC MOLLUSCA Monographs of the Marine Mollusks of the Tropical Western Pacific and Indian Oceans PUBLISHED BY The Department of Mollusks Academy of Natural Sciences of Philadelphia Philadelphia, Pennsylvania, U.S.A. EDITOR R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia CO-EDITORS William J. Clench Museum of Comparative Zoology Harvard College Cambridge 38, Massachusetts Harald A. Rehder United States National Museum Smithsonian Institution Washington 25, D.C. BUSINESS BOARD Alfred J. Ostheimer, 3rd George F. Kline R. Tucker Abbott SPONSORED BY The Natural Science Foundation 1510 Chestnut Street Philadelphia 2, Pennsylvania Publication commenced March 31, 1959 all receipts from the sale of INDO-PACIFIC A MOLLUSCA are invested in the Journal itself or expended directly to promote malacologieal research. Inquiries concerning subscriptions should be ad- dressed to INDO-PACIFIC MOLLUSCA, Depart- ment of Mollusks, Academy of Natural Sciences, Philadelphia 3, Pennsylvania, U.S.A. The Journal is not sent to libraries as an Academy publication ex- change. Contributions of specimens of Indo-Pacific marine mollusks for the advancement of research may be mailed to the Curator, Department of Mollusks at the Academy or to specialists currently working on particular genera or families. Instructions for Binding There are no special instructions for binding, since the numbers are issued in consecutive pamphlet form of four or more pages. Each volume contains approximately 300 pages with an index at the end. Consecutive pagination, for bound copies, appears in the normal place at the top right or left of each page. v Instructions for Looseleaf All pamphlets are punched for a standard-sized _ loo^leaf ring or post binder. If a ring binder is employed, the staples in each pamphlet should be removed and the pages cut; but, if a post binder is used, dismantling is unnecessary unless replacement pages are issued at a later date. Expandable post binders are recommended, and copies may be ob- tained at cost from the editorial office of the Journal. Pagination for looseleaf handling appears at the bottom center of each page. By maintaining the sheets in numerical order, even though there may be large pagination gaps, the Journal will be in biological sequence. Guide tabs to families, genera and special sections, such as “Index” and “News Bulletins,” are available at cost. Other journals of the same size, such as JOHNSONIA (Harvard College, Cambridge, Mass.), may be punched and added to the binder, if one wishes. The Natural Science Foundation The Natural Science Foundation is a tax-exempt scientific and educational organization incorpo- rated in 1953 in Pennsylvania for the purpose of en- couraging and promoting research and education in the natural sciences, particularly in malacology. It sponsors and helps finance INDO-PACIFIC MOL- LUSCA, and has supported numerous expeditions for mollusks in the Indian and Western Pacific Oceans. Copies of annual reports are available upon request. Inquiries concerning membership should be sent to the Natural Science Foundation, 1510 Chestnut Street, Philadelphia 2, Pennsylvania. o^tTHSONIAN Do % R Vi [00 - 003] institution March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Introduction 5 INTRODUCTION INDO-PACIFIC MOLLUSCA is a journal de- voted to the taxonomic revision of the marine mol- lusks of the tropical Western Pacific and Indian Oceans. For two hundred years many thousands of Indo-Pacific mollusks have been described in nu- merous and widely scattered publications with little or no correlation with previous reports nor in rela- tion to species described from other nearby islands and provinces of this rather uniform region. This has resulted in many synonyms, the unwarranted treatment of some forms as full species, wide vari- ation in nomenclature, and, in some cases, consid- erable duplication of research and effort. Purpose of the Journal Although taxonomic malacology, particularly of the Indo-Pacific, is to a great extent still in the de- scriptive and cataloging stages, with many new species yet to be described, it has seemed advisable to begin this series of taxonomic monographs, so that all the available information can be drawn together and a careful evaluation made of the known species. While the monographs in this journal may be considered to be “up to date”, experience tells us that no scientific effort is either perfect or complete. Newly collected material, a re-examina- tion of characters, differences of opinion, or neces- sary revisions in nomenclature all contribute to- wards changes, a better understanding and an im- proved classification of species. Method of Publication To help reduce the confusion caused by addi- tions and changes which inevitably arise when a new monograph stimulates further research, this journal is published in a form which permits it to be either bound in the normal manner or kept in looseleaf fashion. While most of our efforts will be directed towards genera and families that are still in need of revision, there will be some effort to keep previously published monographs “up to date” by adding newly discovered species, important changes in nomenclature or new information con- cerning habitats, geographical records and anatomy. These supplements will be issued at the close of each volume in the form of a Review Number, and those subscribers maintaining their journal in looseleaf form need only slip the additional pages in the appropriate places. Monographs will be un- dertaken in groups where sufficient material and competent specialists are available. Numbers will be published at irregular intervals, as research is completed, but it is hoped that a yearly minimum of a hundred pages will be maintained. Regional Scope of the Journal This journal attempts to cover the species which are mainly confined to the extensive tropical marine region known as the Indo-Pacific. This is a fairly uniform province embracing the Indian and tropical West Pacific Oceans. It extends from the east coast of Africa across the Indian Ocean (to the north half of Australia and the south half of Japan) and eastward through the Hawaiian Chain and Poly- nesia to Easter and Clipperton Islands. In mala- cological terms, the Indo-Pacific may be generalized as that region coinciding with the distribution of such common species as Cypraea caput-serpentis Linne, Conus ehraeus Linne, Nerita albicilla Linne and Antigona reticulata (Linne). Within this region there are several provinces which contain endemic species or exclude certain otherwise widely dis- tributed species. In cases where a genus or family is predomi- nantly Indo-Pacific, the few additional species which occur in other regions of the world will also be included. Pelagic groups, such as Janthina, Recluzia, the Pteropods, and certain cephalopods will be treated from a worldwide standpoint be- cause of their universal distribution. Scientific Names The scientific names employed by the authors of these monographs are those which are considered to be the most acceptable under the present rules and recommendations set down by the International Commission on Zoological Nomenclature. When personal opinion is involved, the name most widely used has been accepted. Synonymies The first reference under the species heading is to the original citation, and all subsequent refer- ences are to known or supposed synonyms. Other references may be included to demonstrate previ- ous general placement or lead to additional infor- mation, such as anatomy, habits, new localities, etc. The geographical name appearing in parentheses refers to the original locality as cited by the author. Measurements Measurements of specimens are made in milli- meters (25.37 mm. = 1 inch). Except where exten- sive measurements may be necessary, we generally list only an example of a large ( the largest we have [00-051] 6 Introduction Editors seen), average (a specimen that appears typical in size for the species and not the average of a long series of measurements ) , and small specimen ( given only when a small specimen can be determined as adult ) . Common Name The usage of common names is a very hazardous one because of the wide variation in local names. However, in most cases we have given the name which appears to be most generally in use. In other cases we have coined suitable names for those species which have never received a vernacular name. Range The range is a general statement of the geo- graphical distribution of a species, and is based mainly upon specimens we have examined, al- though some purportedly reliable records from the literature are used. Because of the great need for collections from many areas in the Indo-Pacific, these ranges are not necessarily final. It should also be realized that the special habitat preferences of the various species, such as for mangroves, reefs, rocky shores or sandy bays, may result in the species being absent in certain large areas not offering a suitable environment. Locality Records All records are based upon specimens examined by the author of a monograph. Only in rare in- stances have we used records from the literature, and in these cases have indicated the identifier. Because of the vast area covered and the numerous localities involved, we have omitted those which are very close to each other, and in other cases abbreviated the information to read, for example, “present on 13 of the 26 islets of Bikini Atoll.” After each record the name of the museum or private collector is included in parentheses. The following abbreviations refer to the major natural history museums containing large collections of Indo- Pacific mollusks: ANSP— Academy of Natural Sciences of Philadelphia Auck. Mus.— Auckland Institute and Museum, N.Z. Aust. Mus.— Australian Museum, Sydney BPBM— Bernice P. Bishop Museum, Honolulu BM— British Museum (Natural History), London Domin. Mus.— Dominion Museum, Wellington, N.Z. MCZ— Museum Comparative Zoology, Cambridge, Mass. Nat. Mus. Viet.— National Museum of Victoria, Melbourne NSF— Natural Science Foundation Expedition, deposited in the ANSP. So. Aust. Mus.— South Australian Museum, Adelaide USNM— United States National Museum, Wash., D. C. Order of Listing Locality Records Our locality records are listed in a general west to east order (Africa to Galapagos), and second- arily from north to south. The following list of localities is a general outline, and does not mention every island. This is also the sequence in which many museums keep their collections of a single species when there are numerous lots in a drawer. Madagascar (and Comores, Aldabra) Seychelles, Mauritius, Reunion, Rodriquez, Maldives, Laccadives, Chagos Ecology Under this section all ecological data have been included, although it is surprising how little infor- mation is available even for the common species. There is also some limitation to the value of some of the published habitat records, since we have no way of knowing if the identifications of the mol- lusks in question are correct or not. Collectors in the Indo-Pacific area can add much valuable infor- mation by recording simple observations on the relative abundance, habitat, habits and living ani- mals of the species collected. A sample of the AFRICA Natal to Somalia (including the islands of Socotra, Pemba, Zanzibar, Mafia) Red Sea and Gulf of Aden Persian and Oman Gulfs INDIAN OCEAN ISLANDS SOUTH ASIA West Pakistan, India, to the Ganges Ceylon Bay of Bengal Islands (Andamans and Nicobars) Burma and Malay Peninsula SOUTHEAST ASIA Korea to southern China; Hainan Japan, Ryukyus, Taiwan Philippines East Indies (Indonesia), Cocos Keeling AUSTRALIA MELANESIA New Guinea, Admiralty, Solomons, to New Caledonia, Lord Howe and Kermadecs MICRONESIA Bonins, Marianas, Palaus, Helen Reef, Carolines, Wake, Marshall and Gilberts POLYNESIA Hawaiian Chain Line Islands, Johnston, Ellice, Samoa, Tonga, the Cooks, Society, Marquesas and the Tuamotus EASTERN PACIFIC Easter, Clipperton and Galapagos [00 - 052] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Introduction 7 species studied should be sent to one of the several museums specializing in mollusks, so that taxono- mists will be able to correlate the ecological infor- mation with the correct species. The following out- line suggests the type of data most needed. Species number or name Geographical locality Date and collector Relative abundance Habitat Above tide line Intertidal Offshore depth Substrate Rock, sand, mnd, coral, algae or other Relation to substrate Atop, buried, attached to, or rock burrower Other T ype of water Motion Temperature Turbidity Other Living habits Solitary or gregarious Nocturnal or diurnal Special behavior Method of feeding Food Method of locomotion Associations with other organisms Breeding and eggs Enemies The living mollusk Shape Color For those interested in recording detailed infor- mation on mollusks collected on coral reefs and atolls, we recommend the reef terminology em- ployed by Joan Demond (Micronesian Reef- associated Gastropods, Pacific Science, vol. II, no. 3, pp. 275-341, July 1957). A similar terminology appeared in mimeographed form in the Atoll Re- search Bulletin, Washington, D. C., no. 46, 3 pp., 2 figs. 1955 (Tracey, Emery and Cloud). Cross section of a coral atoll showing reef terminology. (Modified from J. Demond, 1957). [00 - 053] 8 Introduction Editors *b °0 a % °0 ^ CJ D (M -+-m Generalized map of the approximate limits of the Indo-Pacific region upon which the mono- graphs in this journal are based. In cases where a genus or family is predominantly Indo-Pacific, the few additional species which occur in other regions of the world are also included in the monographs. [00 - 054] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Guide to Contents 9 GUIDE TO CONTENTS The classification of the Phylum Mollusca out- lined below by families is largely taken from J. Thiele’s Handbuch der Systematischen Weichtier- kunde, Jena, vols. 1 and 2, 1929-1935, and from W. Wenz’s Handbuch der Palaozoologie, Berlin, vol. 6, 1639 pp., 1938-1944. This list contains all of the recent marine groups. Names in square brackets refer to land and fresh-water groups which are not covered in this journal. The page numbers refer to the looseleaf pagina- tion found at the bottom of each sheet. Title page 00-001 Editorial Masthead 00-003 Instructions for binding 00-003 List of Issues and Changes 00-005 Introduction 00-050 Guide to Contents by Families 00-100 Phylum MOLLUSCA Class GASTROPODA Subclass PROSOBRANCHIA Order Archaeogastropoda Superfamily Pleurotomariacea Pleurotomariidae 00-150 Haliotidae 00-200 Scissurellidae 00-400 Fissurellidae 00-500 Superfamily Patellacea Patellidae 01-500 Acmaeidae 01-800 Lepetidae 01-950 Superfamily Trochacea Trochidae 02-000 Stomatiidae 03-000 Angariidae 03-100 Skeneidae 03-150 Cyclostrematidae 03-250 Turbinidae 03-500 Phasianellidae 04-000 Superfamily Neritacea Neritopsidae 04-180 Neritidae 04-200 Phenacolepadidae 05-000 Titiscaniidae 05-075 [Hydrocenidae] [Helicinidae] Superfamily Cocculinacea Cocculinidae 05-100 Lepetellidae 05-170 Order Mesogastropoda [Superfamily Cyclophoracea] [Cyclophoridae] [ Viviparidae] [Pilidae] [Lavigeridae] [Superfamily Valvatacea] Superfamily Littorinacea Lacunidae 05-200 Littorinidae 05-250 [Pomatiasidae] [Chondropomidae] Superfamily Rissoacea [Hydrobiidae] [Assimineidae] [Tornidae] Rissoidae 05-400 Adeorbidae ( Vitrinellidae) 06-000 Skeneopsidae 06-100 Rissoellidae 06-150 Superfamily Cerithiacea Turritellidae 06-200 Mathildidae 06-400 Omalaxidae 06-500 Architectonicidae 06-550 Vermetidae 06-700 [Thiaridae] Caecidae 06-850 Planaxidae 07-000 Modulidae 07-050 Potamididae 07-100 Diastomidae 07-200 Cerithiidae 07-250 Cerithiopsidae 07-550 Triphoridae 07-850 [00- 101] 10 Guide to Contents Editors Superfamily Epitoniacea Superfamily Buccinacea Epitoniidae 08-200 Columbellidae (Pyrenidae) 15-500 Janthinidae 08-600 Buccinidae 16-500 Melongenidae 17-000 ( Superfamily Pyramidellacea Nassariidae 17-200 see under Opisthobranchia 30-500) Fasciolariidae 18-000 Superfainily Hipponicacea Superfamily Volutacea Fossaridae 08-700 Olividae 18-800 Vanikoridae 08-800 Mitridae 19-400 Hipponicidae 08-900 Vasidae 20-400 Harpidae 20-600 Superfamily Calyptraeacea Volutidae 20-700 Trichotropidae 09-000 Cancellariidae 21-700 Capulidae 09-100 Marginellidae 22-000 Calyptraeidae 09-200 Superfamily Conacea Superfamily Strombacea Turridae 22-500 Xenophoridae 09-400 Conidae 25-000 Struthiolaridae 09-500 Terebridae 27-000 Aporrhaidae 09-600 Strombidae 09-650 Subclass OPISTHOBRANCHIA Order Onchidiata Superfamily Cypraeacea 07 Qfin Lamellariidae 10-100 Onchidiidae Z l -hflJU Eratoidae 10-200 Order Tectibranchia Cypraeidae 10-400 Superfamily Bullacea Ovulidae ( Amphiperatidae ) 11-400 Pupidae (Acteonidae) 28-000 Ringiculidae 28-500 Superfamily Atlantacea Hydatinidae 28-600 Atlantidae 11-500 Diaphanidae 28-700 Carinariidae 11-600 Notodiaphanidae 28-750 Pterotracheidae 11-700 Bullidae 28-800 Atyidae 29-000 Superfamily Naticacea Aceridae 29-200 Naticidae 11-800 Retusidae 29-300 Scaphandridae 29-400 Superfamily Tonnacea Philinidae 29-600 Cassididae 12-400 Gastropteridae 29-700 Cymatiidae 12-800 Runcinidae 29-750 Bursidae 13-200 Aglajidae 29-800 Tonnidae ( Oocorythinae ) 13-400 Ficidae 13-700 Superfamily Aplysiacea Aplysiidae 29-900 Order Neogastropoda Superfamily Muricacea Superfamily Pijramidellacea Muricidae 13-800 Aclididae 30-500 Magilidae 15-300 Melanellidae 30-600 [00 - 102] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Guide to Contents 11 Stiliferidae 30-800 Entoconchidae 30-900 Pyramidellidae 31-000 Order Pteropoda Suborder Thecosomata Superfamily Cavoliniacea Spiratellidae 31-500 Cavoliniidae 31-600 Superfamily Peracleacea Peraclidae 31-700 Procymbuliidae 31-800 Cymbuliidae 31-850 Desmopteridae 31-950 Suborder Gymnosomata Pneumodermatidae 32-000 Cliopsidae 32-050 Notobranchaeidae 35-100 Clionidae 35-200 Thliptodontidae 35-300 Anopsiidae 35-350 Laginiopsidae 35-400 Order Sacoglossa Oxynoidae 34-450 Caliphyllidae 35-550 Stiligeridae 35-600 Elysiidae 35-700 Limapontiidae 35-800 Oleidae 35-850 Order Acoela Suborder Notaspidea Umbraculidae 35-900 Pleurobranchidae 36-000 Suborder Nudibranchia Superfamily Doridacea Doridoxidae 36-200 Duvauceliidae 36-250 Bathydorididae 36-350 Hexabranchidae 36-400 Polyceridae 36-450 Corambidae 36-900 Vayssiereidae 36-950 Dorididae 37-000 Phyllidiidae 37-400 Superfamily Aeolidiacea 37-500 Doridoididae 37-700 Hedylidae 37-750 Dironidae 37-850 Dendronotidae 37-900 Bornellidae 37-950 Scyllaeidae 38-000 Phyllirrhoidae 38-050 Nectophyllirrhoidae 38-150 Tethyidae 38-200 Zephyrinidae 38-250 Lomanotidae 38-300 Iduliidae 38-350 Notaeolidiidae 38-400 Goniaeolididae 38-450 Madrellidae 38-500 Flabellinidae 38-550 Heroidae 38-700 Pseudovermidae 38-750 Tergipedidae 38-800 Fionidae 38-900 Calmidae 38-950 Aeolidiidae 39-000 Myrrhinidae 39-250 Superfamily Rhodopacea Rhodopidae 39-300 Subclass PULMONATA Order Basommatophora Superfamily Ellobiacea Ellobiidae 39-310 [Otinidac] Superfamily Amphibolacea Amphibolidae 39-500 Superfamily Siphonariacea Gadiniidae 39-600 Siphonariidae 39-700 Superfamily Hygrophila [Chilinidae] Latiidae 39-900 [Physidae] [Lymnaeidae] [Ancylidae] [00 - 103] Arminidae [Order Stylommatophora] 12 Guide to Contents Editors Class AMPHINEURA (Loricata) Superfamily Pectinacea Order Eoplacophora (Lepidopleurida) Dimyidae 55-000 Plicatulidae 55-100 Lepidopleuridae 40-000 Propeamussiidae 55-200 Order Mesoplacophora 40-200 Pectinidae 55-400 Ischnochitonidae Callistochitonidae 41-200 Spondylidae 56-000 Order Isoplacophora Limidae 56-200 Cryptoconchidae 41-700 Superfamily Anomiacea Cryptoplacidae 42-700 Mopaliidae 43-700 Anomiidae 56-500 Order Teleoplacaphora Placunidae 56-700 Chitonidae 44-200 Aulacochitonidae 45-200 Superfamily Ostreacea Callochitonidae 45-700 Ostreidae 56-800 Lepidochitonidae 46-200 Superfamily Gaimardiacea Class SCAPHOPODA Gaimardiidae Juliidae 57-300 57-400 Siphonodentaliidae 47-000 Dentaliidae 48-000 Order Eulamellibranchia Suborder Schizodonta Class PELECYPODA Superfamily Trigoniacea Order Palaeoconcha Trigoniidae [Mutelidae] 57-500 Superfamily Solemyacea [Aetheriidae] Solemyidae 50-000 Order Protobranchia [Superfamily Unionacea] Superfamily Nuculacea Suborder Heterodonta Nuculidae 50-100 Superfamily Astartacea Malletiidae 50-500 Astartidae 57-700 Nuculanidae Order Filibranchia Suborder Taxodonta Superfamily Arcacea 50-600 Crassatellidae Superfamily Carditacea Carditidae 58-000 58-200 Limopsidae 50-900 Condylocardiidae 58-400 Philobryidae 51-200 Superfamily Sphaeriacea Corbiculidae Glycymeridae Arcidae 51-400 51-700 58-500 [Sphaeriidae] Suborder Anisomyaria Superfamily Pteriacea Superfamily Glossacea Isognomonidae 52-700 Kellyellidae 58-900 Pteriidae 53-000 Glossidae 59-100 Pinnidae 53-500 Superfamily Arcticacea Superfamily Mytilacea Arcticidae 59-200 Mytilidae 53-700 Trapeziidae 59-250 [00- 104] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Guide to Contents 13 Superfamily Cyamiacea Cyamiidae 59-450 Sportellidae 59-500 Neoleptonidae 59-600 Superfamily Dreissenacea Dreissenidae 59-650 Superfamily Lucinacea Diplodontidae 59-700 Thyasiridae 59-900 Lucinidae 60-100 Superfamily Erycinacea Erycinidae (Leptonidae) 60-600 Montacutidae 60-800 Galeommatidae 61-000 Superfamily Chamacea Chamidae 61-200 Superfamily Cardiacea Cardiidae 61-400 Tridacnidae 62-000 Superfamily Veneracea Veneridae 62-100 Petricolidae 63-600 Superfamily Mactracea Mesodesmatidae 63-800 Mactridae 64-100 Anatinellidae 64-700 Cardiliidae 64-800 Superfamily Tellinacea Donacidae 64-900 Sanguinolariidae (Asaphidae) 65-200 Semelidae 65-500 Tellinidae 66-000 Suborder Adapedonta Superfamily Solenacea Superfamily My ace a Corbulidae (Aloididae) 67-500 Myacidae 68-000 Superfamily Gastrochaenacea Gastrochaenidae 68-200 Superfamily Pholaclacea Pholadidae 68-300 Teredinidae 68-800 Suborder Anomalodesmacea Superfamily Pancloracea Lyonsiidae 69-300 Pandoridae 69-500 Thraciidae 69-700 Periplomatidae ( Latemulidae ) 69-900 Myochamidae 70-100 Chamostreidae 70-300 Pholadomyidae 70-400 Superfamily Clavagellacea Clavagellidae 70-500 Order Septibranchia Superfamily Poromyacea Verticordiidae 70-600 Poromyidae 70-900 Cuspidariidae 71-200 Class CEPHALOPODA Subclass Tetrabranchia Nautilidae 74-000 Subclass DIBRANCHIA Order Decapoda Superfamily Sepiacea Spirulidae 74-300 Sepiidae 74-400 Sepiadariidae 74-700 Sepiolidae 74-900 Idiosepiidae 75-300 Glaucomidae Solenidae Superfatnily Hiatellacea 67-000 67-100 Superfamily Loliginacea Loliginidae 75-400 Lepidoteuthidae 76-300 67-400 Promachoteuthidae 76-400 Hiatellidae 14 Guide to Contents Editors Superfamily Architeuthacea Cirroteuthidae 77-950 Lycoteuthidae 76-500 Opisthoteuthidae 78-000 Enoploteuthidae 76-550 Suborder Incirrata Octopodoteuthidae 76-650 Neoteuthidae 76-700 Superfamily Bolitaenacea Onychoteuthidae 76-750 Bolitaenidae 78-050 Gonatidae 76-850 Amphitretidae 78-075 Psychroteuthidae 76-900 Vitreledonellidae 78-100 Architeuthidae 76-950 Histioteuthidae 77-000 Superfamily Octopodacea Alluroteuthidae 77-100 Octopodidae 78-150 Bathyteuthidae 77-150 Superfamily Argonautacea Brachioteuthidae 77-200 Valbyteuthidae 77-250 Alloposidae 78-250 Ommatostrephidae 77-300 Tremoctopodidae 78-300 170 0 CA Thysanoteuthidae 77-400 Ocythoidae 78-350 Chiroteuthidae 77-450 Argonautidae 78-500 Joubiniteuthidae 77-550 Cranchiidae 77-600 Glossaries, terminology 80-000 Order Octopoda Bibliographies 81-000 Suborder Cirrata Collecting and museum methods 84-000 Superfamily Vampyroteuthacea Expeditions, history, biographies 85-000 Vampyroteuthidae 77-800 Faunal and insular accounts, check lists 87-000 Laetmoteuthidae 77-850 Natural Science Foundation reports, Superfamily Cirroteuthacea list of consultants 89-000 Stauroteuthidae 77-900 Indices 90-000 [00 - 106] INDO-PACIFIC MOLLUSCA Numbers Issued and available: Vol. 1, No. 1 Introduction and Vasidae, 32 pp. (March 1959) $ 2.24 No. 2 Strombus, 114 pp. (November i960) 7.98 No. 3 Lambis, 28 pp. (September 1961) 1.96 No. 4 Pinnidae, 52 pp. (September 1961) 3.64 No. 5 Turrinae, 120 pp. (March 1964) 8.40 No. 6 Tridacnidae and Strombus, 58 pp. (April 1965) 4.06 No. 7 Turrinae, Terebellum, Strombus, 48 pp. (May 1967) 3.36 No. 8 Cypraea (Zoila); index to Vol, 1, 34 pp. (Dec.1967) 2.38 Permanent Binder 4.50 10 Guide tabs 1.00 If bought complete (Nos. 1-8) with binder and tabs | 39.52 (Foreign address add 48$^ § 40.00) New Subscribers: You may begin a new subscription by purchasing the last number (No. 8, 34 pages) for $2.38, or by purchasing the back numbers. You will automa- tically receive and be billed for future numbers. Back numbers may be purchased at 7 cents per page, as shown above. Because of our limited stock of the earliest numbers, we regret that back numbers can only be sold in reverse sequence; that is, No. 8 must be purchased before No. 7, No. 7 before No. 6, etc. The journal is now available only in looseleaf form. Postage paid by publisher. - J Detach here ORDER FORM FOR INDO-PACIFIC MOLLUSCA I enclose $39.52 for all that has been published, including the binder and guide tabs (foreign: $40.00) I enclose $ for the following: Name : Address : Make personal checks, bank drafts or money orders to "Academy of Natural Sciences", and mail to: A Department of Mollusks Academy of Natural Sciences 19th & The Parkway Philadelphia, Pa. 19103 U.S .A. / \ 1 1 March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 15 THE FAMILY VASIDAE IN THE INDO-PACIFIC by R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia The family Vasidae, a worldwide and warm water group of marine gastropods, belongs to the superfamily Volutacea. Because of radula and colu- mellar plicae differences, the family is divided into two major parts— the subfamily Vasinae which in- cludes the Vase shells and the subfamily Xancinae which contains the large Chank shells. Subfamily Vasinae Shells usually large, heavy, usually prominently sculptured, and with 2 to 6 irregularly-sized colu- mellar plicae on the lower half of the inner parietal wall. Periostracum thin to heavy. Operculum homy, brown, unguiculate and curved. Radula raehiglos- sate, with 3 strong teeth, the central bearing 3 strong cusps, the lateral with two cusps. (In the Xancinae, the eolumellar plicae are located much higher up on the inner parietal wall, and the lateral radular tooth has only one large cusp). Here, we are dealing with all the known Recent species of the subfamily Vasinae, except those found in the Western Atlantic (see Abbott 1950, John- sonia, vol. 2, no. 28). Of the twenty known species, five are Western Atlantic, twelve Indo-Pacific, one South Australian, one Panamic and one South African. Most of these species live in the intertidal zone or in very shallow water, although the South Australian species has been dredged at a depth of 120 fathoms. Some prefer reef flat habitats, while others are found on sandy and/or grassy bottoms. Little is known about their biology, except that they feed on clams (and possibly marine worms). The sexes are separate. The genera now assigned to the subfamily are Eovasum Douville 1920 (Eocene of Africa and South America), Vasum Roding 1798 (Recent, worldwide), Tudicla Roding 1798 (Recent, Indian Ocean) and Tudicula H. and A. Adams 1863 (Re- cent, Indo-Pacific). Wenz (1943, p. 1306) also places the genera Afer Conrad 1858, Metzgeria Norman 1879, Ptychatractus Stimpson 1865 and Piestochilus Meek 1864 in this subfamily, but fur- ther research is evidently needed before accepting these placements. Genus Vasum Roding 1798 Vasum Roding, Museum Boltenianum (2), p. 56. Type by subsequent designation (see Abbott 1950, p. 208): V. turbinellus Roding 1798 = Murex turbinellus Linne, 1758. 1810 Volutella Perry, Arcana or the Museum of Natural History, signature B 1, pi. 2. Type by monotypy: V. divergens Perry 1810 = V. muricatum Born, 1778. 1817 Cynodonta Schumacher, Essai Nouveau Systeme Habit. Vers Test., p. 73. Type by monotypy: Voluta ceramica Linne. (Cynodona on p. 241 is a spelling error. ) 1835 Scolymus Swainson, Elements of Modem Conchology, p. 21; 1840, Treatise Malacology, p. 304. Type by subsequent designation ( Abbott 1950, p. 208 ) : S. cornigerus Chemnitz = turbinellus Linne, 1758. The subgenera employed in the genus Vasum are rather nebulous in character, and it is possible that as other species are discovered, especially among the fossils, the differences will become less recog- nizable. At first glance, there seems little concho- logical similarity between Vasum (Altivasum) flin- dersi of South Australia and Vasum (Globivasum) globulus of the West Indies, yet a more or less gradual transition of generic shell characters may be seen between them in such species as horridum Heilprin (Pliocene of Florida), ceramicum Linne (Indo-Pacific) and capitellum Linne (West Indies). The subgenera of Vasum are: Subgenus Vasum Roding, 1798. Type: turbinellus Linne, 1758. Subgenus Altivasum Hedley, 1914. Type: flin- dersi Verco, 1914. Subgenus Siphovasum Rehder and Abbott, 1951. Type: latiriforme Rehder and Abbott, 1951 (Carib- bean). Subgenus Globivasum Abbott 1950. Type: globu- lus nuttingi Henderson, 1919 (Caribbean). Subgenus Vasum Roding 1798 (for synonymy see under the genus Vasum. The type is Vasum turbinellus Linne, 1758.) The shells in this subgenus are fairly large (2 to 5 inches in length ) , heavy, with slightly to moder- ately produced spines, and with 3 to 5 eolumellar plicae. The spire is moderately high, and the siphonal canal short. Only two of the Indo-Pacific Vasum are widely distributed, these being the coral reef species tur- binellus and ceramicum which extend over most [20 - 403] 16 Vasum R. T. Abbott Vasidae of the Indian and Western Pacific Oceans. The other five species are rather limited in their range, such as V. tubiferum which is only found in the Cuyo-Palawan section of the Philippines and V. rhinoceros which is known only from British East Africa. Key to the Indo-Pacific Vasum sensu stricto a Shell nodulated, without spines truncatum a Shell with blunt or sharp spines b b Inside of outer lip with black spots c b Inside of outer lip without black spots d c Lower third of columella without black armatum c Lower third of columella and lower part of outer lip with black e d Columellar plicae 3; parietal wall yellow-brown rhinoceros d Columellar plicae 5; parietal wall purple-brown tubiferum e Spire high; 3 columellar plicae strong ceramicum e Spire low; 4 to 5 columellar plicae unequal turbinellus (Vasum crosseanum Souverbie is known from a single specimen which we have not seen nor included in this key. ) 3 Plate 1, fig. 1, Vasum ceramicum (Linne), Philippines; Vasum turbinellus form cornigerum (Lamarck), Red Sea; fig. 2, Vasum turbinellus (Linne), New Guinea; fig. 3, fig. 4, Vasum armatum (broderip), Tuamotu Islands. All natural size. [20 - 404] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 17 Vasum (Vasum) turbinellus (Linne, 1758) (PI. 1, figs. 2, 3; pi. 2, fig. C) Range— East Africa to western Polynesia. Remarks— This is the commonest Vasum in the Indo-Pacific and is generally abundant wherever it occurs. It is distinguished by its relatively low spire, by the 3 strong columellar plicae between which are two additional smaller ones, and by the black, slightly raised teeth on the inner edge of the outer lip which are not bifid or in pairs as in Vasum ceramicum. V. armatum which has a pure-white columella and whose two peripheral rows of spines are almost equal in size replaces turbinellus in the central and eastern portions of Polynesia. Since Vasum is a neuter noun, adjectival names of species have a neuter ending; however, the name “turbin- ellus” is a substantive noun, meaning a “top” or “little whirlwind”, and should not be changed to turbinellum. The length of the spines on the shoulder of the shell of turbinellus varies considerably, and through- out the range of the species, particularly in the Red Sea, they may be very long and curled up- ward. These specimens may be referred to as form cornigerum Lamarck, 1822 (PI. 1, fig. 3). Habitat— V. turbinellus lives in shallow water from the low tide mark to a depth of about 30 feet. In some areas, it is fairly common on the reef flats where, during the day, it takes shelter under dead coral rocks. In other areas, it may be found on a bottom of sand, coral rubble and sparse eelgrass. Description— Adult shell 30 to 80 mm. in length ( 1 to 3 inches), solid, heavy, turbinate and strongly spined. Spire moderately elevated. Whorls 7 to 9, the body whorl bearing at the shoulder 8 to 9 stout, blunt, upwardly pointing spines which are slightly open at their ends; immediately below is a similar row of much smaller spines. The middle of the whorl bears 2, rarely 3, spiral cords which may each bear 8 to 15 small knobs. At the base of the shell there are three rows of 7 to 8 bluntly conical spines. Suture indistinct and wavy. Parietal wall glazed, yellowish white with black-brown mot- tlings. Columella with three large plicae between which may be 2 additional, smaller ones. Outer lip slightly thickened, somewhat crenulated, creamy- white with 4 to 6 squarish black spots over the slightly raised, short teeth on the outer lip. Lower third of columella usually with a brownish stain. Umbilicus absent or rarely a small indentation. Color of outer shell grayish with sparse or heavy blackish brown mottlings. Periostracum thin, weakly foliaceous, and grayish yellow or gray-brown in color. Operculum fills most of the inner aperture, corneous, blackish and unguiculate. Measurements (mm.) (including spines) — length width no. whorls 85 73 6 ( large; Dutch New Guinea ) 50 48 6 ( average; Palau Islands) 44 35 6 (small; Marshall Islands) Synonymy — 1758 Murex turbinellus Linne, Systema Naturae, ed. 10, p. 750, no. 466 (in O. Asiatico ad Nussaanan); refers to Rumphius Mus., pi. 24, fig. B, and others. 1767 Voluta turbinellus Linne, Systema Naturae, ed. 12, p. 1195, no. 430; 1955, Dodge, Bull. Amer. Mus. Nat. Hist., vol. 107, art. 1, pp. 127-129. 1811 Volutella nigra Perry, Conchology, London, pi. 26, fig. 1 (African Seas). 1822 Turbinella cornigera Lamarck, Anim. sans Vert., vol. 7, p. 105, no. 7 (Moluques); 1840, Kiener, Co- quilles Vivantes, vol. 6, Turbinelle, pi. 1 1822 Turbinella variolaris Lamarck, Anim. sans Vert., vol. 7, p. 110 (locality not given); 1840, Kiener, Co- quilles Vivantes, vol. 6; Turbinelle, pi. 21, fig. 1 (a young, worn specimen). Plate 2. Single rows of radulae from adult Vasum ( greatly enlarged). Fig. A, Vasum rhinoceros (Gmelin), Zanzibar; B, Vasum armatum (Broderip), Toau Atoll; C, Vasum tur- binellus (Linne), Philippines; D, Vasum ceramicum (Linne), Philippines. [20 - 405] 18 Vasum R. T. Abbott Vasidae Types— Linne’s type locality is “ad Nussaanan,” a place quoted from Rumphius and probably refer- ring to the small islet of Nusa Laut, to the east of Ambon near Saparoea and Haroekoe (formerly Ona) Islands, Indonesia. Possible cotypes of tur- binellus are in the Linnaean Society collection in London (Dodge, 1955, p. 128). Locality records — See accompanying map, p. [20-406] solid dots: specimens examined; open circles: from the literature. Selected records: SOUTH AFRICA: Natal (Natal Mus.). EGYPT: Berenice, Foul Bay (ANSP) SAUDI ARABIA: Jedda (C. Aslakson, ANSP). KENYA COLONY: Mombasa (B. Verdcourt; ANSP). RYUKYU IDS.: Okinawa (W. A. McCarty, ANSP). POLYNESIA: Vailele Bay, Upolu Id.; Samoan Ids. (N.S.F. ); Johnston Id. ( D. Thaanum); Howland Id. (Ted Dranga, ANSP). Vasum (Vasum) armatum (Broderip, 1833) (PI. 1, fig. 4; pi. 2, fig. B) Range— Eastern Polynesia from the Phoenix Is- lands to the Tuamotu Islands (not Hawaiian Is- lands). Remarks— The fusiform shape, white lower colu- mella, and the two rows of equal-sized, conic spines at the shoulder distinguish this Polynesian species from turbinellus. Tryon (1882, Manual of Con- chology, vol. 4, p. 72) considered this species as a synonym of ceramicum. This species appears to be limited to low coral islands and atolls of eastern Polynesia. “Animal above pale flesh-colour, finely mottled with purplish; siphon and locomotive disk salmon- coloured; eyes on a bulbous expansion near tip of tentacles. Operculum yellowish” (Couthouy’s notes in Gould 1852). Habitat— This appears to be a moderately com- mon to abundant species in pockets of sand on the reef flats facing the open ocean in the intertidal zone. Description— Adult shell 23 to 70 mm. in length (usually about 52 mm.) (1 to 2& inches). Solid, heavy, somewhat fusiform, and moderately heavily spined. Spire elevated. Whorls 6 to 7, the body whorl bearing at the shoulder two rows of 7 to 8 almost equal-sized, conic spines which are slit along their anterior faces. The middle of the whorl bears 3 or 4 small, rough, spiral cords, below which is first a single row of small, well-developed spines, and then, at the base, 1 or 2 rows of weak nodules. Suture indistinct and wavy. The entire outer shell has numerous, coarse, raised, wavy threads. Parietal wall glazed, enamel-white, and rarely with brown markings at the top. Columella white, usually with °o °o o *b °o CM O CM [20 - 406] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 19 4 equal-sized, slanting plicae, rarely with a weak fifth. Inside of body whorl with about 8 small, raised spiral cords. Outer lip thickened and with 4 or 5 pairs of black, rounded teeth. Umbilicus absent. Color of outer shell bluish gray to grayish white with diffused mottlings of bluish brown which are usually more concentrated in a band just below the suture and on the middle of the last whorl. Periostracum thin, translucent, weakly foliaceous. Operculum corneous, claw-shape, light yellowish brown in color, and with its outer surface with very coarse, uneven lines of growth which give it a gnarled appearance. Measurements (mm.) — length 74 width 57 no. whorls 7 + ( large; Henderson Atoll ) 36 30 7 ( average; Henderson Atoll ) 26 18 7 (small; Henderson Atoll) Synonymy — 1833 Turbinella armata Broderip, Proc. Zool. Soc. London, vol. 1, p. 7 (ad Insulam Elizabethae ) ; 1847, Reeve, Conch, Icon., vol. 4, Turbinella, pi. 5, fig. 29; 1876, Kobelt, Syst. Conchyl. Cab., vof. 3, no. 3, pi. 16, fig 2 ( good). 1852 Turbinella armata Broderip, Gould, U.S. Exploring Exped., vol. 12, p. 238, pi. 17, figs. 290, 290a, 291 ( animal ) . 1933 Vasum ceramicum var. armata Broderip, Dautzenberg and Bouge, Joum. de Conchyl., vol. 77, p. 206. ( Anaa Atoll). Type Locality— On coral reef, Elizabeth Island [Henderson Id., Tuamotu Islands] Hugh Cuming, collector. The types are in the British Museum. Possible cotypes are in ANSP 35230. Locality records— TUAMOTU ISLANDS: Puka-puka Id. ( Honden Id., Gould 1852, p. 238); Toau Atoll ( H. A. Pils- bry); Takaroa Atoll (H. A. Pilsbry); Fakarava id., Makemo Id., Tikahau Atoll, Raroia Atoll (all USNM). Henderson Id., near Pitcairn (Cuming). Flint Island (USNM). Fan- ning Island ( B. P. Bishop Museum). PHOENIX ISLANDS: Hull Id. (USNM). EASTERN SAMOA: Rose Atoll (USNM). See accompanying map, p. [20-406], Vasum (Vasum) ceramicum (Linne, 1758) (PI. 1, fig. 1; pi. 2, fig. D) Range— British East Africa to eastern Polynesia. Remarks— The Ceram Vase is the largest member of this genus in the Tropical Indo-Pacific province, being exceeded in size only by Vasum (Altivasum) flindersi Verco of South Australia. It is distin- guished from the smaller and commoner V. turbi- nellus by its high spire, three strong, slanting colu- mellar plicae (rarely with one or two additional weak plicae), and the small paired teeth on the in- side of the outer lip. It differs from armatum of eastern Polynesia in having a blackish brown patch on the lower end of the columella. Tryon (1882, Manual of Conchology, vol. 4, p. 72) erroneously considered Latirus vexillulum (Reeve) a synonym. Habitat— Moderately common in surging waters off the front edge of coral reefs, usually in depths of 3 to 30 feet. Description— Adult shell 80 to 140 mm. in length (3 to 5M in.). Solid, heavy fusiform, and rather strongly spined. Spire elevated. Whorls 9 to 11, the body whorl bearing at the shoulder 7 to 10 strong, somewhat triangular, outwardly projecting spines which are slit open on their anterior faces; imme- diately below are two similar rows of much smaller spines. Below this are 2 to 4 small, rough spiral cords. The base of the shell bears 1 or 2 rows of small spines. Suture indistinct and wavy; below it on the upper part of the shoulder there are 4 to 6 distinct, raised, spiral threads. Parietal wall glazed, all white or sparsely mottled with black-brown. Columella with 3 very strong, slightly slanting, squarish plicae between which are rarely 1 or 2 more very weak plicae. There are a dozen or so weak, white, spiral cords within the aperture on the inside wall of the body whorl. Outer lip slightly thickened, crenelated and bearing 5 or 6 pairs of small, black, raised teeth. Lower fourth of the columella usually with a brownish black stain. Umbilicus chink-like and shallow. Color of outer shell whitish with heavy mottlings of black or black-brown. Periostracum thin, varnish-like, trans- lucent brown, somewhat foliaceous. Operculum fills most of the inner aperture, is corneous, claw-shaped, blackish, and with a muscle scar of a little more than /3 the area of the inner side. Odontophone % the length of the shell, and with 150 rows of radu- lar teeth (see pi. 2, fig. D). Measurements (mm.) (including spines) — length width no. whorls 148 94 9 + (large; Guam Id. ) 125 81 9 + (average; Zanzibar) 81 56 7 + (small; Guadalcanal Id. ) Synonymy — 1758 Murex ceramicus Linne, Systema Naturae, ed. 10, p. 751, no. 470 ( O. Asiae ad Ceram). Refers to Rum- phius, pi. 24, fig. A, pi. 49, fig. L; Bonanni, pi. 286; and others. 1767 Voluta ceramica Linne, Systema Naturae, ed. 12, p. 1195, no. 432; 1955, Dodge, Bull Amer. Mus. Nat. Hist., vol. 107 art. 1, p. 130-132. 1807 Turbinellus spinosus G. Fischer (von Waldheim), Museum Demidoff, Moscow, p. 205 (Tranquebar et Nicobar). Refers to Gualtier, pi. 55, fig. D; Lister, pi. 829-51; Chemnitz Conchyl. -Cab., vol. 11, figs. 1725, 1726, the latter being turbinellus Linne. Types— The type locality is Ceram Island, Indo- nesia. The possible type is in the Zoological Museum of the University of Uppsala, Sweden (Dodge, 1955, p. 131). [20-407] 20 Vasum R. T. Abbott Vasidae Locality records— (see accompanying map). Selected rec- ords-. ZANZIBAR: Chango Id.; Pange Id.; Chumbe Id. CEYLON: Hikkaduwa (G. F. Kline, ANSP). NEW GUINEA: Wombrisau, Biak Id. (NSF). SOLOMON IS- LANDS: Lunga, Guadalcanal (ANSP). CAROLINES: Ifaluk Atoll (USNM); Round Rock, Helen Reef (NSF); PALAU ISLANDS: Babelthuap Id. (NSF); Koror Id. (NSF). GILBERTS: Apamama (USNM). RYUKYU IS- LANDS: Okinawa Id. (W. A. McCarthy, ANSP). FIJI ISLANDS: Makuluva Id., and Suva, Viti Levu Id. (USNM ); Bega Id. (ANSP). NIUAFOU ISLAND (USNM). SAMOA: Fagalii Bay, Upolu Id. (NSF). SOCIETY ISLANDS: Moorea Id. (USNM). Vasum (Vasum) tubiferum (Anton, 1839) (PI. 4, bg. 1; pi. 3) Range — Cuyo Islands, west central Philippines (also Mindanao Island ?). Remarks— The Imperial Vase is a very distinctive and highly localized species known, so far, only from one area in the Philippines. It resembles V. turbinellus, but differs in having a deep and funnel- shaped umbilicus, purple to lavender splotches on the parietal wall, a “furry” brown periostracum and lacking black spots on the inside of the outer lip. Conchologically, this species most resembles V. rhinoceros from East Africa. Melvill and Standen (1895, Jour. Conch., vol. 8, p. 104) report this species from Lifu Island, Loyalty Islands, but I suspect these may be turbinellus. I am indebted to Dr. Ramon Lim of Cuyo City, Palawan, for sending preserved specimens for study. Habitat— A Philippine shell collector told us that this is a moderately common species found in very shallow water in quiet bays where there is a sand and eelgrass bottom. Description— Adult shell 60 to 116 mm. in length (about 2M to 4M inches). Solid, heavy, turbinate and strongly spined. Spire rather well elevated. Whorls about 7, the body whorl bearing at the shoulder 8 long, upwardly curving spines which are flaringly open at their ends. Below are 4 or 5 spiral, coarse cords of decreasing size which may bear raised nodules or very small open-faced spines. Base of the shell with a row of well-developed, open-faced spines, below which are two much weaker rows of low, scale-like spines. Suture indis- tinct. Parietal wall well-developed, slightly raised, glazed and colored a light tan with large splotches of chestnut- or purple-brown. Columella bears 5 plicae, the upper, lower and middle ones being the largest. Interior of aperture white and smooth. Outer lip thickened, slightly reflected, coarsely crenulated, and tan or whitish in color. Lower third of columella white. Umbilicus funnel-shaped and deep. Color of outer shell orange-brown to yellow- ish. Periostracum thick, brown, and strongly foli- aceous. Operculum corneous, dark-brown, unguicu- late, and its exterior with fine, irregular growth lines. Measurements (mm.) (including spines) — length width no. whorls 116 90 7 (large; Cuyo Id.) 88 70 7 (average: Cuyo Id.) 74 60 8 (small; Cuyo Id.) Synonymy — 1839 Turbinella tuhifera Anton, Verzeichniss der Conchyl., Halle, p. 70 (no locality); 1876, Kobelt, Syst. Conchyl. Cab., vol. 3, no. 3, p. 155, pi. 9, fig. 3. 1842 Turbinellus imperialis Reeve, Conch. Systematica, vol. 2, p. 181, pk 229, fig. 4 (no locality); 1843, Reeve, Proc. Zool. Soc. London, vol. 10, p. 198 (no local- ity); 1847, Reeve, Conch. Iconica, vol. 4, Turbin- ella, pi. 5, fig. 28 ( Cagayan, Island of Mindanao, Philippines). Types— Anton gave no locality; nor did Reeve in 1842 and 1843 for his imperiale. In 1847, Reeve states that Hugh Cuming collected it at Cagayan, Island of Mindanao, Philippines. I have looked for this species at Cagayan and elsewhere on Min- danao without success, and it is possible that Cum- ing meant Cauayan Island in the Cuyo— Palawan group where this species is known to exist. Reeve’s type in the British Museum is 78 mm. in length, and the purple-brown on the parietal wall has faded to brown. Plate 3. Vasum tubiferum (Anton), Cuyo Id., Philippines. Figs. A, external verge, showing cross-section; B, three- quarter view of odontophore; C, two rows of radular teeth; D, attachment side of operculum. [20 - 408] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 21 Records— Hugh Cuming is supposed to have col- lected this species at Cagayan, Misamis Prov., Min- danao Id., Philippines, although there is some doubt about this. A Philippine shell collector brought us two dozen specimens during the 1958 du Pont— Academy Expedition which he had collected on Cuyo Island, Palawan Province, Philippines. Vasum (Vasum) rhinoceros (Gmelin, 1791) (PI. 4, figs. 3, 4) Range— Kenya Colony and Zanzibar, British East Africa. Remarks— The Rhinoceros Vase is common, but only found, in Kenya and Zanzibar. It is distin- guished by its three single columellar plicae, mas- sive nodules on the shoulder, brown-splotched parietal wall and by the absence of black markings on the outer lip. An unusual color form occurs in some colonies in Zanzibar in which the brown maculations on the outer shell are absent and in which the parietal wall is a pure light-yellow. Habitat— This species occurs in fairly large col- onies in shallow water on a rock, sand and weedy bottom just inside fringing coral reefs. Rarely, it is dredged on a sand and rubble bottom to a depth of 48 feet. Description— Adult shell 45 to 85 mm. (II to 3/2 inches) in length. Solid, heavy, varying in shape from turbinate to sub-fusiform, and with blunt or sharp spines. Whorls 7 to 8, the body whorl bearing at the shoulder one or two rows of 5 to 8 spines which may be large and blunt, open or closed at their ends, or rather pointed and somewhat tri- angular. The middle of the body whorl bears 6 to Plate 4, fig. 1, Vasum tubiferum (Anton), Cuyo, Philip- Figs. 3 and 4, Vasum rhinoceros (Gmelin), Zanzibar; fig. 3 pines; fig. 2, Tudicula rasilistoma new species (young shell), is the yellow form; fig. 4 is normal. All natural size. Queensland. [20 - 409] 22 Vasum R. T. Abbott Vasidae 10 small, indistinct, unevenly-sized spiral cords. Base of shell with a single row of fairly prominent, open-ended spines, below which are 3 or 4 indis- tinct, sometimes spined, cords. Nuclear whorl rather large, bulbous, chalky white, followed by numerous, small axial riblets. Axial sculpture of fine foliaceous growth lines which may develop into fine scales. Parietal wall thick, raised, glazed, and colored either chocolate-brown with tan mottling or a pure light-yellow. Columella with 3 distantly- spaced, rather strong, slightly slanting, purplish white plicae, the lowest being the smallest. Aper- ture elongate, tapering below, and within it is whitish with a large, diffused, mauve-brown splotch and with about 24 fine, spiral ridges. Outer lip thickened, slightly reflected, crudely crenulate, and glossy tan. Umbilicus partially open, narrow, some- times deep, sometimes sealed. Color of outer shell cream with light-brown mottlings and specks; rarely all yellowish. Periostracum thin, translucent brownish, and slightly foliaceous. Operculum cor- neous, unguiculate, light-brown, and its outer sur- face with several weak longitudinal striations. For radula, see pi. 2, fig. A, p. [20-405], Measurements (mm.) (including spines) — length width no. whorls 92 76 8 + (large; Zanzibar) 73 58 8 (average; Zanzibar) 57 40 7 ( small; Kenya ) Synonymy — 1791 Valuta rhinoceros Gmelin, Systema Naturae, ed. 13, p. 3458, no. 128 (ad Novae Guineae); refers to Chemnitz, Conchyl.-Cab., vol. 10, figs. 1407, 1408. 1822 Turbinella rhinoceros Gmelin, Lamarck, Anim. sans Vert., vol. 7, p. 105 (Nouvelle-Guinee); 1847, Reeve, Conch. Icon., vol. 4, Turbinelle, sp. 33 (Zanzibar); 1876, Kobelt, Syst. Conchyl. Cab., vol. 3, no. 3, pi. 16, fig. 1 (good), pi. 6, figs. 2, 3 (poor). Types— Gmelin’s type locality of “New Guinea” was evidently erroneous, and it was not until 1847 that the proper locality of Zanzibar was recorded by Reeve. Gmelin did not have a type specimen. Locality records— KENYA: Malindi; Kilifi; Vipingo; Mom- basa (all Coryndon Mem. Mus. ); Diani Beach (Abbott, USNM). ZANZIBAR: Chumbe Id. (4-6 fms.); Mangapwani (intertidal); Ras Nungwe (1 fm.); Fumba (intertidal); Mnemba Id. (intertidal); Paje; Mnemba Id. (intertidal); Kiwengwa; Chwaka; Chango Id.; Chukwani; Chumbe Id.; off Kisiki Id. (9 fms.). (all Ostheimer, Orr and Thorington, 1957, ANSP ). [20 - 410] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 23 Vasum (Vasum) truncatum (Sowerby, 1892) ( Plate 5 ) Range— Eastern Cape of Good Hope and south- ern Natal, South Africa. Remarks— The South African Vase is evidently a rare species, and, although specimens have been found dead on the beach, Quekett (E. A. Smith, 1903, p. 370) reports it from “deep water”. It is readily recognized by its white shell, almost conic shape and its low, weak nodules. Description— Adult shell 65 to 72 mm. (about 2 to 3 inches) in length, solid whitish, subtriangular and nodulated. Spire flattish, except for the two raised, smooth, papillate nuclear whorls. Whorls 8, the shoulder somewhat carinate and bearing 9 to 11 low but distinct and slightly pointed nodules. Sides of last whorl slightly concave and bearing 5 indis- tinct rows of very weak, rounded nodules. Suture distinct and very wavy. Aperture white within. Columella and thickened outer lip blotched with brown. Upper end of parietal wall bears a whitish, swollen callus. Columellar plicae 4, the uppermost the largest. In young specimens, umbilicus present and chink-like. Periostracum rather thick, decidu- ous, matted, and light-brown in color. Measurements (mm.) — length width no. whorls 65 50 8 (from Sowerby, 1892) 71 52 _ (from E. A. Smith, 1903 50 35 7 (immature, Pondoland) Plate 5. Vasum truncatum (Sowerby), Union South Africa ( natural size; from the Proc. Mai. Soc. London, vol. 5, pi. 15, fig. 3). Synonymy — 1892 Turbinella truncata Sowerby, Marine Shells of South Africa, London, p. 17, pi. 4, fig. 85 (Port Eliza- beth ) . 1902 Turbinella triangularis E. A. Smith, Jour. Conchology, vol. 10, p. 249, pi. 4, fig. 6; 1903, Proc Mai. Soc. London, vol. 5, p. 370, pi. 15, fig. 3 (off Durban and Port Shepstone). 1915 Xancus truncatus Sowerby, Bartsch, Bull. 91, U.S. Nat. Mus., p. 42 (Port Alfred). Types— Port Elizabeth is the type locality. Sow- erby states that the type collected by S. D. Bair- stow was deposited in the Oxford University Museum, England. The type of triangularis is in the British Museum (Natural History), London. Locality records— SOUTH AFRICA: Port Eliza- beth; Port Alfred; Fossil head, Pondoland, (Natal Museum, 3869), all Cape of Good Hope; Port Shepstone; Off Durban, both Natal. Vasum (Vasum) crosseanum (Souverbie, 1875) (Plate 6) Range— Presumably Madagascar. Remarks— There has been no further information on this evidently rare species since it was first de- scribed by Souverbie in 1875. Tryon (Manual of Conchology, vol. 4, p. 71, 1882) erroneously con- V.7 Plate 6. Vasum crosseanum (Souverbie). Madagascar? (natural size; from the Joum. de Conchyl., vol. 24, pi. 13, fig. 1). [20-411] 24 Vasum R. T. Abbott Vasidae sidered this a worn and abnormal specimen of Vasum muricatum (Linne) from the West Indies, but I feel certain that it is a good species awaiting re-discovery. It appears to be characterized by 5 well-developed columellar plicae, bulbous apex, rounded body whorl, yellowish white mouth, and slight umbilicus. This species is provisionally placed in the subgenus Vasum. Description— Shell 95 mm. (3/2 in.) in length, solid, pyriform, very deeply and narrowly umbili- cate and whitish in color with subdued bands of yellowish rose. Whorls 8, the first 4 mammillate and with numerous axial costae. Suture wavy, impressed and with fine fimbriations. Spiral sculpture of numerous uneven, raised, rough threads. Axial sculpture of moderately developed, somewhat pointed nodules on the shoulder of the whorls. Aperture elongate-oval, with a thickened, slightly reflected outer lip and with a strong, raised, yel- lowish white, glossy parietal shield. Columella bears 6 spiral plicae, the 2nd, 4th and 6th being strong, and the 1st, 3rd and the 5th being weak. Siphonal canal short and open along its entire length. Umbilicus narrow and very deep. Opercu- lum and soft parts unknown. Measurements ( mm.) — Length 95, width 65, length of aperture 80, width of aperture 32. Synonymy — 1875 Turbinella crosseana Souverbie, Jour, de Conchyl., vol. 23, p. 297; 1876, ibid, vol. 24, p. 382, pi. 13, fig. 1 ( Mauritius or Madagascar ? ) Types— The type is in the National Museum of Natural History in Paris, France. Records— Unknown, except for Souverbie’s sur- mise that it comes from Madagascar or Mauritius. March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Vasum 25 Subgenus Altivasum Hcdley 1914 This subgenus includes those Vasum species which have evolved in the direction of delicate spinosity, elevated spire, and rather shortened an- terior siphonal canal. Evidently the only Indo-Pacific representative is the subgenotype, flindersi. In the Western Atlantic region, it is represented by the Recent V. capitellum Linne and by two Miocene species (subcapitellum) Heilprin and horridum Heil- prin. Vasum (Vasum) ceramicum Linne of the Indo- Pacific bears a superficial resemblance to members of this subgenus by its possession of an elevated spire, but is excluded by its 5 columella plicae, fewer and stouter spines, and lack of an umbilicus. Description— Shell moderately large, rather heavy and solid, with a well-elevated spire, and bearing several to many spiral rows of long, scale-like spines. Columella with 3 slanting plicae. Umbilicus funnel- shaped and usually deep. Siphonal canal moderately to greatly shortened. Periostracum, thin, coriaceous, brownish yellow in color. Operculum horny, ungui- culate, and with a terminal nucleus. Synonymy — 1914 Altivasum Hedley, Biological Results . . . F. I. S. “En- deavour” 1909-1914, vol. 2, pt. 2, p. 68, pi. 9. Type by monotypy, Latirus aurantiacus Verco 1895, non Montfort 1810 = Altivasum flindersi Verco, 1914); 1914, Hedley, Trans, and Proc. Royal Soc. South Australia, vol. 38, p. 484. 1950 Vasum (Altivasum) Hedley, Abbott, Johnsonia, vol. 2, no. 28, p. 213. Vasum (Altivasum) flindersi Verco, 1914 (PI. 7, figs. 1, 2) Range— From the Gulf of St. Vincent, South Aus- tralia, to Nurina, Western Australia, from 18 to 120 fathoms. Remarks— This is the largest and most attractive of the Indo-Pacific Vasidae. It is considered a rare shell, at least in collections, probably because of the infrequency at which it is dredged by commer- cial fishermen off the southern coast of Australia. This species is readily distinguished from Vasum ceramicum Linne by its numerous, open spines, wide and deep umbilicus, proportionately high spire and salmon-chalk color. Young specimens lack spines and have a narrower umbilicus, thus giving them the appearance of a Latirus. Habitat— Dredged in 18 to 120 fathoms. Description— Shell large, 130 to 160 mm. (5 to 6 inches) in length, solid, strongly spinose, and whit- ish orange to pure white in color. Whorls about 12, the early ones bearing 9 to 11 blunt, axial nodules which in the last 3 or 4 whorls develop into long, slightly recurved, anteriorly open, tubular spines. Nuclear whorls 1/2, slightly swollen, round and smoothish. Spiral sculpture on post-nuclear whorls of 5 to 6 raised threads. Suture wavy, indented and bordered below by a raised thread which becomes well fimbriated in the last 3 whorls. Shoulder of the last whorl bears the largest spines, and below this row are 7 to 8 crowded spiral rows of smaller spines. Aperture ovate, white within. Columella bears 3 small, spiral, slightly slanting plicae. Umbilicus rather large, funnel-shaped and very deep. Perio- stracum thin, translucent-brown and coriaceous. Measurements (mm.) — length width no. whorls 160 98 11 (adult, ANSP ) 130 65 — (fide Hedley, 1914) 95 52 8 (immature, ANSP) Synonymy — 1895 Latirus aurantiacus Verco, Trans. Royal Soc. South Australia, vol. 19, p. 89 [79], pi. 2, figs. 1, la (Back- stairs Passage, South Australia, 18.5 fms. ) 1914 Altivasum flindersi Verco, Trans, and Proc. Royal Soc. South Australia, vol. 38, p. 484. New name for L. aurantiacus Verco 1895, non Montfort 1810. Types— The type locality is Backstairs Passage, South Australia, Australia, in 18.5 fathoms. The holotype, an immature specimen 46 mm. in length, is in the South Australian Museum in Adelaide. Plate 7. Vasum (Altivasum) flindersi Verco, Great Aus- tralian Bight. Fig. 1, adult; fig. 2, young. (Both half natural size). [20-431] 26 Altvasum R. T. Abbott Vasidae Locality records— SOUTH AUSTRALIA: Back- stairs Passage, 18.5 fins. (Verco, 1895); off New- land Head, 20 fms. (Verco, 1914); Gulf of St. Vincent (Hedley, 1914); Great Australian Bight ( ANSP). WESTERN AUSTRALIA: Great Aus- tralian Bight from 126° to 129° East Long., 120 fms. (Hedley, 1914). [20 - 432] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 T udicula 27 Genus Tudicula H. and A. Adams 1863 Recent species of this genus of Vasidae are known only from the Indian Ocean and the north- ern half of Australia, although some probably exist in the East Indies. On the whole, they may be con- sidered uncommon or rare, perhaps because of the difficulty in dredging for them at depths ranging from 4 to 20 fathoms. One species, armigera, is occasionally found living in the intertidal zone. Members of this genus are undoubtedly carnivor- ous, and, so far as we know, live on sand and rocky bottom. There are five known Recent species. A number of fossil species have been described under both T udicla and T udicula, but only sinotecta Lud- brook, 1941 from the Pliocene of South Australia (Trans. Royal Soc. South Australia, vol. 65, p. 97, pi. 5, fig. 14) seems to belong to true Tudicula. Synonymy — 1863 Tudicula H. and A. Adams, Proc. Zool. Soc. London for 1863, p. 429 (as a subgenus of Tudicla). Type by subsequent designation (W. Wenz, 1943, p. 1303 ) : T. armigera A. Adams. Description— Shells medium in size, 30 to 70 mm. in length, moderately solid, usually pyriform or fusiform, sometimes spinose, and with a long, nar- row, anterior siphonal canal. Whorls 4 to 7, usually spinose, rarely smoothish, but always with fine spiral threads. Nuclear whorls rather large and slightly mammillate. Parietal shield usually well- developed. Lower third of columella with 3 or rarely 4, moderately developed, slanting plicae. Operculum corneous, unguiculate, brown, and with a terminal nucleus. Radula rachiglossate, the cen- tral tooth with 3 cusps, the lateral teeth with two cusps. Body and penis similar to those in Vasum. Key to the Indo-Pacific Tudicula a Spire angle less than 85° b a Spire angle greater than 85° c b With 7 shoulder knobs per whorl rasilistoma b With 10 to 12 shoulder spines per whorl armigera c Aperture oval-round; white d c Aperture oval-elongate; orange zanzibarica d Shoulder with 12 to 14 spines spinosa d Shoulder without spines inermis Tudicula (Tudicula) armigera (A. Adams, 1855) (PI. 9, figs. 9, 10) Range— Queensland, Australia. Remarks—1 This handsome Tudicula is moderately common in some parts of Queensland. It is char- acterized by the single row of well-developed shoulder spines, and the two rows of spines on the rather slender siphonal N canal, and by its white parietal wall. The length of spines in adult shells is variable, some being 5 mm. in length, others as long as 14 mm. The color of the shell may vary from yellowish white to reddish brown. This is the type of the genus Tudicula H. and A. Adams. Habitat— This species lives on sand and rubble bottom from low tide mark to a depth of 20 fathoms. Description— Adult shell 55 to 72 mm. (2M to 2% inches) in length, solid, pyriform with a long, thin anterior siphonal canal, and spinose. Color white to yellowish cream, with or without light-brown to purplish brown flecks or small maculations. Nuclear whorls 1/2, proportionately large, rounded, smooth. Plate 8. Figs. 1-3, Tudicula zanzibarica Abbott, paratype. Figs. 4-6, Tudicula inermis Angas, off Broome, Western 1, dorsal view of animal with cross-section of verge; 2, Australia. 4, radulae; 5, side view of central tooth; 6, radulae; 3, operculum. operculum. [20 - 443] 28 T udicula R. T. Abbott Vasidae semi-glossy, and white to purple-brown. First post- nuclear whorl weakly cancellate; the next 5 whorls bearing 6 to 7 raised, finely fimbriated, spiral threads; the lowest thread, located just above the wavy, well-indented suture, bears increasingly longer, open-faced spines ( 10 to 12 per whorl ) . The long siphonal canal bears 2 spiral rows of long, delicate spines. The numerous spiral threads on the body whorl may bear numerous, very small spines. Axial sculpture of 10 to 12 low, rounded folds. Aperture ovate. Outer lip slightly reflected, thick- ened, glazed and weakly crenulate. Parietal shield well-developed, raised, glossy white to cream. Lower third of columella with 3 (rarely with a weak 4th), slightly slanting plicae, the lowermost being the broadest. Operculum corneous, chestnut- brown, unguiculate, and with a terminal nucleus. Measurements (mm.) (including spines) — length width no. whorls 71.4 31.0 6 (large; off Bundaberg) 65.1 30.8 6 (average; Palm Isle) 56.5 28.0 5 (small; Pancake Creek) Synonymy — 1855 Ttidicla armigera A. Adams, Proc. Zool. Soc. London for 1854, p. 221 ( Moreton Bay). 1884 Turbinella (Tudicula) armigera A. Adams, E. A. Smith, Report Zool. Collections H. M. S. “Alert”, London, p. 53, pi. 5, fig. G. Types — The type locality is Moreton Bay, Queensland, Australia. The type is in the British Museum (Natural History), London. Plate 9. Figs. 1 and 2, Tudicla spirillus Linne, east coast of India. Figs. 3 and 4, Tudicula inermis Angas; 3, from Exmouth Gulf, West Australia; 4, holotype (Cornell Univ. Paleo. Mus. 19232). Figs. 5 and 6, Tudicula spinosa H. and A. Adams; 5, off Darwin, West Australia (A. R. Cahn collection at ANSP); 6, Torres Straits, Queensland ( Mrs. W. Barker collection ) . Figs. 7 and 8, Tudicula zanzibarica Abbott, off Zanzibar; 7, para- type (ANSP); 8, holotype (ANSP). Figs. 9 and 10, Tudicula armigera A. Adams; 9, Pancake Creek, Queensland (ANSP); 10, off Bundaberg, Queensland (MCZ). All natural size. [20 - 444] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 T udicula 29 Locality records— QUEENSLAND: Palm Isle in 18 fms. (ex J. Brazier, ANSP); Pancake Creek, Bustard Head (ANSP); off Bundaberg in 20 fins. (MCZ); Moreton Bay, Port Curtis, 0 to 11 fms., and Port Molle in 14 fms. (E. A. Smith, 1884, p. 53). Tudicula (Tudicula) rasilistoma new species (PI. 4, fig. 2, p. [20-409]) (PI. 10, figs. A-C) Range— Known only from northern New South Wales and southern Queensland, Australia. Remarks — The recently discovered “Polished- mouthed” Tudicula from northeast Australia is evi- dently related to T. armigera, but differs in having two brown spots on the parietal wall, in having a shorter, stouter and brown-tipped siphonal canal, and in having 7 to 8 weakly spined or smooth nodules per whorl (instead of 10 to 12 strong spines per whorl). The apex of most adult shells is eroded away. Young specimens have a delicate pink to rose aperture and lack the parietal shield. Habitat— Nothing is known about the habitat or habits of this species, except that it occurs in 30 fathoms of water ( according to shrimp fishermen ) . Description— Adult shell 56 to 72 mm. (2M to 3 inches) in length, solid, fusiform, weakly spinose, and with a moderately short, stout siphonal canal. Color chalky pinkish cream to whitish and overlaid with irregular bands of various shades of brown. Nuclear whorls VA, smooth, rounded and opaque- white. Body whorl with two closely set, peripheral rows of 7 (rarely 8) blunt nodules. The midpoint of the stout anterior siphonal canal bears a row of 5 or 6 fairly long, slender, open-faced spines, below which may be a second row of obsolete spines. Lower third of siphonal canal an almost solid, dark chocolate-brown. Umbilicus chink-like, shallow or absent. Parietal wall well-developed, raised, glossy cream to pinkish and with a brown blotch on the left center and at the posterior or upper end. Colu- mellar plicae 3, the middle one being the largest and most distinct. Outer lip slightly reflected, sharp, strong, but finely and unevenly crenulate. Inner wall, behind the outer lip, with weakly developed, raised, spiral lirae. Periostracum grayish brown, translucent, thin, and microscopically fimbriated. Operculum corneous, unguiculate and dark-brown in color. Measurements ( mm.) — length width no. whorls 72.0 39.0 8 ( holotype ) 59.0 34.5 7 ( Tin Can Bay, Queensland ) 58.0 33.0 5? (paratype, ANSP) Synonymy — I am indebted to Miss J. Hope Macpherson of Victoria who suggested the name and asked that I describe this species. Types— The type locality is off Tweed Heads, northern New South Wales, dredged in 30 fathoms. Holotype (F18189) and paratypes in the Nat. Mus. Victoria. One paratype from Brisbane in ANSP no. 227669. Locality records— QUEENSLAND: Caloundra (Mrs. L. Brown Coffin.); off Brisbane (Nat. Mus. Victoria and ANSP); dredged in Tin Can Bay (ANSP). NEW SOUTH WALES; 30 fathoms, off Tweed Heads ( Nat. Mus. Victoria). Plate 10. Tudicula (Tudicula) rasilistoma new species, holo- type from New South Wales. Immature paratype: A, oper- culum; B, shell; C, nuclear whorls enlarged. (Courtesy of J. Hope Macpherson). Tudicula (Tudicula) spinosa II. and A. Adams, 1863 (PI. 9, figs. 5, 6, p. [20-444]) Range— Northern Territory to Queensland, Aus- tralia. Remarks— This small and attractive species is found in a few fathoms of water along the Queens- land coast and westward to the Darwin area. It is rather rare in collections, but, to judge from the number of specimens sent to us from the Arafura Sea by Colonel A. R. Cahn, it is not uncommon in its habitat. The species is recognized by the single series of small sharp spines on the periphery of the shoulder. Most specimens are cream in color with small and numerous reddish brown flecks, although some may be pure white or with a wide, broken, subdued band of brownish on the body whorl. Habitat— Unknown, except for the fact that it has been dredged in 9 fathoms of water. [20 - 445] 30 T udicula R. T. Abbott Vasidae Description— Adult shell 40 to 50 mm. ( 2 inches ) in length, turnip-shaped, finely spinose, with a long aspinose siphonal canal, and white with yellowish brown flecks. Nuclear whorls 1, glossy, opaque- cream to tan and slightly mammillate. First post- nuclear whorl with fine, axial riblets which are soon crossed by 2, then 3, fine spiral threads. Spire flattish, its angle about 100°, and each whorl with 8 to 10 small, irregular, spiral threads. Suture wavy and marked by broken, former shoulder spines. Shoulder of body whorl carinate and with 12 to 15 short, sharp, flattened spines which are open on their anterior faces. Below each spine, a low axial fold runs to the base of the whorl and is crossed by small, distinct, raised, wavy cords. Aperture oval-round, white. Parietal shield raised, white; base of columella bears 3 well-developed plicae which are set at almost right angle to the axis of the shell. Interior of outer wall with 9 to 11 sharp, fine spiral cords running back into the aperture. Siphonal canal about 2* the length of the entire shell. Periostracum very thin, light-tan, and some- what deciduous. Measurements (mm.) off Darwin ) — length width 50.0 22.5 42.5 19.5 41.2 19.0 Synonymy — 1863 Tudicla (Tudicula) spinosa H. and A Adams, Proc. Zool. Soc. London for 1863, p. 429 (Port Curtis). 1884 Turbinella (Tudicula) spinosa H. and A. Adams, E. A. Smith, Report Zool. Collections H. M. S. “Alert”, London, p. 54, pi. 5, fig. H. 1932 Tudicla inermis Angas, Kuroda, Venus, vol. 3, p. 117, fig. 3 on p. 114 (northern Australia). Types— The type locality is Port Curtis, Queens- land, Australia. The type in the British Museum (Natural History) is 38 mm. in length and entirely white in color. Locality records— QUEENSLAND: Port Curtis (Hugh Cuming); Prince of Wales Channel, Torres Straits in 9 fms. (E. A. Smith, 1884, p. 54). NORTHERN TERRITORY: off Darwin (ex A. R. Cahn, ANSP). Tudicula (Tudicula) inermis Angas, 1878 (PI. 9, figs. 3, 4, p. [20-444]) Range— Western Australia and Northern Terri- tory, Australia. Remarks— This ra^e species differs from spinosa in lacking the small spines on the shoulder of the body whorl. Most specimens we have seen are rather heavily pigmented with light-brown. We have two specimens from off Darwin which have undulations at the shoulder and weak axial folds on the sides of the body whorl, characters which are approaching those found in spinosa. T. inermis may subsequently be found to be a western sub- species of spinosa. The radula figured is from a specimen loaned by the Australian Museum (no. C. 57363) from Exmouth Gulf. See pi. 8, p. [20-443]. Habitat— Unknown, except for the fact that it occurs at a depth of 5 to 11 fathoms. Description— Adult shell 40 to 47 mm. ( 1& inches) in length, turnip-shaped, smoothish, with a rela- tively long siphonal canal; color cream to whitish with light-brown mottlings and flecks. Nuclear whorls 1, glossy, opaque-tan to pinkish brown and slightly mammillate. First two postnuclear whorls with 3 or 4 slightly nodulated spiral threads. Spire flattish, its angle about 100°, and with 8 to 13 irreg- ularly-sized spiral threads. Shoulder of body whorl slightly carinate and with or without weak undula- tions. In some specimens there may be a very weak axial fold below each of the undulations. Base of last whorl with numerous, fine spiral threads. Aper- ture oval-round, white. Parietal shield raised, white; the base of the columella bears 3 well- developed plicae which are set almost at right angle to the axis of the shell. Posterior end of the parietal wall with a small, swollen, white callus. Interior of outer wall with 9 to 11 sharp, fine spiral cords running back into the aperture. Siphonal canal about 21 the length of the entire shell, with brownish flecks, rarely with pink. Periostracum thin, grayish, axially fimbriated and deciduous. Operculum corneous, translucent yellowish brown, unguiculate, with fine growth lines and with a ter- minal nucleus. The head, foot and tentacles are very similar to those in Vasum. In a shell 44 mm. in length, the odontophore is 5.0 mm. in length and bears about 103 rows of teeth. Measurements (mm.) length width no. whorls 41.0 21.0 5.5 ( holotype ) 44.5 20.0 5.6 ( off Broome ) 46.5 21.5 6.0 ( Exmouth Gulf ) Synonymy — I STS Tudicula inermis Angas, Proc. Zool. Soc. London for 1878, p. 610, 2 figs, (from a dealer at Singapore); 1887, E. A. Smith, Annals and Mag. Nat. Hist., series 5, vol. 19, p. 465 (Exmouth Gulf). Types— The type locality given by Angas was Singapore. This is evidently erroneous. The holo- type is in the Paleontological Museum, Cornell University, Ithaca, New York, no. 19232, from the Newcomb collection. Locality records — WESTERN AUSTRALIA: Exmouth Gulf, T. H. Hayes (B.M. and ANSP); between Cape Bossut and Broome, 5 fms., A. A. Livingston (Aust. Mus. ); Pearl bank, 42 mi. W.S.W. of Cape Jaubert in 42 to 66 ft. ( Odhner, 1919). NORTHERN TERRITORY: off Darwin, T. Ino (ANSP). (including spines; all from length of siphon no. whorls 31.5 5 26.0 5 24.3 5 [20 - 446] March 31, 1959 INDO-PACIFIC MOLLUSCA, vol. 1, no. 1 Tudicla 31 Tudicula (Tudicula) zanzibarica Abbott, 1958 (PI. 9, figs. 7, 8, p. [20-444]) (PI. 8, figs. 1-3, p. [20-443]) Range— Zanzibar, British East Africa. Remarks— The Zanzibar Tudicula is characterized by its elongate, apricote-colored aperture and the 7 to 8 delicate, somewhat triangular, purple-brown spines on the last whorl. The close relationship be- tween Tudicula and Vasum is demonstrated in the characters of this species in which the aperture and nucleus are Va^nm-like, but the general shape of the shell, the spines and the radula are more T udicula- like. Habitat— Dredged in 8 fathoms on a bottom of sand, broken shell and wiry grass. Description— Adult shell 28 to 40 mm. ( 1 to 1/2 inches) in length, subtriangular, solid, and spinose. Color chalky-white with a flush of light-orange, and with dark purple-brown on the ends of the shoulder spines and the siphonal canal. Spire moderately elevated, its angle about 90°, its length about K that of the entire shell. Whorls 6, strongly shoul- dered near the top where there is a peripheral row of 7 to 8 large, flattened, somewhat triangular, purple-brown spines which may be narrowly ex- tended. Sides of whorls flat to slightly concave. Nuclear whorls 1, rather large, bulbous, elevated, smooth and yellowish white. First postnuclear whorl rudely cancellated by 4 coarse, unequal, spiral cords and about 8 axial, poorly-developed ridges. Lower third of the last whorl, in the central region of the siphonal canal, bears 2 spiral rows of about 6 elongate spines which may become obso- lete in the last whorl. Microsculpture consists of numerous, small, spiral cords of varying size and of numerous, very fine, axial threads and fimbria- tions of light-tan periostracum. Aperture long, wide above, gradually becoming constricted below. Outer lip thickened, glossy, weakly crenulate, and pale peach in color. Parietal shield well-developed and raised. Columella bears 3 weak, whitish, spiral plicae. Interior wall of last whorl with about 9 very weak, small, spiral ridges. Operculum corneous, rather thick, unguiculate, brown, and with its muscle scar being about M the area of the entire operculum. Animal’s soft parts similar to those of Vasum, with a short, squarish foot; head small, bearing two, short stubby tentacles with the eyes near the ends. Gills with about 400 lamellae. Osphradium well-developed, bi-laminate and about half the length of the gills. Color of preserved soft parts cream with maculations of purple-brown. Odonto- phore long (6 mm. in the 40 mm. -long shell of the holotype), narrow, and with about 145 transverse rows of rachiglossate teeth (see pi. 8, figs. 1-3). Measurements (mm.) (not including spines) — length width no. whorls 40.0 25.2 6.0 ( holotype ) 36.5 20.5 5.5 ( paratype ) 28.0 15.0 5.0 ( paratype ) Synonymy — 1958 Tudicula zanzibarica Abbott, Notulae Naturae (Phil- adelphia), no. 305, pp. 1-4, figs. 1-7 (Zanzibar). Types— The type locality is 1/2 miles W.S.W. of Ras Nungwi, north end of Zanzibar Island. Natural Science Foundation station no. 651, March 4, 1957 (Ostheimer and V. Orr). Holotype in ANSP no. 225261; paratypes in the Coryndon Mem. Mus., B.M., and ANSP. Locality records — Known only from the type locality. Genus Tudicla Rbding 1798 This genus is apparently limited in Recent times to the Bay of Bengal in the Indian Ocean where it is represented by a single species, Tudicla spirillus (Linne). The anatomy is unknown, so that the genus is placed, at present, in the Vasidae solely on conchological grounds. The genus has had several fossil subgenera and six or seven Recent species assigned to it by several authors, but we believe that none of these properly belongs to this genus. Among the possibly related fossil genera are: Tudi- clana Finlay and Marwick 1937 (Cretaceous, New Zealand), Pyropsis Conrad 1860 (probably a Buc- cinidae from the Upper Cretaceous of southeast United States), and others listed by W. Wenz, 1943, vol. 6, p. 1304. The genus Afer Conrad 1858, represented by two Recent West African species ( afer Gmelin and porphyrostoma Adams and Reeve) has been tradi- tionally placed in the Vasidae next to Tudicla, but the shells appear to be more like those of Latirus in the family Fasciolariidae. Its anatomy is un- known. True Tudicla appears first in the Cretaceous of East Africa ( krenkeli Cox, 1925) and Madagascar [20-471] 32 Tudicla R. T. Abbott Vasidae ( hourcqi Collignon, 1951), then the Eocene of Egypt ( rames and thebaica Cuvillier, 1933), the Miocene of Europe ( rusticulus Basterot, 1825), the Pliocene of Australia ( angulata , costata and tur- binata Angas, 1888), and the Pliocene of Karikal, India ( spirillus Linne). See also Cossmann, 1901, Essais de Paleoconch. Comp., pt. 4, pp. 68-72. We have not seen specimens of “Fusns” couderti Petit 1853 or Tudicla fusoides A. Adams 1854, both from China, but they do not appear to belong to true Tudicla as Tryon (1881) and Kiister (1876) believed. Other Recent species, such as armigera A. Adams and spinosa H. and A. Adams, were once assigned to Tudicla, but are actually members of the genus Tudicula and much more closely related to the genus Vasum. Fischer in 1884 (Manual de Conch., p. 619) in- cluded Spirillus Sowerby 1842 as a generic synonym of Tudicla. However, Sowerby (Conch. Manual, 2nd ed., p. 306) did not propose this as a new generic name, but was merely capitalizing Linne’s specific name of spirillus. Description— Shell medium in size, moderately solid, pyriform, with a swollen, slightly bicarinate body whorl, flattish spire, and a long, narrow siphonal canal. Without an umbilicus. Parietal shield raised, with a sharp edge, and glazed. Colu- mella roundly arched and bearing at its base a single, raised spiral cord. Region of the posterior canal with a swollen, button-like callus. Nucleus mammillate, high and with 1M round, swollen, glossy whorls. Operculum corneous, narrowly oval with an apical nucleus, according to H. Adams, 1874. Synonymy — 1798 Tudicla Roding, Museum Boltenianum, Hamburg, pt. 2, p. 145. Type (by subsequent designation, P. Fischer, 1884, p. 619): T. carinata Roding = Murex spirillus Linne, 1767. 1835 Pyrella Swainson, Elements of Conchology, London, p. 21. Type (by monotypy): P. spirilla L. = Murex spirillus Linne, 1767. 1838 Spirillus Sehliiter, Kurzgefasstes Verzeichniss Conehyb, Halle, p. 21. Type (by monotypy): S. rostratus Sehliiter = Murex spirillus Linne, 1767. 1857 Pyrenella J. E. Gray, Guide Syst. Distrib. Mollusca Brit. Mus., pt. 1, p. 11. Type (by monotypy): P. spirilla = Murex spirillus Linne, 1767. Tudicla (Tudicla) spirillus (Linne, 1767) (PI. 9, figs. 1, 2, p. [20-444]) Range— Southeast India and northern Ceylon. Remarks— This is the only known living species in this genus, and it is easily recognized by its turnip shape, bicarinate body whorl, the single fold on the lower part of the columella, and by its mammillate nuclear whorls. Habitat— Unknown, except that it probably lives on a sandy bottom in several fathoms of water. Description— Adult shell 66 to 82 mm. ( 23a to 3)1 inches) in length, moderately solid but strong, pyriform, and with a long, sinuous, smooth siphonal canal. Color of shell shiny cream to pinkish gray with sparse flecks of light-brown which form small squarish spots on the upper carina of the body whorl. Nuclear whorls mammillate, projecting, and of 1/2 round, swollen, pinkish or yellowish, glossy whorls. Spire flattish, its angle about 140°. Post- nuclear whorls 3)2. Spiral sculpture of numerous, raised threads which are squarish and broad on the base of the whorls. Body whorl with a squarish periphery, bounded above by a sharp, wavy keel and below by a series of 6 to 8 round, low nodules. Aperture ovate-round, glossy, pinkish to purple within, and bounded on the outer side by a sharp lip. Inside of outer wall with numerous, fine, raised, spiral ridges. Parietal shield well-developed, smooth and glossy white. Posterior canal region with a swollen, button-like, white callus. Base of columella with a single, strong, spiral plica. Siphonal canal open along its length, long and slightly sinuous. Operculum corneous, elongate- oval, and with an apical nucleus. Radula unknown. Measurements — length width 66.0 36.5 ( small; Ceylon ) 70.5 41.0 (average; Ceylon) 78.0 43.0 (large; Ceylon) Synonymy — 1767 Murex spirillus Linne, Sysyema Naturae, ed. 12, p. 1221, no. 554 (in Tranquebar); 1957, Dodge, Bull. Amer. Mus. Nat. Hist., vol. 113, art. 2, p. 156. 1798 Tudicla carinata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 145 ( refers to Knoor, vol. 6, pi. 24, fig. 3 ) . No locality. 1811 Monoplex capitatus Perry, Conchology, London, pi. 3, fig. 4 (locality unknown). 1838 Spirillus rostratus Sehliiter, Kurzgefasstes Verzeichniss Conchyl., Halle, p. 21 (refers to ‘Pyrula spirillus Lam.”). Types— The type locality given by Linne was Tranquebar, a section of the southeast coast of India. The type is in the British Museum (Natural History), London. Locality records— Known only from the Tranque- bar coast in southeast India and the northern part of Ceylon. Gravely, F. H. (1942, p. 66) reports that this species is occasionally washed ashore on the beaches near Madras, southeast India. [20 - 472] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 33 5^4- as ojQ >-•' 1 M I- I / THE GENUS STROMBUS IN THE INDO-PACIFIC by R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia The genus Strombus is a world-wide, tropical group of mesogastropods belonging, with other Re- cent genera such as Lambis, Terebellum, Tibia, and Rimella, to the family Strombidae. The genus originated during the early Miocene or possibly the Eocene, and became established throughout the warm seas of the world. It flourished during the Pliocene and the early Pleistocene in the area of southeast Asia, but lias since had a gradual diminu- tion in species, until none now survive in the Medi- terranean, only one in West Africa, seven in the Caribbean region and four in the Panamic region on the Pacific side of Central America. Sixty living races occur in the Indo-Pacific of which thirty eight may be considered to be full species. Members of the genus Strombus are for the most part shallow water inhabitants. In all known cases they are herbivorous or detritus-feeders. The vast majority live from the low tide line to a depth of about 20 feet, although about one fourth of the spe- cies occur more commonly from 15 to 30 feet, and two at depths from 6 to 66 fathoms. Few S trombus have been found in water too cold to support the production of coral reefs, and the vast majority of species are found within the limits of the 70° F re- stricted isotherm (water always above 70 F). S trombus are outstanding for their agility of lo- comotion. The foot is narrow, very muscular, and with the flat, creeping sole limited to the anterior third of the foot. The operculum is long, sharp, strong and somewhat sickle-shaped, serving not only as a lever in moving the animal forward, but also as a defensive weapon against predatory crabs and fish. The eye peduncles are long and muscu- lar, and near the distal end give rise to a small, short tentacle. The eyes are large and usually adorned with circular rings of orange, yellow or red color. The animals are dioecious with the adult males bearing a long, open-grooved prong-like penis on the right side of the “back”. The animals generally congregate in large colonies in shallower water dur- ing the warmer months of the year to spawn. The egg masses consist of a long, jelly-like tube to which sand grains become attached. The coiling tube be- comes entwined into a sponge-like mass. The length of unravelled tube may vary from 43 to 74 feet and contain an estimated 185,000 to 460,000 in- dividual eggs. The eggs develop into free-swim- ming veligers within 80 to 100 hours after first be- ing laid. An account of the spawn and veligers of three Western Atlantic Strombus and an excellent bibliography is given by R. Robertson ( 1959, Proc. Mai. Soc. London, vol. 33, pt. 4, pp. 164-171, 1 pi. ). A slight degree of sexual dimorphism is evident in the shells of most Strombus, those of males usu- ally being slightly smaller (see remarks under S. gibberulus gibbosus). The shells of Strombus are characterized by the development in the adult of a large, flaring, gener- ally thickened, outer lip and the presence of a U- shaped notch on the edge of the lip near the anterior end. This feature is referred to as the “stromboid notch,” and it generally serves as a "peep hole” for the protruding right eye. It is prob- ably the convenience of this notch that has led to a reduction in length of the right eye peduncle, so common a feature in most Strombus. Another char- Plate 11. Living Strombus pipus (Roding) emerging from its shell. From left to right: the brown, sickle-shaped oper- culum, underside of foot, yellowish brown proboscis and brownish mauve-spotted eye peduncles. Zanzibar specimen natural size. Photo by Virginia Orr. 34 Strombus R. T. Abbott Strombidae acteristic is the production of slightly thickened, rounded varices in the postnuclear whorls of the spire. These adolescent varices may serve to strengthen the rather fragile young shell. Some spe- cies produce wide adolescent varices, such as those found in the subgenus Gibbenilus, while others produce narrow ones, or on rare occasions none at all. In shell size, adults vary from the 13-inch Strombus (Tricornis) goliath Schroter of Brazil to a dwarf S. (Canarium) maculatus Sowerby only % inch ( 8 mm. ) in length. Strombus quite commonly differ in shell size from colony to colony due probably to a combination of genetic and ecologic conditions. Examples are dis- cussed under the remarks on S. canarium and S. gibbenilus gibbosus. Some unjustified races have been based upon these local dwarfs or giants. Old adults commonly thicken the shell, especially the outer lip, and may lay down an aluminum-like glaze, features which also have been the basis for unjustified subspecies. The internal anatomy of several Indo-Paeific spe- cies has been treated in detail by B. Haller ( 1893, Morphologisches Jahrbuch, Leipzig, vol. 19, pp. 577-588, pis. 18-20 ) and by R. Bergh ( 1895, Zoolo- gische Jahrbiicher, Anat. und Ontog., Jena, vol. 8, pp. 342-378, pis. 22-23). Subgenera of Strombus There is always difficulty in defining limits for genera and subgenera that will be satisfactory to all workers. Strombus presents no exception. Our evaluation is based upon shell, radular, penial, and opercular characters, as well as the fossil history. Among the shell characters employed are sculpture of apical whorls, production of axial and spiral sculpture in later whorls, development of the outer lip, general shape and texture of shell, the colora- tion and sculpturing of the columella and of the inside of the outer body wall, and the development of the siphonal canal. Unfortunately, the expres- sion of these characters arises in varying combina- tions throughout obviously different stocks, so that some shell characters become misleading. The modifications of the penis have served usefully in some cases, some being simple prongs (as in the subgenus Canarium) , others being strongly bilobed (as in the subgenera Dolomena, Euprotomus, and Lentigo ) . The radula throughout the genus Strombus shows little diversity, and, again, sufficient inter- and intra-specific variations occur to reduce this charac- ter to secondary importance. The lateral cusps of Plate 12. Living animal of Strombus canarium Linne. 1, the “stromboid notch”. 2, anterior siphonal canal. 3, eye peduncle. 4, tentacle. 5, proboscis and mouth. 6, anterior end of foot. 7, operculum. 8, posterior canal. Plate 13. Portion of egg masses of Strombus with some sand grains removed to show internal coils of eggs. Fig. 1, S. ra- ninus Gmelin, Bahamas. Fig. 2, S. costatus Gmelin, Ba- hamas. Both about x 40. (From R. Robertson, 1959, p. 167). some species have made possible the rather con- venient acceptance of the subgenera Strombus sensu stricto and Doxander. In other groups, such as Gibberulus, the absence of the tiny, but distinct peg at the inner base of the lateral has been help- ful. Curiously, some of the radular characteristics are correlated with the marine provinces, rather than with assumed phylogenetic relationships. [09 - 832] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 35 Plate 14 Figs. 1, 2 3,4 5, 6 7, 8 9, 10 11, 12 13 14 15 16, 17 18, 19 Strombus uurisdianae aratrum (Roding). Cairns. Strombus aurisdianae Linne. Balabac Id. Strombus bulla (Roding). Cebu Id. Strombus vomer vomer (Roding). New Caledonia. Strombus vomer hawaiensis Pilsbry. Hawaii. Strombus vomer iredalei Abbott. Western Australia. Strombus decorus decorus ( Roding). Mozambique. S trombus decorus persicus Swainson. Arabia. Strombus luhuanus Linne. Dutch New Guinea. Strombus fasciatus Born. Red Sea. Strombus dilatatus swainsoni Reeve. Pacific. 20 Strombus variabilis Swainson (color form). Palau. 21, 22 Strombus variabilis Swainson. Balabac Id. 23 S trombus dentatus Linne. Okinawa Id. 24 Strombus dilatatus form orosminus Duclos. Cebu. 25 Strombus dilatatus dilatatus Swainson. Mindanao. 26 Strombus gibberulus gibbosus (Roding). Cebu Id. 27 Strombus gibberulus albus Morch. Red Sea. 28 Strombus gibberulus gibberulus Linne. Zanzibar. 29 S trombus terebellatus Sowerby. Pacific Ocean. 30 Strombus fragilis (Roding). Western Pacific Ocean. ( all approximately % natural size ) [09 - 833] 36 Si trombus R. T. Abbott Strombidae Plate 15. Egg mass (fig. a), egg strand (fig. b), and en- larged section showing single row of eggs ( fig. c ) of S trom- bus macula tits Sowerby. (from Ostergaard, 1950, p. 95, fig. 17). Plate 16. Two of forty transverse rows of radular teeth of Strombus gibberulus gibbosus (Roding) from a New Cale- donian specimen, x 70. Little is known about the species characters in the egg masses and veligers, but preliminary ob- servations suggest that these may be additionally useful. Our subgenera are not of equal rank, as would be expected when attempting to put arbitrary limits on nature's gradual and many-faceted avenues of evolution. The subgenera Doxancler, Labiostrombus (in our restricted sense), Gibberulus and Cono- murex seem to be well defined and sufficiently separated from their morphological neighbors. Within Canaria in and Tricornis are several groups of species which are obviously closely related, but other species have a mixture of these group charac- ters. We have refrained from creating a new series of monotypic subgenera or genera, feeling that this would obscure the relationships. Over-zealous ne- ontologists would do well to consult the fossil rec- ord, and paleontologists should not be tempted to recognize some higher categories simply on the basis of long stretches of time. Below we are listing the accepted subgenera and species of Strombus. Those in [brackets] are from other oceans, and those preceded by a dagger f are fossil. Abbreviations: W.A. is Western Atlantic; E.P. is Eastern Tropical Pacific or Panamic region. The list contains 89 fossil and living Indo-Pacific species and subspecies, of which 60 are living. We recog- nize 38 full species for the Recent Indo-Pacific and a world total of 50 Recent full species. List of Recognized Taxa Genus Strombus Linne, 1758 Strombus s.s. Linne, 1758 [pugilis Linne, 1758] Type W.A. [alatus Gmelin, 1791] W.A. [gracilior Sowerby, 1825] E.P. Laevistrombus Kira, 1955 canarium Linne, 1758. Type f varinginensis Martin, 1899 subsp. f martini Oostingh, 1935 f overbecki Cox, 1948 f karikalensis Cossmann, 1903 f glaber Martin, 1879 Tricornis Jousseaume, 1886 tricornis Humphrey, 1786. Type thersites Swainson, 1823 \maximus Martin, 1883 latissimus Linne, 1758 taurus Reeve, 1857 sinuatus Humphrey, 1786 jjunghuhni Martin, 1879 f mekranicus Vredenburg, 1928 \ infiat us Martin, 1879 f tjilonganensis Martin, 1899 [raninus Gmelin, 1791] W.A. [peruvianas Swainson, 1823] E.P. [gallus Linne, 1758] W.A. [costatus Gmelin, 1791] W.A. [galeatus Swainson, 1823] E.P. [goliath Schroter, 1805] Brazil [gigas Linne, 1758] W.A. Dilatilabrum Cossmann, 1904 [f fortisi Brongniart, 1823] Type. Eocene Canarium Schumacher, 1817 urceus Linne, 1758. Type subsp. orrae Abbott, 1960 labiatus (Roding, 1798) subsp. olydius Duclos, 1844 subsp. f gendinganensis Martin, 1879 klineorum Abbott, 1960 microurceus (Kira, 1959) mutabilis Swainson, 1821 subsp. ochroglottis Abbott, 1960 subsp. f ostergaardi Pilsbry, 1921 [09 - 834] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 37 maculatus Sowerby, 1842 \unifasciatus Martin, 1884 f spolongensis Martin, 1916 fusiformis Sowerby, 1842 erythrinus Dillwyn, 1817 subsp. rugosus Sowerby, 1825 haemastoma Sowerby, 1842 scalariforrnis Dnclos, 1833 helli Kiener, 1843 dentatus Linne, 1758 fragilis (Roding, 1798) terebellatus Sowerby, 1842 subsp. afrobellatus Abbott, I960 Dolomena Iredale, 1931 plicatus (Roding, 1798) subsp. columba Lamarck, 1822 subsp. sibbaldi Sowerby, 1842 subsp. pulchellus Reeve, 1851. Type f palabuanensis Martin, 1899 f deperditus J. de C. Sowerby, 1839 dilatatus Swainson, 1821 subsp. swainsoni Reeve, 1850 subsp. \taiwanicns Nomura, 1935 subsp. \fennemai Martin, 1899 f rembangensis Martin, 1899 labiosus Wood, 1828 subsp. \tcschi Cox, 1948 f rutteni Altena, 1942 marginatus Linne, 1758 subsp. succinctus Linne, 1767 subsp. robust its Sowerby, 1874 subsp. Septimus Duclos, 1844 f togopiensis Cox, 1948 f sedanensis Martin, 1899 f javanus Martin, 1879 variabilis Swainson, 1820 subsp. athenius Duclos, 1844 minimus Linne, 1771 Labiostrombus Oostingh, 1925 epidrornis Linne, 1758. Type f denti Cox, 1948 1 kerned jingensis Martin, 1916 Doxander Iredale, 1931 vittatus Linne, 1758. Type subsp. japonicus Reeve, 1851 subsp. \madiunensis Martin, 1899 subsp. f deningeri P. J. Fischer, 1921 subsp. campbelli Griffith and Pidgeon, 1834 f triangulatus Martin, 1879 listen T. Gray, 1852 Lentigo Jousseaume, 1886 lentiginosus Linne, 1758. Type pipus (Roding, 1798) fasciatus Born, 1778 [lotus Gmelin, 1791] West Africa f preoccupatus Finlay, 1927 [granulatus Swainson, 1822] E.P. Euprotomus Gill, 1870 aurisdianae Linne, 1758. Type subsp. aratrum (Roding, 1798) bulla (Roding, 1798) vomer (Roding, 1798) subsp. hawaiensis Pilsbry, 1917 subsp. iredalei Abbott, 1960 Conomurex P. Fischer, 1884 luhuanus Linne, 1758. Type decorus ( Roding, 1798 ) subsp. persicus Swainson, 1821 Gibberulus Jousseaume, 1888 gibberulus Linne, 1758. Type subsp. gibbosus (Roding, 1798) subsp. albus Morch, 1850 f Oostrombus Sacco, 1893 f problematicus Michelotti, 1861. Type. Eocene Distribution of Strombus A comparison of the distribution and speciation of the genus Strombus with that of another meso- gastropod group is possible in the case of the Cypraeidae because of the recent monograph by Sehilder and Schilder (1938, Proc. Mai. Soc. Lon- don, vol. 23, pp. 119-231). Allowances should be made for the excessive generic and racial splitting by those authors. Many of their races are not accepted today, nor do some of their marine sub- regions have much to support their continued recognition ( Sulu Sea region, Sumatran region, Bermudian region). Nevertheless, some interesting comparisons are possible. The living, world-wide Strombidae ( Strombus, Lambis, Terebcllum, Tibia, and Rimella) contains about 75 full species (if strong subspecies are added, 100). The genus Strombus contains 50 full species. There are about 165 full species in the genus Cypraea (Cypraeidae of Schilder and Schil- der, 1938, but see Kay, 1960). This ratio of about one Strombus species to 3 Cypraea species does not maintain itself throughout the various parts of the world tropical seas. One obvious reason for a higher proportion of Cypraea in such places as South Aus- tralia, South Africa, and the Mediterranean is that the Strombidae are more tropical in habitat prefer- ence. About 20 species of Cypraea live in waters presumably too cold for Strombus. Compared to the world ratio, there are proportionately more Strombus than Cypraea in the Panamic and Carib- bean regions. The slightly cooled waters of the [09 - 835] 38 S trombus R. T. Abbott S trombidae Plate 17 Figs. 1, 2 Strombus thersites Swainson. Okinawa Id. 3,4 Strombus taurus Reeve. Rongelap Atoll. 5 Strombus latissimus Linne. Mindanao Id. 6, 7 Strombus tricornis Humphrey. Red Sea. 8 Strombus sinuatus Humphrey. Okinawa Id. 9, 10 Strombus pipus (Roding). Mindanao Id. 11, 12 Strombus lentiginosus Linne. Sehouten Ids. 13 Strombus vittatus campbelli Griff, and Pidg. Queensland. 14 Strombus vittatus vittatus L. Cebu Id. (left, form australis Schroter; right, typical). 15 Strombus canarium Linne (typical form). New Caledonia. 16 Strombus canarium L. (form turturellus Roding) ). Samar Id. 17 Strombus epidromis Linne. Mindanao Id. 18 Strombus vittatus japonicus Reeve. Ise, Japan. ( all approximately b natural size ) [09 - 836] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 39 Plate. 18 Figs. 1,2 Strombus plicatus columba Lamarck. Zanzibar. 3 Strombus plicatus pulchellus Reeve. Luzon Id. 4, 5 Strombus minimus Linne. Cebu Id. 6,7 Strombus marginatus marginutus Linne. Bay of Bengal. 8, 9 Strombus marginatus robustus Sowerby. Japan. 10, 11 Strombus marginatus Septimus Duclos. (left, Solo- mons; right, Luzon Id. ) 12 Strombus plicatus plicatus (Roding). Gulf of Suez. 13, 14 Strombus marginatus succinctus Linne. Ceylon. 15, 16 Strombus plicatus form sibbaldi Sowerby. 17, 18, 19 S trombus labiosus Wood. Philippines. ( all natural size ) [09 - 837] 40 Strombus R. T. Abbott Strombiclae Mediterranean and West Africa support 10 Cypraea but only one Strombus in West Africa. On the other hand, the Indo-Pacific is proportionately richer in Cypraea than in Strombus. A closer comparison of the distribution within the Indo-Pacific permits several generalizations. The Cypraea are not only more numerous in species for each small geographic area, but also are more evenly distributed. The rich areas (the Western Pacific Arc, the Bay of Bengal and central East Africa) are the same for each genus, but much more pronounced for Strombus. The areas of high endemicity (i.e. Red Sea, Bay of Bengal, northern Australia, Hawaii and Polynesia ) are about the same for each genus. The reduction in the number of species eastward from the Western Pacific Arc into Micronesia and Polynesia is very pronounced in Strombus, much less so in Cypraea. Both genera have but one species in Easter Island. There are no close relatives of the Indo-Pacific S trombus in the Panamic province, but one Cypraea (Isabella [or controversa] subspecies mexicana Stearns, 1893) has made the migration. The Western Pacific Are Our patterns of Indo-Pacific distribution of Strom- bus species are similar to those outlined for Cypraea by Schilder and Schilder (1938), with one major difference, in that those authors have failed to recognize the importance and nature of the West- ern Pacific Arc. This is a zoogeographic area run- ning from the northern Ryukyn Islands south through Taiwan, the Philippines, Indonesia and southeastward through New Guinea, to New Cale- donia and Fiji. The center of species occurrence is in the central and southern Philippines where 26 races of Strombus are present. The Western Pacific Arc has a representation of 46 per cent of the Indo- Pacific and 72 per cent of the Pacific races of Strom- bus ( in contrast to 10 and 15 per cent respectively in the case of the Hawaiian fauna, and 11 and 18 per cent respectively in the case of the Society Is- lands ) . The following species or races of Strombus are endemic to the Western Pacific Arc: * Plate 19. Number of species and races of Strombus in each area of the Indo-Pacific region. Insufficiently collected areas have been omitted, and some of the present censuses will increase by about 10 percent when additional records are found. Total races: 60; Indian Ocean 22; Pacific Ocean; 39. [09 - 838] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Stromb us 41 (Tricornis) latissimus (Dolomena) minimus (Dolomena) dilatatus (Labiostrombus) epidromis (Dolomena) septimus (Euprotomus) bulla (Euprotomus) vomer s.s. Other species of Strombus occurring mainly in the Western Pacific Arc have advanced (or have not retrenched their former more widespread dis- tributions) in several directions. Fifteen species maintain themselves in Queensland, Australia. Some of these may represent Pliocene or even Mio- cene invasions from the north, but others may be Quaternary migrations along westward flowing cur- rents from the New Caledonia and Brampton Reef areas. The color patterns of Strombus canarium support the latter possibility for some species. Only three species of the Western Pacific Arc oc- cur to the west. Two of these, canarium and labiatus, are present in the Bay of Bengal and are also capable of sustaining themselves in the slightly cooler waters of Honshu Island, Japan. These spe- cies are always associated with large islands or con- tinental shores where nitrogenous-rich waters pre- vail. The third species, aurisdianae, which occurs abundantly throughout the middle of the Indian Ocean from East Africa to Sumatra, may be a case of a recent advancement from Africa into the West- ern Pacific Arc as far as Okinawa and the Solomons. It occurs relatively abundantly in African fossil beds, but not in Indonesian beds. Furthermore, it is the only species in the Western Pacific Arc which stops at the Solomons. Curiously, the two advanc- ing “horns”, one in northern Queensland, the other in the Ryukyus have specimens which show a tend- ency towards melanism. The Queensland popula- tions are so drastic in this and other characters, that we recognize them as the subspecies aratrum. The Indian Ocean populations show great instability in shell characters, and we believe represent potential isolation and a development of an Indian Ocean species or subspecies for the geologic future. A more advanced case of the aurisdianae- like spread, is seen in Strombus luhuanus. Its fossil rec- ord is also better in Africa, and, in the Recent In- dian Ocean, is represented by the distinct species decorus. The latter’s instability has resulted in a northwestern Indian Ocean subspecies, persicus. S. luhuanus has spread beyond the Western Pacific- Arc as far north as Honshu Island, Japan, as far south as Sydney, Australia, and as far east as Pal- myra Island. All three extremities are noted for shells which tend towards either a stunted condi- tion or a reduction of the typical, bright water- melon red in the aperture. If the central Western Pacific Arc populations were to become extinct, there might well develop a situation similar to that which occurs in the peripheral distribution of S. vomer and its subspecies (see below). Other Western Pacific Arc species which exist to the eastward in varying distances are: fragilis, gib- berulus gibbosus , microurceus, pipus, sinuatus, tere- bellatus terebellatus, and variabilis. We believe that pipits is in a period of shrinkage, since it occurs in small numbers in an isolated pocket in Tahiti, but mainly extends only from the Ryukyus to the Solo- mons. S. pipus is common only in the Sulu Sea. On the other hand, gibberulus gibbosus, a Pacific sub- species, is maintaining itself as an abundant species throughout the vast central Pacific, and has died out only in the Hawaiian Chain since the Pliocene. The stunted, smaller, less colorful specimens living in the atolls of the central Pacific suggest that gib- berulus gibbosus is destined to extinction in the geologic future. Two striking cases of discontinuous distribution exist in the Western Pacific Arc. One of these ap- pears to be of relatively recent origin, the other of more ancient origin. The large, colorful and hand- some Strombus tliersites has been found on numer- ous occasions in the waters of New Caledonia and Okinawa, Ryukyu Islands. Despite many years of collecting in the intervening areas where many rare shells, such as Conus gloriamaris, Vasum tubiferum, Cypraea guttata, etc., have been found, this large shell has not turned up in the Philippines, Indo- nesia or New Guinea. It is fairly safe to say that it does not occur, or at least is extremely rare, in these intervening areas. This is also true of S. vomer vomer which is moderately common in the Ryukyus and New Caledonia. Both of these species, or very close relatives of them, occur in the Pliocene of In- donesia, Taiwan, and the Lau Islands of Fiji. It ap- pears that both of these species were widely spread throughout the Western Pacific in previous geo- logic times. Each, today, has been reduced in dis- tribution. In the case of vomer, three recent isolated ge- netic pools appear to have survived from the once widespread Pliocene range. Probably the oldest of these is the living subspecies iredalei which is limited to the Dampierian province from Western Australia to the Gulf of Carpentaria in northwest Queensland. A second subspecies, hawaiensis, is now limited to the Hawaiian Chain from Midway to Hawaii. It is closer morphologically to the West- ern Pacific Arc vomer vomer and the Pliocene Tai- wan fossil than to iredalei. The third subspecies is [09 - 839] 42 Strombus R. T. Abbott Strombidae Plate 20 Figs. 1, 2 Strombus erythrinus elegans Sowerby. Noumea. 3 Strombus enjthrinus erythrinus Dillwyn. Red Sea. 4 Strombus erythrinus (dwarf form). New Guinea. 5 Strombus erythrinus erythrinus Dillwyn. Mauritius. 6 Strombus haemastoma Sowerby. Wotho Atoll. 7, 8 Strombus belli Kiener. Oahu Id., Hawaii. 9, 10 Strombus mutabilis ochroglottis Abbott. Mauritius. 11, 12 Strombus erythrinus rugosus Sowerby. Polynesia. 13, 14 Strombus maculatus Sowerby. Hawaiian Ids. 15, 16 Strombus mutabilis Swainson. Cebu Id. 17 S trombus mutabilis form zebriolatus A. and L. 18, 19 Strombus labiatus oltydius Duclos. Indian Ocean. 20, 21 Strombus klineorum Abbott. Ceylon. Holotype. 22 Strombus labiatus (Roding) (smooth form). Cebu. 23 Strombus labiatus (Roding) ( ribbed form ) . Samar. 24,25 Strombus microurceus (Kira). Okinawa. 26, 27 Strombus urceus Linne. Cebu Id. 28 Strombus urceus orrae Abbott. Western Australia. 29 Strombus urceus form ustulatus Sclium. China. 30 Strombus fusiformis Sowerby. Zanzibar. [09 - 840] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 43 the typical vomer vomer which is now, in Recent times, beginning to die out and separate into two remotely isolated populations. Given time, they too may become subspecifically separated. In the above two cases and in S. pipits, we see ac- tual or potential examples of differentiation occur- ring at the extremities of a range after extinction has occurred in the center of distribution. There are, however, suggestive examples of new subspe- cies occurring while the parent species is still abundant and in a widespread and continuous range. Examples are variabilis athenius, erythrinus rugosns, aurisdianae aratrum, gibberulus albus, the marginatus complex and mutabilis ochroglottis. These subspecies are by no means equal in strength. The morphological gap is stronger perhaps due more to genetic differences in some cases, while in other cases the differences are subtle and possibly brought about in part by special geographical eco- logic conditions. In some cases, it would appear that a species has developed on the edges of the Western Pacific Arc and then re-invaded the East Indies. Such may be the case of the more easterly located bulla which now overlaps the range of its very close relative aurisdianae. Some of the shell specimens in the Philippines show a possible hybridization. These two species are normally kept apart by their respec- tive bathymetric ranges, one living in shallow, in- tertidal waters, the other from about 10 to 40 feet of water. In general, we have noted that individuals of one species are usually in shallower water in the Philip- pine area than they are in Micronesia and Poly- nesia. This is especially pronounced in the case of certain Wake Island and Hawaiian Strombus, Cy- praea and Terebra. The possible development of a deeper-water physiological race in the central Pa- cific would make possible a return invasion into the Western Pacific Arc without genetic swamping by the original parent stock. The converse or opposite direction is also possible. Discontinuous distributions in which one small center occurs in the Zanzibar-Mauritius-Seychelles triangle of the western Indian Ocean and in which a second, larger center occurs in the western Pacific are noted in the case of Strombus pipus, S. haema- stoma, S. dentatus, Strombus plicatus columba (In- dian Ocean) and S. plicatus pulchellus (Western Pacific Arc). A comparable case exists in the dis- tribution of S. terebellatus which has its typical race in the Western Pacific Arc and its Indian Ocean subspecies from Mozambique, Zanzibar, northern Madagascar and the Red Sea. Schilder and Schilder ( 1938, vol. 23, p. 178) found this same Indian-Pacific Ocean discontinuity in Cypraea ( Callistocypraea ) testudinaria Linne. [his typical testudinaria and testudinosa should be combined]. The distribution of the Pacific populations of C. testudinaria is the same as that of S. gibberulus gibbosus. This cowrie and the latter Strombus do not have the same bathymetric range, although all are associated with coral reef or coral sand habitats. Continental and Oceanic Species It is well-known that each species has its own ecological requirements, and that the distribution of such environmental factors as temperature, sub- strate, water conditions and, in the case of herbi- vores, the plant life, will all determine the geo- graphical range of the species. Although little is known about the exact ecological preferences of Strombus, we have been struck by the existence of two basic types of species that occur not only in the Strombidae but also throughout most of the marine families of mollusks and some other invertebrates. A large proportion of the Indo-Pacific and the West- ern Atlantic Strombus fall into one or the other of these two groups: 1 . Species limited to the rich, nitrogenous shores of continents or well-vegetated vol- canic islands (Strombus) : (Pacific) canarium dilatatus epidromis labiatus latissimus marginatus minimus urceus vittatus ( Caribbean ) alatus pugilis goliath 2. Species mainly living in clear, oceanic waters surrounding small coral islands, submerged banks or in similar oceanic conditions bor- dering large islands and continents ( Strom- bus): (Pacific) dentatus erythrinus fragilis gibberulus lentiginosus luhuanus decorus maculatus microurceus mutabilis sinuatus taurus ( Caribbean ) gigas costatus On the whole, the species in the second group are more widely distributed. More cases of ende- micity are developed in the first group; and some other families are almost exclusively continental or [09 - 841] 44 Strombus R. T. Abbott Strombidae large-island-dwellers, such as the Melongenidae and most of the Donacidae and Solenidae. Neither phylogenetic relationship, method of reproduction, size of the animal, relative abundance nor bathy- metric range seem to be correlated with the type of distribution. Several Strombus and about one fourth of the two hundred species in other families whose distributions we have studied seem either to be intermediate in their habitat preference or are in- sufficiently understood. Origin of the Recent Indo-Pacifie Fauna The genus Strombus first appeared in the early Miocene in the ancient Tethys Sea. It appears in the fossil beds of the Caribbean, southern Europe and the Indo-Pacific. It does not appear in any fos- sil beds which are considered by most paleontol- ogists to represent a cooler water fauna. Although the fossil record does not seem to be as full or as complete as the Recent one, it would appear that the Pliocene saw the first great development of Strombus, not only in the development of new sub- genera, but also in the geographical spread of the various species. Some time during the Pleistocene, there was an almost universal shrinkage in the ranges, particularly in the eastern and central sec- tions of the Pacific Ocean. A few typically Indo- nesian Pliocene species remained trapped in iso- lated pockets, such as Cyrtulus serotinus Hinds, Lambis robustus Swainson (formerly pseudoscorpio Lamarck) in the Societies, and such as Strombus vomer hawaiensis in Hawaii. In the main, however, the amount of speciation and the extent of distribution of these species of Strombus have remained the same from the Plio- cene to Recent times. Continental species closely resembling canarium and epidromis have remained in the southeast area of Asia. No fossils have turned up, even in the latest drill cores in the Marshalls, which would suggest that any great degree of new speciation had developed in the Central Pacific- area during the Neogene. Ladd ( 1960, Amer. fourn. Sci., vol. 258-A, p. 137) has speculated to the con- trary and suggests that the predominance of spe- cies in our Western Pacific Arc is the result of a gradual “piling up” of species as they were drifted westward by ocean currents. We support the op- posite view that the parent stocks and the majority, although not all, of late Pliocene and Pleistocene races originated in the Western Pacific Arc. A few undoubtedly originated in the Indian Ocean, while others may have started on the eastern edge of the arc. But even such endemic Micronesian and Poly- nesian species as Strombus maculatus have their ex- tremely close relatives in the Miocene of Indonesia. If Ladd’s over-emphasis of the importance of the transportation powers of the equatorial currents and winds were to be applied to the Western Pa- cific Arc, we should certainly find a greater con- centration of Micronesian-like species in the Ryu- kyu Islands and perhaps even the Marianas. In considering the migration of marine species, two factors must not be overlooked— the many millions of years involved, and the work of the counter cur- rents. How else would several hundreds of Indo- Pacific species have found their way from either the central Pacific or the Western Pacific Arc all the way to the Tuamotu, Society and Marquesas Islands? Nor are there many malacologists ready to suggest that the few obviously Indo-Pacific ele- ments in the Panamic Province ( Conus dalli, Cy- praea Isabella mexicana. Conus sponsalis mix, etc.) are the “seeds" of the Indo-Pacific fauna. In the Caribbean, the concentration of the number of spe- cies certainly does not correspond with the activi- ties of the Gulf Stream or other major currents. In fact, many formerly widespread Miocene species have retreated southeasterly against the currents and exist today only in the area of northwestern Venezuela ( Cypraea mus Linne, Strombina pu- milio Reeve, etc.). In view of the presently known fossil record, re- cent distributions, and the migratory history of marine species in the Indo-Pacific and other oceans, we believe that the central part of the Western Pacific Arc (i.e. the Philippines or Indonesia) is the center of origin of most of the Pacific races and of some of the Indian Ocean races. The paucity in number of species in such areas as eastern Poly- nesia, the central atolls of the Indian Ocean, the Bahamas and such small coral islands as Grand Cayman Island (West Indies), Easter Island (east- ern Polynesia ) and Cocos-Keeling ( Indian Ocean ) are probably due to poor ecological conditions and not to the fact that they are isolated from migrant- carrying currents. [09 - 842] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 45 Doubtful species of Strombus Faustino ( 1928, Monograph 25, Bureau of Sci- ence, Manila, pp. 211-214) lists 30 species of Strom- bus from the Philippines. He erroneously listed Strombus hebraeus Linne and Strombus textile Linne, doubtlessly meaning Linne’s two well-known species of Conus. Some early authors prior to 1842 employed the genus Strombus to such modern genera as Tunis, Rissoina, Thiara, Faunus and Morum. Strombus elatus Anton, 1839 Range— Indo-Pacific? Remarks— It would appear from Anton’s descrip- tion that this species is related to vittatus Linne or labiosus Wood. It was not figured. Anton lists Strombus turritus Lamarck, so it is not that form of vittatus. Synonymy— 1839 Strombus elatus Anton, Verzeichniss tier Conehylien, Halle, p. 85, no. 2799 (no locality). Strombus sulcatus Anton, 1839 Range— Mexico [?]. Remarks— If from Mexico, this species may be a synonym of Strombus gracilior Sowerby or S. pu- gilis Linne. It was not figured and its description could apply to several species. Non sulcatus Hol- ten, 1802. Synonymy- 1839 Strombus sulcatus Anton, loc. cit., p. 85, no. 2798 ( Mexico ) . Strombus tubercularis Anton, 1839 Range— Unknown; Indo-Pacific? Remarks— This species was not illustrated and it’s description meagre, although it is quite possible that it is a synonym of the dwarf, knobbed form of Strombus decorus (Roding) from East Africa. Synonymy- 1839 Strombus tubercularis Anton, loc. cit., p. 86, no. 2813 (no locality). Strombus parvulus Krumbeck, 1906 Range— Tertiary (weisser Versteinerungskalk) Tripoli. Remarks— This is such a poorly preserved fossil that it is very doubtful that it is a Strombus. Synonymy- 1906 Strombus parvulus Krumbeck, Palaeontographica, Stuttgart, vol. 53, p. 118 ( Djebel Tar, Tripoli). Strombus labrosus Menke, 1829 Range— Unknown. Remarks— This is an unfigured dubious species which Morch says is a variety of luhuanus Linne. Synonymy- 1829 Strombus labrosus Menke, Verz. Conch. -Samml. Mals- burg, p. 59, no. 1209; 1871, Morch, Malak. Blatt., vol. 18, p. 127. Strombus glabratus Sowerby, 1842 Range— West Africa from Angola to French Guinea. Remarks— This species evidently belongs in the family Nassariidae and is the type of the genus Naytia H. and A. Adams, 1853. It is a synonym of Naytia oblicpia (Kiener, 1841). However, Sowerby’s name must continue in use, since Kiener’s Ruc- cinum obliquum is a homonym of obliquum Gme- lin, 1791. Synonymy— 1842 Strombus glabratus Sowerby, Thesaurus Concliylio- rum, vol. 1, p. 32, pi. 8, figs. 66, 67 (no locality). Strombus moisei Cuvillier, 1930 Range— Upper Eocene, Bartonian of Egypt. Remarks— I doubt if this is a Strombidae, and, if I interpret the four columellar plicae correctly, it should be placed in the family Vasidae. Synonymy- 1930 Strombus moisei Cuvillier, Memoires l’lnstitut d’Egypte, vol. 16, pp. 253 and 323, pi. 20, figs. 15 and 16 (Ain Mouca, Bartonian of Egypt). Strombus altispirus King, 1953 Range— Miocene of Zululand, South Africa. Remarks— Not demonstrated, in my opinion, to be a Strombus. Synonymy— 1953 Strombus altispirus King, Trans, and Proc. Geol. Soc. South Africa, vol. 56, p. 77, fig. 9, pi. 12, fig. 32 ( Sapolwana, Zululand, Miocene ) . Strombus mimasakensis Yokoyama, 1929 Range— Miocene or Pliocene of Japan. Remarks— Based upon a single large broken shell which could be a Melongena, or possibly a Strom- bus resembling the fossil S. maximum Martin from Java. Synonymy- 1929 Strombus mimasakensis Yokoyama, Journ. Faculty Sci. Imp. Univ. Tokyo, sect. 2, vol. 2, pt. 8, p. 366 ( Mimasaka, Chugoku, Japan). [09 - 849] 46 Doubtful species R. T. Abbott Strombidae Strombus martapurensis K. Martin, 1889 Range— Tertiary of Indonesia. Remarks— This does not appear to be a Strombus, and possibly not even a member of the family. Synonymy— 1889 Strombus martapurensis K. Martin, Jaarbock Mijn- wegen in Nederland. Oost-Indie, Amsterdam, vol. 18, p. 66, pi. 20, fig. 13; 1889, Samml. Geol. Reichs- Mus. Leiden, 1st series, vol. 4, p. 189, pi. 20, fig. 13. Strombus bivaricosus Nomura, 1935 Range— Pliocene of Taiwan. Remarks— N omura likened this possible juvenile form to S. sondeianus Martin. It may also be the young of S. taiwanicus Yabe and Hatai, 1941. Synonymy- 1935 Strombus bivaricosus Nomura, Science Reports Tohoku Imperial Univ. Sendai, 2nd ser., vol. 18, no. 2, p. 178, pi. 8, figs. 14 a, 14 b (Byoritu Beds, Pliocene, 1000 meters S.E. of Hakusyaton, Taiwan). Strombus sondeianus Martin, 1906 Range— Pliocene of Java, Indonesia. Remarks— This is possibly the young of S trombus fennemai Martin. Synonymy— 1906 Strombus sondeianus K. Martin, Samml. Geol. Reiclis- Mus. Leiden, n. Folge, vol. 1, pt. 9, p. 319, pi. 45, figs. 739, 740 (Sonde, Java, Pliocene). Bibliography The majority of the bibliographic references to Strombus appear in the synonymies under the vari- ous species. Some of the references, listed by author, date and pagination, which appear in the sections on locality records may be located in the Zoological Record. The standard iconographs which dealt with S trombus during the nineteenth century are listed in the synonymies (Sowerby, 1842; Kiener, 1843; Duelos in Chenu, 1844; Reeve, 1850- 51, and Tryon, 1885). A useful bibliography of fos- sil Indonesian species is found in van der Vlerk, 1931 (see below). Abbott, R. Tucker. 1949. Sexual Dimorphism in Indo-Pacific Strombus. Nautilus, vol. 63, no. 2, pp. 58-61, 2 graphs. [S. gibberulus Linne and S. mutabilis Swainson.] Beets, C. 1940. Mitteilung fiber Messungen an Strombidae aus der Sammlung-Schroder von Nias. Geologie und Mijnbouw, Jaargang 21, no. 2, pp. 17-25. [Bionomics of shell of Strombus labiosus Wood.] Bergh, R. 1895. Beitriige zur Kenntniss der Strombiden, besonders der Gattung Terebellum Klein. Zoologische Tahrbiicher, Abt. Anatomie und Ontogenie der Thiere, Jena, vol. 8, pp. 342-378, pis. 22 and 23. [Anatomy.] Chadwick, George H. 1899. An Attempt to Define the Natu- ral Groups of Strombs. Nautilus, vol. 13, no. 7, pp. 76-78, no. 8, pp. 93-96. Clench, W. J. and R. T. Abbott. 1941. The Genus Strombus in the Western Atlantic. Johnsonia, vol. 1, no. 1, pp. 1-15, 10 pis. Fischer, Paul. 1861. Note sur les organes visuels des Strom- bus. Journal de Conchyliologie, vol. 9, pp. 213-220. Issel, A. and C. Tapparone Canefri. 1876. Studio Monografico sopra gli Strombidi del Mar Rosso. Annali del Museo Civico di Storia Naturale di Genova, vol. 8, pp. 337-366. Kobelt, W. 1876. Catalog der Gattung Strombus Linne. Jahrbiicher der Deutschen Malakozoologischen Gesell- schaft, vol. 2, pp. 255-262. Robertson, Robert. 1959. Observations on the Spawn and Veligers of Conehs (Strombus) in the Bahamas. Proceed- ings Malacological Society of London, vol. 33, pt. 4, pp. 164-171, pi- 11 and text figs. 1 and 2. Robertson, Robert. [1961], The Feeding of Strombus, a Herbivorous Marine Gastropod. Notulae Naturae, in press, van der Vlerk, I. M. 1931. Caenozoic Amphineura, Gastro- poda, Lamellibranchiata, Scaphopoda. Leidsche Geologi- sche Mededeelingen, Leiden, deel 5, pp. 206-296. [Strom- bus on pp. 246-248.] [09 - 850] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 47 Subgenus Strombus Linne, 1758 Type: Strombus pugilis Linne, 1758 The typical subgenus of Strombus is not, in our opinion, represented in the Recent Indo-Pacific. It exists in tropical American waters— pugilis Linne 1758, and alatus Gmelin, 1791, of the Caribbean and Florida, respectively; and g racilior Sowerby, 1825, of the tropical Eastern Pacific. This stock has been in the Caribbean for a long time in the form of such Miocene species as Strombus proximus Sowerby, 1850, bifrons Sowerby, 1850, and pugi- loides Guppy, 1873. There appears to be some close eonchological kinship between the above species and the Indo- nesian Miocene species S trombus varinginensis Martin. From this stock, it appears that Strombus canarium of the Recent Indo-Pacific could have arisen, especially when the sculpture and color pat- terns of the Caribbean Miocene proximus and ca- narium are compared. However, I find no very close relationships between the radulae and soft parts of pugilis and canarium, and, hence, have arbitrarily accepted Laevistrombus as a valid sub- genus for canarium. Strombus sensu stricto is characterized by solid, medium-sized shells with a broadly fusiform shape, a single row of shoulder spines and a smooth colu- mella. The operculum is strongly serrated. The accessory pad of the penis bears a peculiar prong. The central tooth of the radula is quadrate; the lateral very stout, conic and without smaller cusps; and the two marginals thin, narrow and with 5 to S denticles. The Western Atlantic species were treated by Clench and Abbott in Johnsonia, vol. 1, no. 1, pp. 1-16, 1941. I would modify that treatment by ac- cepting alatus Gmelin, 1791, as a full species, and by considering nicaraguensis Fluck, 1905, as merely a form of pugilis. Synonymy— 1758 S trombus Linne, Systema Naturae, ed. 10, p. 742, no. 289. Type by subsequent designation ( Montfort, 1810): Strombus pugilis Linne; also possibly La- marck 1799, p. 72 who listed only pugilis Linne. 1838 S trombella F. Schliiter, Kurzg. Syst. Verz. Conch., p. 22. Type by monotypy: S. pugilis Linne. Subgenus Laevistrombus Kira, 1955 Type: S trombus canarium Linne, 1758 This subgenus appears to be limited to the Indo- Pacific, and contains only one living species— cana- rium Linne. Six fossil species, described from the Miocene and Pliocene of Indonesia and India, ap- pear to belong to this group. The shells are thick, generally rotund and in- clined to be smoothish. The columella is smooth, the outer lip thick. The operculum bear 7 to 8 small serrations. The penis is a simple prong, usually dark-maroon or brown in color. The central radular tooth is ovoid; the lateral with a basal peg, and the marginals thin and with only 2 or 3 cusps. The group does not seem to have survived in, or Plate 21. Strombus pugilis Linne, type of the genus Strom- immature. 3, enlargement of apex. 1 and 2, slightly reduced. bus Linne, 1758. Santo Domingo, West Indies. 1, adult. 2, [09 - 855] 48 Laevistrombus R. T. Abbott S trombidae possibly ever reached, East Africa or the Red Sea. Odhner, 1919, probably erroneously reported ca- narium from Madagascar. Synonymy— 1955 Laevistrombus Kira, Coloured Must, of the Shells of Japan, Osaka, ed. 1, p. 31; 1959, ibid., ed. 2, p. 36. Type hereby designated: Strombus canarium Linne. S trombus canarium Linne, 1758 (PI. 17, figs. 15, 16; pis. 12, 22, 23, 24) Range— Southern India to Australia and Mela- nesia, and north to Japan. Remarks— This well-known species is usually abundant wherever it occurs. It is not a coral-water species, but rather is associated with larger islands and continental shores where sandy mud exists. Its heavy, smooth brownish shell, its white smooth aperture and its thick, winged outer lip are the dis- tinguishing characters. This species is very often variable in size, shape and color pattern. The length of adult shells in some colonies ( fapen Id., Dutch New Guinea) may not exceed 40 mm., while on nearby Soepiori Id., Schouten Id., Dutch New Guinea, the median adult length is 86 mm. Some colonies exhibit a wide range in size (Tabaco, Albay Prov., Luzon Id., Philippines) from 4 2 to 98 mm. There are two color forms which may occur in the same individual at different stages of growth, one of a network of light or dark-brown streaks, the other of a unicolor wash of light (or rarely dark) yellow-brown. The latter color form is more common in larger specimens. Plate 22. Figs. 1-5, Strombus canarium Linne, Gubat, Lu- zon Icb, Philippines. Fig. 1, almost mature, remaining figures The netted form is the typical canarium ; the plain color phase is the form turturella (Roding) and Isabella Lamarck. The spires of the shells may also vary, some being high and with more angular peri- pheries, others being short and more flat-sided. Although some authors have considered the above forms as separate species, I can find many inter- grades, combining characters, and no geographical pattern. Shells from New Caledonia and Queens- land are commonly (although not all) very darkly colored. The aluminum-like glaze on the apertural lip is a sign of maturity or old age which is prob- ably also correlated with diet or water conditions. Habitat— Lives in large colonies on sandy mud and algae bottoms from low tide mark to a depth of about 20 feet. Risbec ( 1935, vol. 60, p. 409 ) describes the egg mass as a long, entangled, single, gelatinous tube with a single row of ova. Description— Adult shell 31 to 97 mm. ( 1M to 3/i inches) in length, solid, heavy, globose, smooth, light-brown and with a flaring lip. Nuclear whorls 3, smooth, translucent whitish, yellow or tan, and rounded. Postnuclear whorls brown, moderately rounded, and with either 7 to 9 spiral cords or wide, incised lines. Numerous axial riblets may be present to form a gross reticulate pattern. By the second or third whorl, only incised spiral lines are present, and the remaining whorls are sculptureless, except for 6 to 15 weak spiral threads or incised lines at the base of the shell. Apex with 4 to 16 swollen, whitish, rounded, former varices, the early ones being spirally incised or corded, the later ones inclined to be smoothish. Penultimate whorl flat- tish, rounded, or barely angular. Last whorl roundly represent immature specimens. All natural size. [09 - 856] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 49 swollen at the shoulder. Color of shell variable; either a uniform light yellow-brown or chestnut, or densely covered with a fine, zigzag network of darker brown. Columella straight, swollen, glazed, smooth and enamel-white. Outer lip thick, rounded, and its upper end projecting slightly upward. It and the parietal wall may have a brown or gray, aluminum-like glaze. Siphonal canal short. Strom- boid notch very shallow. Periostracum rather thick, reticulated, yellow-brown and fimbriated at the su- tures. Operculum stromboid, dark-brown, one third the length of the shell, slightly arching and with 7 to 8 weak serrations. Radula ribbon 4 mm. in length, with 40 to 45 rows of teeth, and amber to wine-red in color. For- mula: 3-1-3; 1-5 to 1-9 (plus peg); 3; 2. Verge simple and with a broad swollen distal end; dark- maroon in color. Posterior mantle filament small. Measurements (mm.)— length width no. whorls 97.0 62.1 9 + 90.1 51.0 9 + 67.5 40.0 10 31.0 22.2 6 + Synonymy— 1758 S trombus canarium Linne, Systema Naturae, ed. 10, p. 745, no. 438 (In O. Asiae); 1767, ed. 12, p. 1211, no. 507; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 275-276. 1798 Lambis turturella Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 65, no. 833. Refers to Conchyl.-Cab., vol. 3, fig. 817. 1822 S trombus Isabella Lamarck, Anim. sans Vert., vol. 7, p. 207 ( Grandes Indes). Refers to Conehyl.-Cab., vol. 3, fig. 817. 1834 Strombus vanikorensis Quoy and Gaimard, d’LTrville’s Voy. L’ Astrolabe, Zoologie, vol. 3, p. 73, pi. 51, figs. 7-9 (Vanikoro). 1834 Strombus taeniatus Quoy and Gaimard, ibid., p. 75, pi. 51, figs. 14-15 (Amboine). 1876 Strombus gibbus “ Martini” Issel and T.-Canefri, An- nali Mus. Civico Storia Nat. Genova, vol. 8, p. 344; 1895, R. Bergh, Zoologische Jahrbiicher, Jena, vol. 8, pp. 359-362, pi. 23, figs. 50-53 (anatomy). 1885 Strombus vanicorensis Quoy, Tryon, Manual of Conch., Phila., vol. 7, p. 1 10. Types— “ The type specimen of canarium, marked by Linnaeus, is found in the Linnaean collection in London at the Linnaean Society of London" ( Dodge, 1956, p. 276). We hereby designate Amboina, In- donesia, as the type locality, since Linnaeus merely gave "In O. Asiae”. Nomenclature— Under our remarks we discussed our reasons for considering Isabella Lamarck as merely a form of canarium. Even if accepted as a subspecies, Lamarck’s Isabella would have to take the earlier name of turturella (Roding, 1798). Selected records ( see accompanying map, pi. 23 ) . Solid dots: specimens examined; open circles: literature records) — INDIA: Goa (J. E. Bridwell, USNM); Pamban and Tuti- corn. Gulf of Manaar ( E. Thurston, 1895, p. 125 ) . CEYLON : Karaitivu Id., and Trincomalee (George and Mary Kline, NSF). ANDAMANS: Port Blair (W. N. Carpenter, USNM). SINGAPORE: muddy sand, algae, intertidal at Tanak Merah Besar ( R. D. Purchon, ANSP). THAILAND: Gulf of Siam: Sutamarat; Taluei Id.; Bangbert Bay; Koh Samui; Koh Samit; Lem Sing (all USNM). CHINA: Cheefoo (Yen- tai) (Ping and Yen, 1932, p. 48 [accurate?]). HONG KONG: S.W. of Urn Id., Rock}' Harbour (A. J. Staple, ANSP). JAPAN: Shirahama, Wakayama Pref., Honshu Id. (T. Habe, ANSP). PHILIPPINES: common on most of the islands: Luzon; Leyte; Samar; Catanduanes; Negros; Basi- lan; Panay; Palawan; Balabac; Polillo; Busuanga; Mindoro; Mindanao; Cuyo (ANSP and USNM). INDONESIA: (see Oostingh, 1923, p. 82 for many records). Sarawak, Borneo (MCZ). AUSTRALIA: Queensland: Port Douglas (Tony Marsh, ANSP); Bedford Beach, Cooktown (MCZ); Hay- man Id., Cumberland Group (MCZ). NEW CALEDONIA: Touho Bay; Baie des Prunes; Baie de FOrphelinat ( all G. and M. Kline, 1958, NSF). NEW HEBRIDES: Lamap, Mallicolo Id. (G. Massoulard, ANSP). [Records, such as Odhner, 1919, p. 35, for the western part of the Indian Ocean are probably based on mixtures or misidentifications.J ( giant; Luzon Id. ) ( large; Schouten Ids. ) ( average; Luzon Id. ) (small; New Guinea) Plate 23. Geographical distribution of Strombus canarium Linne. [09 - 857] 50 Laevistrombus R. T. Abbott Strombidae Plate 24. Strombus canarium Linne, New Caledonia. Fig. 1, central radular tooth (two views). 2, lateral, showing basal peg. 3, inner marginal. 4, outer marginal. 5, verge. 6, operculum. Fossil records— Schepman (1907, p. 185) reports this spe- cies from the Posttertiary of the Celebes. Altena ( 1942, vol. 22, pp. 47-49) reports canarium from the Pliocene of New Guinea, Indonesia and the Philippines, but certainly sonv of these are varinginensis. Dickerson’s Vigo Group, Luzon Id. specimens are probably varinginensis. True canarium occurs in the upper Pliocene of Niki-Niki, Timor, Indonesia (Tesch, 1920, p. 48, pi. 129, fig. 165). Tesch also claims that it occurs at Nias Island (off west Sumatra); Sonde, Java; Gorontalo, Celebes; Fialarang, Timor; and Cheribon, Java, although I have not seen these specimens nor figures of them. Beets (1948, Basteria, vol. 12, p. 8) records five Quaternary specimens from Goenoeng Mendong, eastern Borneo. Abrard ( 1946, p. 59, pi. 4, fig. 23) records this spe- cies as Isabella from the Nua River Pliocene on Malekula Id., New Hebrides. Strombus varinginensis Martin, 1899 Range— Pliocene of Java and Borneo, Indonesia, and Luzon Island, Philippines. Remarks— This and the following fossil species are related to the Recent Strombus canarium Linne or possibly vittatus Linne. Synonymy— 1899 Strombus (Strombus) varinginensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 184, pi. 30, figs. 426-429 (Tji Djadjar, Cheribon Residence, Java, here designated as the type lo- cality ) . 1922 S trombus canarium L., Dickerson, Philippine Journal of Science, vol. 20, p. 202, pi. 5, fig. 3 (Vigo Group, Upper Miocene, Luzon Id.). 1935 Strombus (Labiostrombus) varinginensis varinginensis K. Martin, Oostingh, Wetenschappelijke Mededeel., no. 26 (Dienst van den Mijnbouw in Nederl.-Indie), p. 56; 1948, Cox, Schweizerische Palaeontologische Abhandl., vol. 66, p. 25, pi. 2, figs. 3, a, b ( Dent Peninsula, Borneo). Plate 25. Figs. 1 and 5, Strombus varinginensis Martin. Types from the Pliocene of Java, (from K. Martin, 1899, pi. 30, figs. 427, 428). Figs. 2 and 6, S. varinginensis mar- tini Oostingh ( type of S. isabella var. thersites Martin, non Swainson; from K. Martin, 1899, pi. 30, figs. 423, 424; Pliocene of Java). Figs. 3 and 4, Strombus glaber Martin. Types from the Upper Miocene of Java, (from K. Martin, 1879, pi. 9, fig. 6). All natural size. Plate 26. Strombus varinginensis Martin. Dent Haven, British North Borneo. Pliocene, (from Cox, 1948, pi. 2, figs. 3a, 3b). x 2. Strombus varinginensis subspecies martini Oostingh, 1935 Range— Pliocene of Java, Sumatra, Borneo and Quarternary of New Guinea. Remarks— This was originally named thersites Martin, 1899 (non Swainson, 1823). Oostingh con- sidered it a subspecies of varinginensis; Cox raised it to specific rank despite the presence of the two, only slightly differing, in the same fossil bed in Bor- neo. It is probably only a form. [09 - 858] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 51 Synonymy— 1899 Strombus Isabella Lam. var. thersites K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 184, pi. 30, figs. 423-425 (Java, Pliocene). Non Swainson, 1823. 1935 Strombus (Labiostrombus) varinginensis martini Oost- ingh, Wetenschappelijke Mededeelingen, no. 26 ( Dienst van den Mijnbomv in Nederlandsch-Indie), p. 57 (new name); 1942, Oostingh, Leidsche Ge- ologische Mededeel., vol. 22, p. 49. 1948 Strombus ( Labiostrombus ) martini Oostingh, Cox, Schweizerisch Palaeontologische Abhandl., vol. 66, p. 24, pi. 2, figs. 9a, b (Borneo, Pliocene). Plate 27. Strombus martini Oostingh. Dent Haven, British North Borneo. Pliocene, (from Cox, 1948, pi. 2, figs. 9a, 9b). Natural size. Strombus overbecki Cox, 1948 Range— Pliocene of Borneo. Remarks— Quite possibly a malformed specimen of varinginensis Martin. Synonymy- 1948 Strombus ( Labiostrombus ) overbecki Cox, Schweizer- isch Palaeontologische Abhandl., vol. 66, p. 26, pi. 2, fig. 7 (7 km. inland from Dent Haven, Dent Peninsula, Borneo). Plate 28. Strombus overbecki Cox. Dent Haven, British North Borneo. Pliocene. Adult, holotype. (from Cox, 1948, pi. 2, figs. 7, a, b, c). Natural size. Strombus glaber Martin, 1879 Range— Upper Miocene of Java, Indonesia. Remarks— Allied to S. varinginensis Martin. From the Tjilanang beds, Upper Miocene of Java (van der Vlerk, 1931, p. 246 ). Synonymy- 1879 Strombus glaber K. Martin, Die Tertiarschichten auf Java, Leiden, p. 49, pi. 9, fig. 6 (Java); 1905, ibid., n. Folge, vol. 1, pt. 9, p. 319, pi. 45, fig. 738 ( Palabuan-ratu, Java, Pliocene). Strombus karikalensis Cossmann, 1903 Range— Fossil (Pliocene) of southeast India. Remarks— Allied to S trombus varinginensis mar- tini Oostingh. Other than its smoothish spire, this shell resembles the Recent S. vittatus vittatus Linne. Synonymy— 1903 Strombus karikalensis Cossmann, Journ. de Conchyl., vol. 51, p. 164, pi. 6, figs. 12, 13 (Karikal, French India ) . [09 - 859] 52 Laevistrombus R. T. Abbott Strombidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [09 - 860] November 23, 1960 Strombus 53 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Subgenus Tricornis Jousseaume, 1886 Type: Strombus tricornis Humphrey, 1786 Members of this subgenus, whose type is tri- cornis Humphrey, are variable in size, shape of the outer lip, and details of the apical sculpturing. The subgenus has within it several species-groups, but, because so many of the so-called generic characters appear in random combination within almost any one species, it would be impossible to recognize ad- ditional subgenera without erecting one for each species. In general, the subgenus Tricornis is char- acterized by large, heavy shells which have promi- nent axial knobs, a smooth columella, and a fairly thick, usually deciduous periostracum. The oper- culum is proportionately broad and its edge smooth or with very weak, almost obsolete, serrations. The radular teeth are strong; the central with a large middle cusp, flanked on either side by 1, 2 or 3 smaller cusps. The lateral lacks a distinct, basal peg, and the two marginals are relatively broad with 4 to 6 large cusps. The tropical Western Atlantic contains costatus Gmelin, 1791, raninus Gmelin, 1791, goliath Schroter, 1805 (=Dillwyn, 1823, Sowerby, 1842), g alius Linne, 1758, and gigas Linne, 1758. The tropical Eastern Pacific contains peruvianas Swain- son, 1823, and galeatus Swainson, 1823 (allied to goliath, Recent, Brasil). The Indo-Pacific contains tricornis Humphrey, 1786 (from the Red Sea), thersites Swainson, 1823, taurus Reeve, 1857 and sinuatus Humphrey, 1786. The earliest fossil recorded is inflatus Martin, 1879, from the Lower Miocene of Java. It, herklotsi Martin, 1880, and tjilonganensis Martin, 1899 from the Miocene of Java, and trigonus Gratelonp from the European Dax Lower Miocene are probably forerunners of tricornis and costatus. Also related are Strombus haitensis Sowerby, 1849, ( Miocen \ Santo Domingo), g alliformis Pilsbry and Johnson, 1917 (Miocene, Santo Domingo), dominator Pi’.s- bry and Johnson, 1917 (Miocene, Santo Domingo), chipolanus Dali, 1900 (Florida, Pliocene), and leidyi Heilprin, 1887, (Pliocene of Caloosahatchee, Florida). The latter is represented by a recent form or subspecies in Bermuda’s isolated, cooler water. S trombus fortisi var. valdetubercidatns Innocenti (Boll. Soc. Geol. Itah, vol. 47, p. 60, pi. 2, fig. 6) re- ported from the Eocene of Ronca, Italy, is based upon a poor and almost unidentifiable specimen. Strombus bravardi Borchert, 1901, from Brazil is not a Strombus. Synonymy— 1886 Tricornis Jousseaume, Le Naturaliste, Paris, 1st series, vol. 3, 8th year, no. 28, p. 220. Type by monotypy: T ricornis tricornis Lamarck = tricornis Humphrey, 1786; 1888 Jousseaume, Mem. Soc. Zool. de France, vol. 1, p. 174. 1921 Lobatus “Swainson” Iredale, 1921, Proc. Mai. Soc. London, vol. 14, p. 208. Type by monotypy: Strom- bus bituberculatus Lamarck = raninus Gmelin, 1791. 1929 Aliger Thiele, Handbuch Systemat. Weichtierkunde, Jena, vol. 1, pt. 1, p. 254. Type by monotypy: Strombus gallus Linne. 1940 Eustrombus (subgenus of Strombus) Wenz, Hand- buch der Palaozoologie, lief. 6, band 6, p. 945. Type by original designation: S trombus gigas Linne. Nomenclature— Iredale stumbled into Lobatus and first validated it in 1921 when he erroneously assumed that the name had appeared as a genus in 1837, presumably belonging to Swainson. In the anonymous “Catalogue of the Foreign Shells in the Possession of the Manchester Natural History So- ciety (Manchester ?, 1837) there appears on page 75 the line “[Strombus] bituberculatus ( Lobatus , Swainson).” S. lobatus Swainson, 1822 is known to be a synonym of bituberculatus Lamarck, and the name in parentheses probably refers to this species, since on page 76 another synonym is listed with a capital letter: “[Strombus] exustus ( Tapilio ) La- marck,” and not meant as a genus name. Strombus tricornis Humphrey, 1786 (PI. 17, figs. 6, 7; pi. 29, figs. 1, 2) Range— Red Sea and Gulf of Aden. Remarks— The rather heavy, strongly knobbed shell with its single spine protruding from the up- per and outer corner of the outer lip is distinct for this moderately common species from the Red Sea area. The elongate, axially pinched knob on the center of the dorsum is always the largest. The color of the aperture and outer shell is quite vari- able, the former ranging from white to tan to weak rosy tan, and the latter from white to strongly flecked and suffused with soft browns. Old speci- mens are apt to have a brownish aluminum-like glaze around the aperture. Adults vary greatly in size. We have been unable to verify records in the old literature of this species occuring in Reunion, Mauritius, the Seychelles and the West Indies. It is interesting to note that a figure was published of this species over 400 years ago. Habitat— Little is known about its habitat, ex- cept that it occurs in shallow water just below the low tide line. [09- 871] 54 Tricornis R. T. Abbott S trombidae Plate 29. Figs. 1 and 2, Strombus tricornis Humphrey, immatures. Port Sudan, Red Sea. Roth natural size. Description— Adult shell 64 to 125 mm. ( 2b to 5 inches) in length, solid, moderately heavy, with large blunt spines on the body whorl, and with a long, triangular projection on the upper and outer end of the outer lip. Color of shell whitish with varying degrees of black-brown to light brown Hecks, maculations or axial streaks. Rarely albin- istic. Whorls 10. Nuclear whorls 2, smooth, translu- cent white. Postnuclear whorls carinate, bearing numerous axial riblets which become small knobs (12 to 16 per whorl) just above the minutely in- dented suture in later whorls. Early whorls with 15 to 20 microscopic threads which become obsolete in the last whorl. Spire rarely with 2 or 3 small, whitish former varices. Shoulder of body whorl with 3 to 4 rather large, pyramidal knobs, the last one usually small, and the second to last one being the largest, slanting and axially pinched. Parietal wall glazed. Columella smooth, white or tinged with tan, rose or lavender. Interior of aperture, smooth enamel-white, rarely tan, orangish or pink- ish, and having at the top of the body whorl a sunken, spiral depression or trough. Outer lip thick, somewhat wavy and with its upper and outer end produced into a long triangular to elongate spine which is longer than the spire. Siphonal canal short. Stromboid notch broadly U-shaped. Periostracum moderately thin, somewhat rough, and apt to flake off when dry. Operculum and soft parts unknown. Measurements (mm.)— (excluding spine) length width no. whorls 63.9 46.0 9 (small; Red Sea) 101.5 82.1 9 + ( average; Red Sea 125.0 96.2 8 + (large; Red Sea) Synonymy— 1558 Murex lacteus Rondelet, De Natura Aquatilium Car- men . . . Lugduni, p. 37 (non-binomial). 1786 Strombus tricornis Humphrey, Portland Catalogue, London, p. 5, no. 50 (no locality); refers to Chem- nitz Conchyl.-Cab., vol. 3, figs. 843-845 and Lister, fig. 873. 1807 Strombus tricornis G. Fischer, Museum Demidoff, Moscow, p. 188 (Jamaique); refers to Davila, p. 183, and Chemnitz Conchyl.-Cab., vol. 3, figs. 843- 845. 1816 Strombus tricornis Lamarck, Le Liste, p. 3; pi. 408, fig. 1 of Encycloped. Method, (no locality); 1954, P.-H. Fischer, Jour, de Conchyl., vol. 94, no. 4, pp. 152-153. 1844 Strombus orientalis “Jonston”, Duclos, in Chenu, Illus. Conchyl., vol. 4, Strombus, p. 15, pi. 18, figs. 5 and 6, pi. 21, figs. 1 and 2 (Les Antilles). 1844 Strombus pertinax Duclos, in Chenu, Illus. Conchyl., vol. 4, p. 15, pi. 29, figs. 1, 2 (no locality given). Types— Humphrey’s type was sold at public auc- tion in 1786, and its whereabouts is unknown to us. No locality was given, so we hereby designate the Red Sea as the type locality. G. Fischer’s type may be in the zoological museum of the University of Moscow. He referred to a figure in Chemnitz of our tricornis, but his locality of “Jamaique suggests an error in datum or that he had S trombus gallus Linne from the West Indies. [09 - 872] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 55 Nomenclature— The author of tricornis is some- times listed as Solander, sometimes as Humphrey. We prefer the latter, since we believe he wrote the Portland Catalogue and merely consulted So- lander’s manuscript card file for names. Records— RED SEA: Aqaba, Israel (Fischer, 1870, p. 162). Ras Domeirah, Assab Bay, Dahlac Archipelago ( Issel and T.-Canefri, 1876, p. 340). Port Sudan (ANSP). 20 miles north of Jidda, Saudi Arabia ( C. Aslakson, ANSP). Ras Banas, Egypt (USNM). Massara, Eritrea (MCZ). GULF OF ADEN: Djibouti (MCZ); Berbera (USNM). [Records for the Seychelles, Amirantes, Reunion, Andamans, Philippines and West Indies are unverified and the latter two certainly erroneous.] Fossil records— SUDAN: raised coral reefs, Port Sudan. Pleistocene (Hall and Standen, 1907, p. 67). EGYPT: Pleis- tocene, beach 80 ft. alt.. Wadi Gueh ( R. B. Newton, 1900, p. 508). FRENCH SOMALIA: Pleistocene, Loyada; Dou- meira; Ras Doumeira (Abrard, 1942, vol. 18, p. 63, pi. 6, fig. 35). Strombus thersites Swainson, 1823 (PI. 17, figs. 1, 2) Range— Ryukyu Islands, New Caledonia (and Society Islands?) Remarks— This is one of the rarest of the large Strombus of the Indo-Pacific. It is heavy and mas- sive like latissimus, but has a less developed outer lip, is lighter in color, and its angle of spire is about 45, rather than 70 to 75, degrees. The knobs on the spire are fewer, more pronounced and more or less evenly developed throughout the entire spire. The two Society Island records are open to question. It may occur in the areas between the Ryukyu Islands and New Caledonia. It occurred in Fiji during the Pliocene. Habitat— Kira ( 1959, p. 37 ) reports that it occurs at a depth from 5 to 10 fathoms. Description— Shell 110 to 146 mm. (4 to 6 inches) in length, massive, with a heavy outer lip, its spire with small knobs, and its body whorl with one large and 2 small knobs. Color of shell whitish with sparse, somewhat zigzag, light yellow-brown streaks. Aper- ture and columella enamel white, rarely with an aluminum-like glaze. Whorls about 11; spire angle about 45 degrees. Nuclear whorls not observed. First few postnuclear whorls with numerous axial riblets. The last 4 whorls (but not the body whorl) each bear just above the wavy suture 9 to 11 promi- nent, smoothish, rounded knobs which gradually increase in size. Spiral sculpture absent or extremely weak. A few former varices may be present in the apex. Last whorl with one large, longitudinally pinched knob on the left side of the shoulder. The dorsum bears one to three slightly raised, weak nodules. Columella, aperture, and outer lip smooth and with a white enamel glaze. Top of outer lip moderately produced. Middle of outer lip thick, glazed and with 5 to 7 light brown color bars. Stromboid notch deep and well-developed. Perio- stracum very thin, smooth and translucent tan. Operculum and soft parts unknown. Measurements (mm.)— length width no. whorls 145 90 8? ( from Wilkins; lectotype 146 86 9 (large; Okinawa) 142 94 9 (large; New Caledonia) 134 78 9 ( average; Okinawa ) 110 78 8 (small; Okinawa) Plate 30. Geographical distribution of Strombus thersites land, Australia. Swainson. There are also two literature records for Queens- [09 - 873] 56 Tricornis R. T. Abbott Strombidae Synonymy- 1823 Strombus thersites Swainson, The Philosophical Maga- zine, vol. 62, p. 401 (New Caledonia); 1951, Wil- kins, Proc. Mai. Soc. London, vol. 28, pt. 6, p. 238, pi. 29. 1828 Strombus thersites Wood, Index Testaceol. Suppl., London, id. 14, pi. 4, fig. 17 ( locality unknown ) . 1842 Strombus thersites Gray, Reeve, Conchologia System- atica, vol. 2, pi. 249, fig. 1, p. 206. 1842 Strombus ponderosus Philippi, Abbild. Beschr. Con- chy!., vol. 1, pt. 1, p. 7, pis. 2 and 3 (Oceanus Paeificus ) . 1885 Strombus (Euprotomus) ponderosus Phil., Tryon., Manual of Conchology, vol. 7, p. 111. Types— New Caledonia is the type locality. Wil- kins, 1951, designated and illustrated the lectotype (Cracherode collection, 1799) now in the British Museum (no. 1950. 11. 15. 1). Locality records (see accompanying map, pi. 30)— RYU- KYU ISLANDS: Amami Id. (Kira, 1959, p. 37); Okinawa Id. (A. A. Scott and A. R. Calm, ANSP). NEW CALE- DONIA: (ANSP; BM; MCZ ) ; Touho (D. Getz). Brampton Reef, 19° 51' S; 158° 20' E, (J. Brazier, 1871, p. 585). LOYALTY ISLANDS: Chepenehe, Lifu (D. Getz). SOCI- ETY ISLANDS: Island of Oheteroa, H. Cuming (Wilkins, 1951, p. 239); (also Schmeltz in Mus. Godefroy, 1874, cat. 5). AUSTRALIA: Ribbon Reef, Qld. ( T. Hartley in litt.). Fossil records— Ladd and Hoffmeister, 1945, p. 361 report a fossil which is closely allied to thersites from Lakemba Id., Lau Group (Fiji) in the Futuna limestone (Miocene or Pliocene). They erroneously likened it to the Red Sea S. tricornis. Strombus maximus Martin, 1883 Range— Upper Miocene of Java, Indonesia. Plate 31. Strombus maximus Martin. Java, Indonesia. Mio- cene. (from K. Martin, 1883, pi. 9, fig. 1). x Plate 32. Strombus maximus Martin. Upper Miocene of Java, Indonesia, (from K. Martin, 1899, pi. 28, fig. 407). V V. [09 - 874] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 57 Remarks— This massive, foot-long (270 mm.) Strombus is possibly the progenitor of the Recent Strombus ther sites. The apex is very similar to that of thersites, but is unlike that of S trombus gigas Linne of the West Indies as Martin had suggested. The dorsal side of the body whorl bears 3 or 4 large, pyramidal tubercles or spines, a feature ab- sent in thersites. Upper Miocene of Java (Tjilanang beds ) according to van der Vlerk, 1931, p. 247. Synonymy— 1883 Strombus maximus K. Martin, Samml. geol. Reichs- Mus. Leiden, 1st ser., vol. 1, p. 195, pi. 9, fig. 1 (Tertiary of Java: Gunung Sela and Tjidamar; Ba- tavia); 1899, Martin, ibid., n. Folge, vol. 1, pt. 1, p. 175, pis. 28, 29, figs. 407, 407a (Solo, Java). Strombus latissimus Linne, 1758 (PI. 17, fig. 5; pi. 33, fig. 1) Range— Ryukyu Islands to the Philippines and to Fiji. Remarks— This is the largest and heaviest of the Indo-Pacific Strombus, although it is smaller than either S. goliath Schroter, 1805 (Archiv. Zook Zoot., vol. 4, p. 139) of Rrazil or galeatus Swainson, 1823 from the Eastern Pacific. S. latissimus is consid- ered uncommon, perhaps because it lives well be- low the low tide line. It is readily recognized by its heaviness, by its broad, thick outer lip which sweeps up beyond the spire, and by its brownish outer coloring. It might be confused with S. thersites but the latter has a shorter lip, and its spire angle is about 45 degrees (instead of 70 to 75 degrees as in latissimus) . Habitat— Little is known of the habitat, although Philippine collectors have informed me that they must dive in 12 to 24 feet of water to collect it. Kira ( 1959, p. 37 ) reports that it lives from 5 to 10 fathoms in the Ryukyu Islands. Description— Adult shell from 120 to 204 mm. (5 to 8 inches ) in length, massive, heavy, rotund, with a broad, thick, flaring lip, yellow-brown to choco- late-brown in color, and with a smooth, tan and whitish aperture. Whorls 11. Nuclear whorls un- known. Angle of spire about 70 to 75 degree. Early whorls pinkish, strongly carinate and with numer- ous small knobs (22 to 17) on the periphery just above the indented suture. Spiral threads 8 to 12, but absent in later whorls. In the penultimate whorl these knobs become larger, rounded and fewer, rarely absent. Rody whorl with one very large, but low, rounded knob at the shoulder on the left side of the whorl not far from the slightly glazed parie- tal wall. Dorsum roundly swollen. Outer side of lip Plate 33. Fig. 1, immature S trombus latissimus Linne, Philippines. Fig. 2, Strombus sinuatus Humphrey, Okinawa Id., Ryukyu Islands. Fig. 3, immature Strombus taurus Reeve, Rongelap Atoll, Marshall Islands. All slightly re- duced. [09 - 875] 58 T ricornis R. T. Abbott Strombidae Plate, 34. Geographical distribution of Strombus latissimus Linne. with about 20 indistinct, whitish spiral ridges. Outer lip massive, spreading upward beyond tbe spire and partially obscuring the apex. Edge of lip thick and turned inward. Stromboid notch deep and pro- nounced. Columella thickened with a tan glaze and smooth. Aperture white and smooth within, becom- ing tan to smoky brown at the edges. Color of shell cream with dense mottlings of dark- or light- brown. In the penultimate whorl the dark-brown is arranged in irregular, crowded, axial streaks. Perio- stracum moderately thick, smoothish, translucent tan and Hakes off when dry. Operculum and soft parts unknown. Measurements (mm.)— length width no. whorls 200.0 147.2 8+ (large; Philippines) 160.2 110.0 11 (average; Okinawa) 111.1 91.0 8+ (small; Jolo Id., P. I Synonymy— 1758 Strombus latissimus Linne, Systema Naturae ed 10 p. 745, no. 436 (In O. Asiae); 1767, ed. 12, p. 1211, no. 505; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 272-273; 1843, Kiener, Coq. Vivantes, vol. 4, Strombus, pi. 4, fig. 2. 1798 Lambis latissimus Linne, Roding, Museum Boltenia- num, Hamburg, pt. 2, p. 65, no. 827. Refers to Conchyl.-Cab., vol. 3, figs. 832 and 835. 1798 Lambis picta Roding, loc. cit., p. 62, no. 793. Refers to Conchyl.-Cab., vol. 3, fig. 874 (an immature specimen ) . 1817 Pterocera alata Schumacher, Essai Nouv. Syst. Hab. Vers Testaces, Copenhagen, p. 221. Refers to Con- chyl.-Cab., vol. 3, figs. 832 and 835. Not Strombus alatus Gmelin, 1791. Types— Evidently Linnaeus did not possess a specimen when he first described the species. A young specimen, possibly to be considered a type, is in the Linnaean collection in Uppsala, Sweden. Since the description was based in part upon the fig- ure given by Rumphius (pi. 36, fig. L), we hereby select Amboina, Indonesia, as the type locality. Records (see accompanying map, pi. 34)— RYUKYU IS- LANDS: Amami Id. (Kira, 1959, p. 37); Okinawa Id. (Mrs. A. A. Scott, ANSP). TAIWAN: Kasyo-to (Kuroda, 1941, p. 97). PHILIPPINES: Borongan village, east side of Samar Id. (ANSP); near Cebu City, Cebu Id. (MCZ); Lubang Id., Mindoro (P. de Mesa, MCZ); Zamboanga, Mindanao Id. (ANSP); Davao Bay, Mindanao; Cuyo Id., Palawan (du Pont- Academy Exped., 1958, ANSP); Jolo Id., Sulu Archipelago (ANSP). INDONESIA: Amboina (Rumphius, 1741, p. 110). NEW CALEDONIA: Touho Bay ( M. Leveque, ANSP). FIJI: reefs off Levuka, Ovalau Id. (MCZ); Kamba Point, Viti Levu Id. (H. S. Ladd, USNM ). Fossil records— None reported. Strombus taurus Reeve, 1857 (PI. 17, figs. 3, 4; pi. 33, fig. 3) Range— Known only from the Marshall and Mari- anas Islands (and possibly the Admiralty Islands). Remarks— For nearly a hundred years this species was known from only two or three specimens. It is now known to be not uncommon in the Marshall Islands thanks to the keen observations and careful collecting being done by such men as Richard C. Willis and John Roberts, Jr. It probably occurs in the Admiralty Islands and other nearby areas. Strombus taurus is closely related to the thersites- latissimus-sinuatus complex. It has a four-inch-long, heavy shell which is characterized by one long and one short spine at the top of the thickened, wavy outer lip and by the lavender-purple blotch deep within the tan and white aperture. The young of taurus are distinguished by a nar- row, bright lavender or rose band just below the suture. Habitat— Willis and Roberts have kindly supplied the following interesting account: “It occurs in fairly large numbers at Eniaro and Gogon Islands, Rongelap Atoll, at a depth from 4 to 15 feet on a bottom of broken rubble of coral, fine sand and brown furry algae. Specimens were always found in pairs and where the tidal currents form terraces on the north and northeast islands of the atoll. Nearby there were usually other species of Strom- bus ( lentiginosus ) and Lambis ( truncata and chiragra ), although the taurus seem to congregate together. The majority of several dozen specimens collected were pockmarked by erosion, lime de- [09 - 876] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 59 posits, and tube worm \V ennetus snails] growths. H. T. Ward ( 1960, p. 1 ) collected a live pair in 80 feet of water in Guam, Marianas, on a hard rock and algae bottom and in company with Lambis truncate. Description— Shell, excluding spine, 80 to 102 mm. (3 to 4 inches) in length. Spine on outer lip adds another 5 to 25 mm. Heavy, massive, rudely knobbed, with one long and one short spine at the top of the outer lip, and a brownish purple splotch deep within the white and tan aperture. Outer shell creamy white with yellow-brown maculations. In the young there are two spiral bands of alternating brown and white, irregular spots near the mid- portion of the base of the shell, and a narrow, white-speckled band of bright violet or lavender just below the finely indented suture. Whorls 10 to 11, the apex usually eroded away. Nuclear whorls 2, smooth and translucent whitish or pinkish. First two postnuelear whorls with numerous, crowded axial riblets crossed by about a dozen fine spiral threads. The latter persist weakly to the penulti- mate whorl. By the third whorl small knobs appear just above the suture ( 10 to 15 per whorl ) and con- tinue to the last whorl. Dorsum of body whorl at the shoulder bears two large, pyramidal knobs and anteriorly a swollen ridge. Below the smaller knob and on the middle of the dorsum is a peculiar, large, elongate, irregularly formed and obliquely placed knob. Parietal shield glazed, very swollen below, and colored yellow-brown. Columella whit- ish above, lavender-tan below with 1 to 3 weak spiral white teeth at the base. In old specimens, the parietal shield runs up on to the spire and is edged above with 1 or 2 short finger-like blades. Aperture constricted, brownish purple to lavender deep within, white further out, and finally soft brown to tan on the outer lip. Deep within the aperture on the upper end and on the body wall is a thickened, rounded, spiral ridge. Outer lip thick, very wavy, glazed with light-brown to creamy tan. Above are two well-developed spines pointing upward, the innermost being twice as long as the outer one. Siphonal canal moderately short, purplish within, and slightly recurved, but not twisted. Stromboid notch very deep and well-developed; below it is a thick, protruding, glazed flange which may have 2 or 3 weakly developed, whitish teeth. Periostracum thin, smoothish, translucent tan, but usually worn away in adults. Operculum stromboid, light-brown, arching, with no or badly worn serrations. Radula ribbon reddish brown, 10 mm. in length, with about 45 to 50 rows of teeth. Formula: 2-1-2; 1-3 (no peg); 6; 7. Verge simple, red-brown with cream maculations. Tentacles brown with white Hecks (preserved specimen). Measurements (mm.)— (length excludes spines) length width no. whorls 80.0 76.2 6 + ( small; Rongelap Atoll ) 89.0 70.1 10 ( average; Rongelap Atoll 101.0 77.1 7 + ( large; Rongelap Atoll ) Synonymy— 1857 S trombus taurus Reeve, Proc. Zool. Soc. London, for 1857, p. 207, pi. 37, fig. 3 (in color) (“Amirante Islands, a group of the Seychelles”); 1885, Tryon, Manual Conch., Philadelphia, vol. 7, p. Ill (Ad- miralty Islands); 1960, Ward, Hawaiian Shell News, vol. 8, no. 4, p. 1 (Guam Id.). Types— The type which was in Sir David Bar- clay’s collection was sold in 1891 and is probably in private hands. It has not been found in the British Museum. The type locality is “Amirante Islands” in the western Indian Ocean. This may be an error and I suspect the original label read “Admiralty Islands” which are close to the known range of this Micronesian species. Tryon who corresponded with Sir David Barclay changed the locality in the Manual of Conchology to “Admiralty Islands”. Un- fortunately, I have been unable to find any refer- ence to this in Barclay’s letters, but I suspect the change was made with cause. Records— MARIANAS: Asan Point, Guam Id. (Mrs. D. L. Hiatt, in litt. ); north side of breakwater, Pier Area, S.W. Tinian Id. (A. R. Bronson, MCZ). MARSHALL ISLANDS: Rongelap Atoll: Eniaro Id. (Willis and Roberts); Kabelle Id., 20 feet (Dick Willis, ANSP); Gogon Id. (Dick Willis); Kieshieehi Id., Arbar Id., and Eniaetok Id. (USNM). Bikini Atoll: J mi. S.W. of S.E. Point, Bikini Id.; 4 miles south, Bikini Id., 25 fathoms, dead; Chieerete Id. ( all J. P. E. Morrison, USNM). ADMIRALTY ISLANDS: (Tryon, 1885, p. 111). [Amirante Islands, Indian Ocean (Reeve, 1857) may be erroneous.] Fossil records— None reported. Plate 35. Strombus taurus Reeve, Marshall Islands. Fig. 1, central radular tooth. 2, lateral. 3, inner marginal. 4, outer marginal. 5, verge. [09 - 877] 60 Tricornis R. T. Abbott Strombidae Strombus sinuatus Humphrey, 1786 (PI. 17, fig. 8; pi. 33, fig. 2) Range— Ryukyu Islands to Australia, Micronesia and Melanesia. Remarks— This handsome species is readily recog- nized by its brownish purple aperture and the 3 or 4 thin, finger-like blades at the top of the outer lip. It is uncommon to moderately common throughout its range. In the Bohol-Cebu area of the Philippines it appears in shallow water from April to May in large numbers, some several thousand specimens having been collected commercially in 1957. This species, despite its peculiar Lambis- like lip, shows close affinities to latissimus, thersites and taurus. Habitat— This species lives on coral sand and algae bottoms from low tide mark to a depth of 10 fathoms. In the Marshalls it occurs both in the la- goons and outside. Mr. Richard C. Willis and Air. John Roberts, Jr. have very kindly supplied the fol- lowing notes on the occurrence of sinuatus on Enieatok Island, Rongelap Atoll, Marshall Islands. “(In September, 1959) shells not plentiful with one or two every 200 feet. Water 4 to 20 feet in depth, clear to slightly murky, temperature 85° F. Bottom of loose broken coral and algae, the latter also growing on the shells. In fresh specimens, the aper- ture is a delicate lavender, overlaid with radiating streaks of orange in young specimens.” Description— Shell 82 to 113 mm. ( 3M to 4)2 inches) in length, solid, moderately heavy, with a brownish purple aperture and 3 or 4 finger-like blades at the top of the outer lip. Color of outer shell whitish with heavy mottlings of yellow-brown, and usually with 3 to 5 narrow spiral bands of alter- nating white and brown Hecks on the body whorl. Whorls 10 to 11. Nuclear whorls 2, smooth, trans- lucent tan to pinkish. First 2 postnuclear whorls with numerous, crowded axial riblets running from suture to suture and crossed by 15 to 20 fine, but strongly raised, spiral threads. The latter persist in strength to the penultimate whorl and cross the knobs and 15 to 20 swollen whitish, former varices on the spire. The remaining whorls are well shoul- dered with neat, rounded, spirally threaded knobs (about 9 to 11 per whorl). Above this row of knobs the whorl is concave. The last whorl has an indis- tinct, low, long knob on the left side, thus giving the parietal wall a flattish appearance. Anteriorly, are 2 small knobs, followed by a large pyramidal one on the dorsum, and finally by a last and smaller one. Below this series of large shoulder knobs, are 2 spiral rows of 2 to 4 small, round, beads. Parietal wall slightly glazed. Columella moderately thick- Plate 36. Geographical distribution of Strombus sinuatus Humphrey. Open circles indicate literature records; solid dots, specimens examined. ened, smooth and brownish-tan. Deep interior of aperture brownish purple to deep purple, blending into soft-brown towards the smooth edge of the outer lip. Middle section of outer lip incurved, glossy, and with brownish color bands. Upper end of lip adnate to the spire and having 3 to 5 ( usually 4) thin, whitish, tongue-like blades. Siphonal canal short, but twisted to the right. Stromboid notch deep and flaring. The flange below it may have 4 to 5 small white teeth. Periostracum moderately thin, semi-glossy, translucent yellowish and apt to flake off when dry. Operculum stromboid, strongly arched and with weak serrations. Soft parts unknown. Measurements (mm.)— length width no. whorls 113.5 69.9 10 + (large; Cebu Id.) 88.8 55.4 10 ( average; Bohol Id. ) 82.5 53.8 7 + ( small; Solomon Ids. Synonymy— 1786 Strombus sinuatus Humphrey, Portland Catalogue, London, p. 189, no. 4022 ( no locality ) ; refers to Seba, vol. 3, pi. 62, fig. 3 and Argenville [Favanne], pi. 22, fig. A2. 1798 Lambis lobata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 65, no. 826. Refers to Conchyl.-Cab., vol. 10, figs. 1506, 1507. Not lobata Roding, loc. cit., p. 68, no. 872. 1807 Pterocera palmata G. Fischer, Museum Demidoff, Moscow, p. 191 (no locality). 1817 Strombus laciniatus “ Chemnitz” Dillwyn, Descr. Cat. Recent Shells, London, vol. 2, p. 663 ( East Indian Seas). Refers to Conchyl.-Cab., vol. 10, figs. 1506, 1507; 1850, Reeve, Conch. Icon., vol. 6, Strombus, pi. if, fig. and sp. 25. [09 - 878] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 61 1822 Strombus cristatus Lamarck, Anim. sans Vert., vol. 7, p. 202 (no locality). Refers to Seba and Favanne ( loc . cit.), and Conchyl.-Cab., vol. 10, figs. 1506, 1507; 1843, Kiener, Coquilles Vivantes, vol. 4, Strombus, pi. 11. Types— The whereabouts of Humphrey’s type of sinuatus is unknown to us. ft was sold at the auc- tion of the shell collection of the Duchess of Port- land in London in 1786. No locality was given. We designate Cebn City, Cebu Island, Philippines as the type locality. The type of Pterocera palmate G. Fischer, if not destroyed by fire, may be in the zoo- logical museum of the University of Moscow ( see K. V. W. Palmer, Nautilus, vol. 70, no. 1, 1956). Nomenclature— The author of sinuatus is given as Humphrey by some workers, and as Solander by others. Either refers to the Portland Catalogue. Records (see accompanying map, pi. 36)— RYUKYU IS- LANDS: Tanegashima Id. (Kuroda in MSS); Yaeyama Id. (A. R. Calm Colin., ANSP); Okinawa Id. (Mrs. A. A. Scott, ANSP). TAIWAN: Karenko (Kuroda, 1941, p. 98). PHILIPPINES: near Cebu City, Cebu Id. (A. B. Franco, ANSP); Badang, Sorsogon Prov., Luzon Id. (du Pont- Acad- emy Exped., 1958. ANSP); Panglao Id., northern Bohol Id. (E.' Zambo, ANSP); Surigao Id. ( Elera, 1896, p. 249); Siasi, Sulu Archipelago ( W. E. Old, Jr., coll n. ). INDO- NESIA: Amboina, Moluccas (MCZ); Sailus Ketjil, Paster- noster Ids., off Sumbawa Id. ( Schepman, 1909, p. 148). AUSTRALIA: reefs off Townsville Queensland ( fide Tony Marsh in lift.); Northwest Cape, on reef. Western Australia (A. Whitworth, coll.). NEW GUINEA: Mios Woendi Atoll. Padaido Ids. ( NSF, 1956). SOLOMONS: Choiseul Id. (W. J. Eyerdam, ANSP). NEW CALEDONIA: (USNM); Touho ( fide D. Getz, in lift.). FIJI: reefs off Levuka, Ovalnu Id. (MCZ); Kamba Point, Viti Levu Id. ( H. S. Ladd, USNM). MARIANAS: Guam Ld. (fide T. Montgomery, in litt.). CAROLINES: Hall Ids., near Truk Id. (Mrs. R. T. Callemore). MARSHALLS: lagoon, Aomaen Id., and Namu Id., and Bokororyuru Id., all Bikini (J. P. E. Morrison, USNM); Eniaetok, Rongelap Atoll (LtSNM). [No vtlienti- cted Hawaiian records, as yet.] Fossil records— None reported. Strombus junghuhni Martin, 1879 Range— Miocene of Java, Indonesia. Remarks— This species was based upon a single and broken specimen. From the illustration of the type, it would appear that this is the closest repre- sentative of Strombus sinuatus that has been found in the Tertiary. The absence of details of the outer lip, however, leaves this relationship still in doubt, although the characters of the spire are fairlv close. This is Miocene according to van der Vlerk, 1931, p. 246. Synonymy— 1879 Strombus junghuhni K. Martin, Die Tertiiirschichten auf Java, Leiden, p. 47, pi. 9, fig. 1 (Java, Tertiary). Strombus mekranicus Vredenburg, 1928 Range— Lower Miocene of West Pakistan. Plate 37. Strombus junghuhni Martin. Miocene of Java, In- donesia. Type, (from K. Martin, 1879, pi. 9, fig. 1). Natural size. Remarks— l concur with Vredenburg’s remarks that this species “is nett very closely related to any living or fossil species, though it belongs to the same group that includes such forms as the recent Strombus bubonius Lamk. [= lotus Gmelin] and Strombus tricornis Lamk. [Humphrey].’’ It is based upon very poorly preserved specimens. Synonymy- 1928 Strombus mekranicus Vredenburg, Memoirs Geol. Sur- vey of India, vol. 50, pt. 1, p. 315, pi. 3, figs. 13, 14, pi. 4, fig. 1 (Mekran beds: north of Talar Gorge, on the road from Kej to Gwadar, base of the sand- stones constituting the Talar Mountains [West Paki- stan]. Plate 38. Strombus mekranicus Vrendenburg. Mekran, Ta- lar, Pakistan. Miocene, (from Vredenburg, 1928, pi. 3, figs. 13, 14). Natural size. [09 - 879] 62 Tricornis R. T. Abbott Strombidae Strombus inflatus Martin, 1879 Range— Lower Miocene and Upper Miocene of Java, Indonesia. Remarks— Strombus herklotsi Martin 1880 and Plate 39. Figs. 1 and 2, S trombus inflatus Martin. Tji Longan, Java, Indonesia. Miocene. ( from K. Martin, 1883, pi. 9, fig. 3). Figs. 3 and 4, S. tjilonganensis K. Martin. Java, Miocene, (from Martin, 1899, pi. 28, figs. 410, 410a, type). All natural size. Plate 40. Strombus tuberosus Martin. Tji Longan, Java, In- donesia. Miocene, (from K. Martin, 1883, pi. 9, fig. 2). Nat- ural size. tuberose Martin 1883 from the same locality in Java are possibly synonyms. I know of no Recent Indo- Pacific species resembling it, although Strombus costatus Gmelin, 1791 of the West Indies is some- what similar. Synonymy— 1879 Strombus inflatu-s K. Martin, Die Tertiarschiehten auf Java, Leiden, p. 48, pi. 9, fig. 3 (Java, Tertiary); 1883, Martin, Samml. geol. Reichs-Mus. Leiden, 1st series, vol. 1, p. 197, pi. 9, fig. 3. 1883 Strombus tuberosus K. Martin, Samml. geol. Reichs- Mus. Leiden, 1st ser., vol. 1, p. 196, pi. 9, fig. 2 (Tji Longan near Selatjau, Java, Tertiary); 1921, K. Martin, loc. cit.. n. Folge, vol. 1, pt. 2, p. 468, pi. 60, figs. 57, 58 (Tji Talahab and Tji Angsana, Java, Pliocene). [Lower Miocene: Njalindoeng beds and upper Miocene: Tjilanang beds, fide van der Vlerk, 1931, p. 247.] 1899 Strombus (s. str.) herklotsi K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 178, pi. 29, figs. 413, 414 (Java). Strombus tjilonganensis Martin, 1899 Range— Upper Miocene of Java, Indonesia. Remarks— Allied to inflatus Martin, above, and possibly a form of it. Synonymy- 1899 Strombus (s. str.) tjilonganensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 177, pi. 28, figs. 410, 411, pi. 29, fig. 412 (Selatjau, Java, Pliocene). Subgenus Dilatilabrum Cossmann, 1904 Type: Strombus fortisi Rrongniart, 1823 Remarks— To our knowledge, this subgenus does not occur in the recent or fossil record of the Indo- Pacific. The subgenus contains two or three species known only from the Italian Eocene of Europe. The shells are large, heavy, with a large flaring lip and presumably with a poorly developed or obsolete stromboid notch. The shells closely resemble some members of the subgenus Tricornis, and Cossmann himself felt it might not be worth recognizing. We have not seen specimens of this subgenus, and can give no opinion as to its relationships or its age. If a Strombus, it is the earliest known record for the genus. Synonymy— 1876 Oncoma Mayer-Eymar, Reitr. geol. Karte Schweiz, no. 14 (2b), p. 57. Non Fieber, 1861. [Not seen.] 1904 Dilatilabrum Cossmann, Essais de Paleoconchologie Comparee, Paris, 6th book, p. 12. Type: Strombus fortisi Brongn. of the Eocene of Europe. New name for Oncoma Mayer-Eymar, non Fieber, 1861. [09 - 880] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 63 Subgenus Canarium Schumacher, 1817 Type: Strombus urceus Linne, 1758 The 18 living subspecies and species in this sub- genus are limited to the Indo-Pacific. Several, like mutabilis Swainson, are very widely distributed, while others, like helli Kiener and klineorum are very restricted in range. Although some of the spe- cies appear to fall into more or less natural groups on mere conchological grounds, such as into the urceus-labiatus-erythrinus group or the mutabilis- microurceus-maculatus group, we are combining a rather wide selection of species because of concho- logical intergrades and very similar anatomical fea- tures. Despite the slender and smooth shell of terebellatus, we believe it has no close phylogenetic relationship with the genus Terebellum. Species like fragilis bridge the gap from terebellatus to mutabilis and, thence, to urceus. The shells in this subgenus are rather small and without a strongly flaring or winged outer lip. The penis is a simple prong with a slightly swollen dis- tal portion. The radula consists of delicate teeth with the central bearing 5 cusps ( the middle one being the largest); the lateral usually has only 3 or 4 (rarely 5) cusps and a small basal peg. The oper- culum has 5 to 9 well-developed serrations. A number of fossil species occurs in the Miocene and Pliocene of southeast Asia and the East Indies. The most ancient of these appear to be related to mutabilis and labiatus. Nothing resembling dentatus or fusiformis occurs in the Pliocene or earlier. Strombus liocyclus Dali, 1915, from the Silex beds of Tampa, Florida [Miocene] quite likely belongs to this subgenus, and is the only recorded Carib- bean representative. Synonymy— 1817 Canarium Schumacher, Essai Nouveau Systeme, Copen- hagen, p. 219. Type by monotypy: Canarium ustu- latum Schumacher = Martini, vol. 3, figs. 803, 805 = urceus Linne, 1758. 1840 Strombidea Swainson, A Treatise on Malacology, Lon- don, pp. 138 and 140. Type by monotypy: urceus Linne, 1758. 1888 Conorium Jousseaume, Mem. Soc. Zool. de France, vol. 1, p. 174. Type by monotypy: Conorium mu- tabilis Swainson [error for Canarium?]. Strombus urceus Linne, 1758 (PI. 20, figs. 26-29; pi. 41, figs. 1-6) Range— Southeast Asia and the Ryukyu Islands to Australia and Melanesia. Remarks— This species is close in appearance to labiatus (Roding), but is distinguished from the latter by its more drawn out siphonal canal, which is generally dark blue-black within and without, and by the smooth central portion of the usually white, rarely black, and rarely orange columella. The two species do not live in the same immediate habitat (see under labiatus). Our interpretation of what Linnaeus meant by urceus is discussed under Types and Nomenclature below. S. urceus is a very variable species. In addition to size, sculptural and color variations that appear within a single colony, there are other geograph- ical dines and groups of morphological variations Plate 41. Strombus urceus Linne. Figs. 1 and 2, Mindanao form. 5, immature of S. urceus orrac Abbott, paratype, Id., Philippines. 3, Geelvinck Bay form, Dutch New Guinea. Broome, Western Australia. All x 1.5. 4, shouldered form, Aitape, New Guinea. 6, Yap Island [09- 891] 64 Canarium R. T. Abbott Strombidae limited to certain rather discrete geographical areas. Some of these latter cases may well be considered subspecies; others may be due to environmental conditions common to these circumscribed areas. We have refrained from breaking the species np into numerous subspecies because we feel that many more large samples are needed from many more localities. Experimental work is also sorely needed. The most distinct of the variations, with a fairly well-defined geographical distribution, is the heavy, high-spired, few-knobbed, square-lipped form so common along the northern and especially north- western portion of the Australian continent. This we accepted as a valid subspecies and name it Strombus urcens orrae. Dominant in the Singapore area, and decreas- ingly so northward into the Gulf of Siam to Hong Kong, and sparingly in the Ryukyus and only sporadically in the central Philippines is the form with a black-colored columella and aperture: form ustulatus Schumacher, 1817 (see pi. 20, fig. 29). This dark-mouthed character is probably an ex- ample of a single gene cline, and not due to en- vironmental conditions. In Tabaco Bay, Albay Province, Luzon Island, an area where many species are particularly darkly colored ( Conus eburneus form poly glottis Weinkauff ), specimens of Strombus urceus do not have darkly pigmented apertures. The deeper water form ( 1 to 25 fathoms ) of urceus intergrades with the larger more robust intertidal form. It ranges in size from 18 to 26 mm. in length, is usually quite elongate, light weight and is sparsely and weakly colored. The blue-black on the siphonal canal may be greatly reduced or absent. An interesting form is dominant in the north- eastern waters of New Guinea which, when sub- stantiated with samples from surrounding areas, may prove to be a distinct subspecies. We refer to this as the “Geelvink form. The lower third of the last whorl is solid blue-black. The body whorl is smooth, rounded at the shoulder and bears only 2 or 3 very weak nodules. The parietal wall is smooth, glistening and appearing to be light-tan, but under a lens is seen to be a beautiful, fine, zigzag, axial pattern of delicate orange-brown. The spire is finely beaded and bluish black. We have also seen the form from the Palaus and Ponape Island in the Carolines (see pi. 41, fig. 3). An additional and evidently closely related form appears to have received the name incisus Wood, 1828. We have this form from the Solomons and from three other localities whose authenticity of data is questionable— Majuro Atoll, Saipan and a large series from “Fiji." It may subsequently be considered a subspecies when more is known about its anatomy and range. In shape, the shell is quad- rate, with a broad shoulder bearing 6 to 7 short, strong, axial, slanting ribs and a very swollen, squarish, upper end to the outer lip which is ele- vated to the height of the suture above. The poste- rior canal is in the form of a minute channel or “knife-cut" located towards the edge of the outer lip. The flange below the stromboid notch is very broad and large and may extend as far forward as the tip of the siphonal canal. The color of the shell is whitish with the lower fourth of the body whorl blue-black to brownish black and the dorsum with 2 or 3 diffused bands of lead to brownish gray. The spire is minutely beaded, the apex blue-black to brownish. Shell length 19 to 30 mm. (see pi. 41, fig. 4). Habitat— Occurs in colonies on sand or sandy mud bottom from the low tide mark to 20 fathoms. It is sometimes associated with sparse algal bot- toms, and is not found intermingled with Strombus labiatus (Roding). Description— Adult shell 19 to 61 mm. (usually about 50 mm.) in length, solid, smooth to axially nodulated, vari-colored, and with an all white, all black, black-rimmed or rarely yellowish-orange columella which is always smooth, except at the top and bottom fourth. Color of outer shell very variable, ranging from brown, whitish, cream, or with greenish, yellow-brown maculations, flecks, broken bands and axial streaks to a solid pink, orange or yellow. Siphonal canal almost always tipped with bluish black within and without. Whorls either rounded or, especially in the spire, squarish. Nuclear whorls 3, bulimoid, glossy, smooth translucent tan or bluish black, rarely colorless. First postnuclear whorl with about 8 to 10 micro- scopic, incised lines. Subsequent whorls become shouldered and with 11 to 17 small axial, usually knobbed or beaded, riblets per whorl. Last whorl with 3 to 8 low knobs at the shoulder, the last 2 or 3 on the dorsum being the largest. Axial plications are almost always absent on the parietal area. Base of shell with 8 to 10 incised lines or low, flat, broad cords, the latter never raised, rounded or strong. Siphonal canal somewhat elongate, slightly reflected upward and moderately twisted. Columella smooth, except for 6 to 15 raised spiral lirae at the top third and 1 to 8 distinct lirae at the lower fourth and set almost at right angle to the axis of the columella. [09 - 892] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 65 Interior of body whorl with about 50 to 70 fine, purple-brown spiral lirae which may or may not ex- tend to the edge of the outer lip. Deep interior of aperture yellowish white or pure blackish or blushed with purple-brown. Columella either all white, all black, yellowish or orangish or combina- tions of these colors. Stromboid notch shallow to deep. Periostracum thin, smoothish, translucent grayish to yellowish, and usually worn away on the body whorl. Operculum stromboid, one third the length of the shell, not arching, brown, and with about 12 fine, sharp serrations. Radula ribbon 4 mm. in length, with about 38 rows of teeth. Formula: 2-1-2; 1-3 (plus peg) or rarely 1-2 (with peg); 4 or 5; 5 or 6. Verge simple, variegated in color and with white and red em- bedded granules. Posterior mantle filament 3 mm. Edge of mantle with a series of red lines. Eye with 1 red ring. Body variegated in color giving it a marble-like effect. Measurements (mm.)— length width no. whorls 61.5 23.9 10 ( giant; Singapore ) 56.4 24.1 9 + (large; Balabac Id.) 48.5 21.6 9 + ( average; Cebu Id. ) 32.3 14.7 10 ( small; Luzon Id. ) 19.0 17.8 8 ( dwarf; deep water) form incisus Wood 31.0 17.1 8 (large; Fiji) 22.7 12.0 8 (average; Fiji) 18.0 9.5 8 (small; Fiji) Synonymy— 1758 S trombus urceus Linne, Systema Naturae, ed. 10, p. 745, no. 440 (In O. Asiae); 1767, ed. 12, p. 1212, no. 512; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 284-285; 1842, Sowerby, The- saurus Conchyl., vol. 1, pi. 7, figs. 34-37, 41, 42; 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 15, fig. 2, pi. 30, figs. 2, 3; 1844, Duclos, in Chenu’s Illustr. Conchyl., vol. 4, pi. 5, figs. 8, 9, 13-15, pi. 24, figs. 5-6. ?1807 Lambis reticulata Link, Besehr. Natur.-Samml., Univ. Rostok, pt. 2, p. 109. Refers to Conchyl. -Cab., vol. 3, fig. 806. 1817 Canarium ustulatum Schumacher, Essai Nouv. Syst.. p. 219. Refers to Conchyl. -Cab., vol. 3, figs. 803. 805. 1828 S trombus incisus Wood, Index Testaceol. Supplement, London, p. 14, pi. 4, fig. 12 (no locality). [Quad- rate form.] 1844 S trombus anatellus Duclos, in Chenu’s Illustr. Con- chyl., vol. 2, pi. 4, figs. 11, 12, pi. 21, figs. 8, 9 (Japan). [Smooth-shouldered form.] ?1839 Strombus crassilabrum Anton, Verzeichniss Con- chylien, Halle, p. 87, no. 2820 (no locality). 1885 Strombus (Canarium) muricatus “Martini”, Watson, Report . . . H.H.S. Challenger, vol. 15, p. 417; 1909, Schepman, in Weber’s Siboga-Expeditie, Leiden, vol. 28, pt. 2, p. 151. 1938 Strombus (Canarium) plicatus Lamarck, Adam and Leloup, Mem. Mus. Royal d’Hist. Nat. Belgique, special series, vol. 2, fasc. 19, p. 112, pi. 1, figs. 8d and e only (non Lamarck). 1946 Strombus ustulatus form laevis Dodge, Amer. Mus. Novitates, N. Y., no. 1314, pp. 2 and 7, figs. 1 and 6. ?1946 Strombus (Canarium) gendinganensis Martin, Abrard, Annales de Paleontologie, Paris, vol. 32, p. 61, pi. 4, fig. 27 (Pliocene, New Hebrides). Types and Nomenclature— No species of Linnaeus has undergone more alternating interpretations by various authors than urceus. Linnaeus’ description could fit any of three species, and his figure refer- ences could be either of two species. The specimen, marked for this species in the Linnaean collection, has never been illustrated, although Hanley ( 1855, p. 275) likened it to Sowerby’s Thesaurus Conch., vol. 1, pi. 7, fig. 45 which is what we call mutabilis Swainson. I cannot help but believe that this speci- men was marked as urceus at a later date by some- one other than Linnaeus. This was done in the case of other species, according to Dodge, Hanley, and 2 Plate 42. Fig. 1, radula of Strombus urceus urceus Linne and. fig. 2, its verge, Palau Islands. 3, verge of S. erythrinus erythrinus Dillwyn and, fig. 4, its central radular tooth, New Caledonia. 5, verge of S. belli Kiener, Oahu Island. 6, oper- culum of S. fragilis (Roding), Palau Islands. 7, operculum of S. dentatus Linne, Zanzibar. [09 - 893] 66 Canarium R. T. Abbott S trombidae Plate 43. Geographical distribution of Stwrnbus klineorum Abbott and Strombus urceus urcens Linne and its Australian subspecies, orrae Abbott. others. None of Linnaeus’ figure references are to mutabilis-\ike shells, despite the fact that perfectly good pictures of it occur in the works referred to by Linnaeus (Rumphius, pi. 37, fig. W; Seba, pi. 62, figs. 42, 43). However, Linnaeus’ figure refer- ences (Rumphius, pi. 37, fig. T; Petiver, pi. 98, fig. 19; Gualtieri, pi. 32, fig. G; and in the 12th edition, Seba, pi. 60, figs. 28, 29 and pi. 62, figs. 45, 41, and 47 ) are all dorsal views and match specimens before us of either of two species which depend upon good apertural views for identification. No amount of arguing is going to settle the point on the basis of the figures. We hereby select shell no. 300 which is illustrated on the microfilm of the Linnaean types in the Museum Ludovicae Ulricae (M. L. U. 288) as the neoholotype or lectotype. This conforms with the urceus concept of Lamarck, Deshayes, Sowerby (Thes. Conch., vol. 1, pi. 7, figs. 34-37), Reeve (Conch. Icon., pi. 11, fig. 24b), Tryon and Dodge. Kiener’s ( 1843 ) best figures are pi. 30, fig. 2, and pi. 31, fig- lb. We designate Cebu Island, Philip- pines, as the type locality. With so much confusion attending the use of the name urceus, it is impossible to know in most cases what certain later authors had in mind, unless they illustrated their shells. For this reason we have not used literature information in working out the dis- tribution of urceus. The smooth shouldered form turns up sporadi- cally as probably a minor genetic form in colonies in the Philippines, East Indies, Melanesia and Singa- pore. It is of no taxonomic significance and has had two names given to it: form anatellus Duclos, 1844 and form laevis Dodge, 1946. Selected records ( see accompanying map, pi. 43. Solid dots: specimens examined; no literature records used) — HONG KONG: Rocky Harbour, Tai She Wan (A. J. Staple, ANSP). THAILAND: Bandon Bight; Koh Chang; Bangbert Bay; Hualpa Id.; Koh Samet; Koh Samui; Koh Tao (all USNM). SINGAPORE: Tanah Merab Besar ( R. D. Purchon, ANSP). RYUKYU ISLANDS: Yaka Beach, Ishikawa, Oki- nawa Id. (USNM). PHILIPPINES: common throughout the islands: Luzon, Mindoro, Lubang, Catanduanes, Negros, Marinduque, Cuyo, Cebu, Bohol, Samar, Masbate, Cami- guin, Palawan, Mindanao, Basilan, Tawi Tawi, Jolo, Bala- bac, Busuanga, Burias, and Sanga Sanga Island (all USNM, ANSP, or MCZ). DUTCH NEW GUINEA: Geelvink Bay: off Rowo Id., Aeori Ids., 1 fm.; Japen Id., Ambai Ids., 8 fms.; (all NSF). PALAUS: South side of Malakal Harbor, 2 fms.; reef, Karamando Bay, Babeltlmap Id. (all NSF). CAROLINES: Ponape (V. Wertley, ANSP); Yap Id. (C .O. Kile, ANSP); reef at Mutunlik, Kusaie Id. (USNM). SOLO- MONS: Pavuvu Id., Russell Ids. (USNM); Florida Id. (MCZ); Sun, Malaiti Id. (W. J. Eyerdam, ANSP). [Unsub- stantiated records: Fiji; Saipan; Majuro Atoll, Marshalls; Tuticorn, India (USNM).] Fossil records— No bona fide records with the possible ex- ception of Abrard’s 1946 Pliocene record from the New Hebrides. The shell figured looks like true urceus in shape, and the columella appears to be smooth. If truly an urceus. it indicates that the eastern limits of this species have drifted westward since the Pliocene. Strombus urceus subspecies orrae new subspecies (PI. 20, fig. 28; pi. 41, fig. 5) Range— Western Australia to the Gulf of Carpen- taria, Australia. Remarks and description— This new subspecies is characterized and distinguished from the southwest Pacific typical urceus by its heavier, more quadrate, less colorful shell, its stronger and fewer nodules and its shorter siphonal canal. Its penultimate whorl bears 7 to 11 knobs (instead of as many as 10 to 16 as in typical urceus). The top end of the outer lip is square and thickened; the columella always white and the inner aperture yellow with sparse brown lirae. Body whorl with 7 to 9 nodules at the shoulder, the first one or two on the dorsum being large, swollen and prominent. The young show 3 narrow, broken spiral bands of brownish black on the body whorl. Stromboid notch shallow. Nuclear whorls 2%, translucent yellowish and glossy. I take pleasure in naming this subspecies after Miss Virginia Orr, Assistant, Department of Mol- lusks at the Academy of Natural Sciences, who col- lected this subspecies in Australia. Measurements (mm.)— length width no. whorls 42.1 20.8 9 ( holotype, ANSP ) 47.8 21.1 10 ( para type, ANSP ) 21.2 9.5 9 ( paratype, ANSP) Habitat— Intertidal on sand flats. Synonymy— No previously published names or [09 - 894] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 67 figures known. Some references to Australian urccus Linne may be this subspecies. Types— Holotype in ANSP no. 247756. The type locality is Augustus Island, Western Australia, Aus- tralia. Collected by B. Bardwell. Paratypes in MCZ and ANSP nos. 232673, 233409, 240131, 240137. Locality records (and other paratypes)— WESTERN AUS- TRALIA: Broome (MCZ); Gantheaume Point, Broome; La Grange Bay (both V. Orr, 1958, ANSP); Augustus Id. (ANSP and MCZ). NORTHERN TERRITORY: Cape Le- veque (MCZ); Shell Id., and East Point, and Port Darwin, Darwin (all MCZ); Quail Id., 35 mi. west of Darwin (MCZ); Nightcliff Point, Darwin (USNM). Allaru Islet, west of Port Essington, Cobourg Peninsula (MCZ). GULF OF CARPENTARIA: Groot Eylandt and Bickerton Id. ( USNM ) . Fossil record, s— None reported. Strombus labiatus (Roding, 1798) This well-known species has had a remarkably complicated nomenelatorial history. Various authors have treated it as a form or as a subspecies of urceus Linne. Our own studies lead us to conclude that it is a discrete species having two subspecies— the typical labiatus from southeast Asia and the southwest Pacific— and the olydius Dnclos from the western part of the Indian Ocean. The chief differ- entiating characters of these two subspecies are: S. labiatus labiatus— columella yellowish- to red- dish-orange with fine, slightly raised, spiral lines of darker color. The lirae on the inner wall of the body whorl are usually darkly colored. S. labiatus olydius— columella light-mauve or whitish violet with rather wide, usually raised, spiral fines of white. The lirae on the mauve inner wall of the body whorl are usually white. The species appears to be confined to large vol- canic islands. Only one specimen of labiatus labiatus is known from the Micronesian islands, that being a live specimen taken in 1955 by Mr. Ostheimer on Babeltlmap Island, Palau Islands. It is relatively uncommon in New Caledonia and Fiji, but very abundant in the Philippines. Plate 44. Living animal of female Strombus labiatus ( Rod- ing). (from Quoy and Gaimard, 1833, pi. 51, fig. 5). Strombus labiatus subspecies labiatus (Roding, 1798) (PI. 20, figs. 22, 23; pi. 44, pi. 45, figs. 1-6) Range— Southeast Asia to Japan, Australia and Melanesia. Remarks—' This is an abundant and well-known species in the central part of its range. It is very variable in size, coloration and sculpture. It may be strongly or weakly ribbed and spirally corded. It is distinguished from S. urceus in having an orangish columella which is generally weakly lirate along its entire length (instead of whitish or black-bordered and smooth at the center ) , in having a more trun- cate siphonal canal (instead of being drawn out), and in usually having strong, axial plications and sometimes in being spirally corded over the entire outer surface. The geographical range of urceus is more restricted than that of labiatus. Habitat— S. labiatus is always associated with an algal bottom which is generally intertidal or rela- tively shallow. S. urceus is not as closely associated with algal bottoms, and may be dredged on pure sandy mud to a depth of 20 fathoms. The latter is seldom found on fringing reef flats. Colonies of these two species are never found co-existing, and among about 50 accurately localized collecting sta- tions in the ANSP collection the two species were never taken at the same, immediate station. Description— Shell 16 to 47 mm. (% to 1/4 inches) in length, solid, axially plicate to smoothish, vari- colored, and with a yellowish to orange columella which has darker-colored, weak spiral lirae. Color of outer shell very variable, ranging from brown, greenish, gray, or yellowish maculations, flecks, broken bands and axial streaks to a solid light- yellow or orangish. Whorls 8 to 10. Nuclear whorls 3, bulimoid, glossy, smooth, translucent whitish, tan or brown-banded. First postnuelear whorl with about 10 microscopic, spiral incised lines. Next whorl with fewer incised lines, but with a weak subsutural cord and with about 20 weak, rounded axial ribs which in later whorls become noduled at the carinate shoulder. Penultimate whorl with 12 to 16 axial, well-shouldered ribs; whorls rarely rounded and rarely without ribs. Last whorl with 8 to 12 axial plications which are crowded, long and slanting on the rounded parietal wall, but on the dorsum are developed into 2 or 3 shoulder knobs. Last whorl rarely smooth and with a rounded shoulder. Base of whorl with 8 to 10 flat spiral cords, which in some specimens are raised and rounded and extend over the entire surface of the last three whorls. Siphonal canal truncate below. [09 - 895] 68 Canarium R. T. Abbott Strombidae Plate 45. Strombus labiatus (Roding). Figs. 1-4, adults, Cebu Island, Philippines. 5 and 6, immatures, Cebu Island, Philippines. All natural size. Columella narrow, with a sharp left edge, g’ossy, orange to yellowish orange and with about 30 to 40 spiral stripes of darker brownish orange. On these are raised lirae throughout the length of the colu- mella in some specimens, and raised lirae only at the upper and lower end in other specimens. The upper lirae are usually uneven, broken or inter- twining. Interior of body whorl with numerous, crowded, raised brownish purple, spiral lirae. The coloration extends almost out to the edge of the lip. Stromboid notch well or poorly developed. Spire with none or 1 to II very small, whitish, axial, swol- len, former varices. In young specimens there is usually a brownish or purplish color patch on the upper and inner portion of the parietal or columel- lar wall. Periostracum thin, smoothish, translucent grayish and flakes off when dry. Operculum strom- boid, not arching, light-brown, with 8 to 9 well- developed serrations, and with a longitudinal con- cave channel on the outer side. Radular ribbon 4 to 5 mm. in length and with about 38 rows of teeth. Formula: 2-1-2; 1-3 (plus peg) and 1-2 (plug peg); 4; 5. The inner marginal is considerably smaller and broader than the outer marginal. Animal (preserved) mottled with red maculations and with a red ring around the eye. Posterior mantle filament 2 to 2.5 mm. in length. Measurements (mm.)— length width no. whorls 47.0 24.2 8 + ( giant; Okinawa Id. ) 44.0 23.1 8 + ( large; Okinawa Id. ) 35.1 17.5 10 ( average; Luzon Id. ) 22.1 12.8 8 ( small; Luzon Id. ) 17.5 10.0 8 ( dwarf; Schouten Ids Synonymy— 1798 Lambis labiatus Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 63, no. 806 (no locality). Refers to Conehyl.-Cab., vol. 3, figs. 804-805. ?1807 Lambis reticulata Link, Beschr. Natur.-Sammlung, Rostock, vol. 2, p. 108. Refers to Conehyl.-Cab., vol. 3, fig. 806. 1816 Strombus plicatus Lamarck, Le Liste, Paris, p. 3; En- c-yclop. Method., pi. 408, fig. 2a and b (no locality); 1822, Anim. sans Vert., vol. 7, p. 210 (Moluques); 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 31, fig. 1 (not fig. lb). Non Roding 1798, non Miihlfeld 1829. 1850 Strombus dentatus Linne, Reeve, Conch. Icon., vol. 6, pi. 9, fig. 17. Not Linne, 1758; 1844, Duclos, in Chenu’s Ulustr. Conchyl., vol. 4, pi. 5, figs. 1-6. 1925 Strombus (Canarium) plicatus Lamarck, Oostingh, Mededeel. Landbouwhoogeschool Wageningen, vol. 29, pt. 1, pp. 59-69, form B only. ?1931 Canarium otiolum Iredale, Records Australian Mus., vol. 18, no. 4, p. 212, pi. 23, fig. 6 (Sydney Har- bour, N.S.W., Australia) [probably an immature], 1946 S trombus uslulatus form plicatus Lamarck, Dodge, Amer. Mus. Novitates, no. 1314, p. 2, fig. 3. 1959 Canarium dentatum Linne, Kira, Coloured Illus. Shells of Japan, Osaka, 2nd ed., p. 35, pi. 15, fig. 6. Types— Roding’s species is based upon figures 804 and 805 of vol. 3 of Martini’s Conchy lien-Cabinet. Roding gave no type locality. Martini mentioned Amboina, Indonesia, which we hereby designate as the type locality. Figure 806 is probably also labiatus, but 803 is an urceus, probably from the mainland of southeastern Asia. The former is the type figure for reticulata Link. The type of plicatus Lamarck is figured in “mirror image” on plate 408, fig. 2, a, b of the Tableau Encyclop. Method., pt. 23. It is the common Philippine plicate form. Nomenclature— Various names have been applied to this species by various authors. S. plicatus La- marck, 1816 is frequently seen in the literature. That name is a homonym of plicatus ( Roding, 1798). S. urceus of authors (not of Linne) and ustulatus of authors ( not Schumacher ) have also been used. See further discussion under the nomen- clature of urceus. Selected records ( see accompanying map, pi. 46. Solid dots: specimens examined. No literature records used) — ANDAMAN ISLANDS: Port Blair (W. N. Carpenter, USNM ) . THAILAND: Koh Chang, Gulf of Siam (USNM). JAPAN: Shirahama, Wakayama Pref., Honshu Id. (T. Habe, ANSP). PHILIPPINES: abundant throughout most, if not all, of the islands. INDONESIA: Keledjitan, Bantam, Java (USNM); Amboina, Moluccas (MCZ). AUSTRALIA: Port Douglas, Queensland (Tony Marsh, ANSP). NEW CALE- DONIA: Touho Bay and reef at Yate (both G. and M. Kline, 1958, NSF). NEW HEBRIDES: Aore Id. (MCZ). [09 - 896] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 69 PALAU ISLANDS: reef, Karamando Bay, Babelthuap Id., rare (NSF, 1955). FIJI: Lambasa, Vanua Levu Id. (ANSP); Buca Bay, Vanua Levu Id. ( R. T. Abbott, MCZ). Nataka- lau, Ovalau Id. (T. Dranga, MCZ). [The USNM has one Guam specimen of doubtful authenticity.] Fossil records— No authentic fossil records exist, since none of the records for “plicatus Lamarck, etc. have been illustrated. Strombus labiatus subspecies olydius Duclos, 1844 (PI. 20, figs. 18, 19) Range— Central East Africa to Mauritius and Ceylon. Remarks and description— This subspecies is limited to the western part of the Indian Ocean. It is uncommon in collections. In external characters, such as sculpture and color, it shows no appreciable difference from the East Indian labiatus labiatus. The aperture, however, consistently shows a strik- ing difference both in color and sculpture. The columella is light-mauve to brownish purple with broad, raised, enamel-white spiral lirae. In imma- ture specimens the lirae may be obsolete at the center of the columella and coloration may be limited to only a few brownish purple, fine streaks. Inside of body whorl and outer lip heavily pig- mented with mauve over which are numerous, irregular, raised, fine spiral lirae. Operculum strom- boid, similar to labiatus, and with 9 serrations. Eye in preserved specimens with a narrow red ring, and posterior to this a narrow yellowish one. Radula 4 mm., with 35 rows, and similar to labiatus and urceus. Formula: 2-1-2; 1-4 (plus peg); 4; 5. The body has a marble-like color pattern similar to that of urceus. Habitat— Found on intertidal, weedy, dead coral reefs. Measurements ( mm.)— length width no. whorls 47.5 19.5 9 (large; Zanzibar) 38.7 20.8 9 + ( average; Zanzibar ) 31.5 14.5 8 + (small; Madagascar) Synonymy- 1844 Strombus olydius Duclos, in Chenu’s Illustrations Con- chyliologiques, Paris, vol. 4, Strombus, p. 4, pi. 5, fig. 7 ( no locality ) . 1929 S trombus urceus Linne, var. olydius Duclos, Dautzen- berg, Faune des Colonies Frangaises, Paris, vol. 3, pt. 4, p. 470. Types— The type of olydius is presumably in the Museum d’Histoire naturelle de Geneve. No local- ity was given by Duclos. We hereby designate Nossi-be, northwest Madagascar, as the type locality. Nomenclature— We know no synonyms of olydius. It has been put in the synonymy of plicatus La- marck and urceus Linne by most previous authors. Records— KENYA: Kikambala, 15 mi. north of Mombasa (Coryndon Mus. no. 2582). TANGANYIKA: Mboa Magi (R. T. Abbott, USNM). MOZAMBIQUE: Port Amelia (LISNM). ZANZIBAR: Kiwengwa; Mangapwani; Fumba; Mazizini (all NSF, 1957). MADAGASCAR: Nossi-be (A. Chavane, ANSP; MCZ). MAURITIUS: (N. Pike, MCZ). CEYLON: Hikkaduwa (R. Jonklass, ANSP). Fossil records— None reported. Strombus labiatus subspecies gendinganensis Martin, 1899 Range— Fossil (Pliocene) of Java, Indonesia, the Philippines and possibly Taiwan. Remarks— This subspecies differs from the living labiatus in usually having the axial riblets and the elongate shoulder and parietal plications greatly re- Plate 46. Geographical distribution of Strombus labiatus la- hiatus (Roding) and Strombus labiatus olydius Duclos. [09 - 897] 70 Canarium R. T. Abbott S tromhidae duced. Many of the Pliocene literature records of plicatus Lamarck in the Indonesian area are prob- ably this subspecies ( see Altena’s listings, 1942, vol. 12, p. 55-57). From his description (but not illus- tration), it would appear that genclinganensis occurs in the Pliocene, Byoritu Beds of Taiwan (Nomura, 1935, p. 179). Dickerson’s Strombus dentatus sonde is a slightly more plicate form of this subspecies. For Abrard’s 1946 Pliocene New Hebrides record, see under urceus Linne. Synonymy— ] 899 Strombus ( Canarium ) genclinganensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 187, pi. 30, figs. 432, 432a (typical), 433, 433a (atypical) (Pliocene, java); 1942, Altena, Leidsche Geologische Mededeelingen, vol. 12, p. 57 (Pliocene, Java); 1922, Dickerson, Philippine Jour. Science, vol. 20, no. 2, p. 225, pi. 5, fig. 4 (Vigo Group, Upper Miocene [? Pliocene], Bondoc Penin- sula, Luzon Id., Philippines). 1899 S trombus dentatus Linn, var., K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 188, pi. 30, fig. 437 [or is labiatus Roding] (Java, Plio- cene ) . 1922 Strombus dentatus sonde “Martin” Dickerson, Philip- pine Jour. Science, vol. 20, no. 2, p. 225, pi. 5, fig. 7 (Vigo Group, Bondoc Peninsula, Luzon Id.). Strombus klineorum new species (PI. 20, figs. 20, 21) Range— Known only from Ceylon. Remarks— This distinctive new species is so far known only from the area around Ceylon. It closely resembles labiatus Boding in morphological charac- ters but appears consistently different in the follow- ing characters: parietal wall smooth and somewhat flattened; outer half of columella salmon red to red- dish yellow, the inner half being a solid purplish black; penultimate whorl with fewer shoulder knobs; and the shoulder of the outer lip extending to the shoulder above. Habitat— Found along the shores of Ceylon in 2 to 5 feet of water on a sandy mud and rocky bot- tom in company with Conus, Cypraea and Bursa (G. and M. Kline). Shallow water within fringing reef, common (B. Jonklass). Description— Adult shell 17 to 37 mm. to VA inches ) in length, smoothish, except for strong knobs at the shoulder, with a brightly colored aper- ture, and resembling labiatus. Whorls 7 to 9. Nu- clear whorls 3, glassy, smoothish, translucent tan and moderately rounded. Postnuelear whorls with 7 to 10 fine incised spiral lines which in the third whorl become crossed by 16 to 19 rounded, evenly spaced axial ribs which become increasingly nodu- lated at the central shoulder. The penultimate whorl has usually 11 (rarely 14) small knobs and the body whorl only 3, of which the last is very large and blunt and located }i of a whorl back from the swollen varix. Ventral wall peculiarly flattened and smooth. Suture slightly impressed and bordered below by a weak, small spiral cord only in the first 3 or 4 postnuelear whorls. Color of outer shell olivaceous brown with suffused maculations of light yellow-brown. Center of body whorl with 2 or 3 narrow spiral hands of tiny clusters of opaque- white dots. Aperture squarish at the top, broader below. Upper end of outer lip shouldered by a large swelling. Columella slightly arching, with a strong border on its left. The outer border of the columella and the inner narrow border of the outer lip are bright carmine to yellowish red. The entire inner half of the columella and a broad band inside the outer lip are very dark blackish purple. Deep within the aperture whitish to cream. Columella smooth at the center, but with spiral teeth at the lower and upper fourth. Inside of body whorl with numerous, raised, fine, black or gray spiral lirae. Tip of short, straight siphonal canal without a black stain. Periostracum thin, translucent, and smoothish. Operculum stromboid, slightly less than % the length of the shell, not arching, light-brown, with about 8 small serrations, and with a weak central rib on the attachment side. Body coloration similar to that of urceus. Measurements (mm.)— (all adults fi-om Ceylon) length width no. whorls 35 20 8.5 (holotype, ANSP 247621) 36 20 8.5 (paratype, ANSP 211121) 27 15 8.0 ( paratype, ANSP 211121 ) 26 14 7.0 (paratype, ANSP 211173) 19 10 7.0 (paratype, ANSP 211173) Synonymy— 1844 Strondms urceus L., Duclos, in Chenu, Illustr. Con- chyl., vol. 2, pi. 5, figs. 10, 11. Non Linne 1758; 1850, Reeve, Conch. Iconica, vol. 6, pi. 11, fig- 24a. Types— The type locality is Powder Bay, near Trincomalee, Ceylon. Collected by George F. and Mary Kline on January 17, 1957. Holotype is ANSP no. 247621, paratypes in ANSP 211121 and 211173; paratype in MCZ 224889. The species is named with pleasure after the Klines who have been inde- fatigable collectors for the Department of Mollusks at the Academy. Locality records (see map, pi. 43)— CEYLON: Kacheri, Powder Bay, and south shore of Fort Frederick, all Trin- comalee (NSF). Akurala, S.W. end of island (R. Jonklass, ANSP). [Literature records from Ceylon and southern India of S. plicatus Lamarck, S. urceus Linne, etc. may be this species.] [09 - 898] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 71 Strombus microurceus (Kira, 1959) (PL 20, figs. 24, 25) Range— Southern Japan to the East Indies, Aus- tralia, Melanesia and to Samoa. Remarks— This small and attractive species has several characters which separate it from closely re- sembling mutabilis and urceus. It is a Southwest Pacific species rather than a widely dispersed one like mutabilis. S. microurceus has a distinctive colu- mella. The outer half is yellowish to whitish orange; the inner half is very dark brownish purple. Across the latter run about 25 to 30, fine, short, spiral lines of yellowish which usually coincide with minutely raised spiral lirae. The latter are usually rather weak or absent near the middle of the columella. Deep within the aperture, the inside of the body whorl is whitish to yellowish, but towards the outer lip there is a broad, well-defined, longitudinal band of dark brownish purple which is crossed by about 50 to 60 whitish, raised, spiral lirae. The exterior of the si- phonal canal has a small purplish patch, a feature absent in mutabilis. Just behind the outer lip and just below the suture, there is usually a peculiar elongate patch of grayish green or bluish gray. We have seen one specimen presumably from Zanzibar which closely resembles microurceus in many respects, except that the columella is smooth- ish and white in the center, and the shoulder of the last whorl bears several evenly-sized, axially- pinched, short knobs. When more material is at hand, this may prove to be a valid Indian Ocean subspecies of microurceus. Habitat— This is a relatively uncommon species, or at least sparse in numbers wherever it occurs. It has been collected on sandy and rocky areas just below low tide and dredged in depths from 1 to 12 fathoms on sand and dead coral bottoms. Description— Adult shell 14 to 27 mm. (about /2 to 1 inch) in length, solid, smoothish, somewhat quadrate in shape and resembling mutabilis. Whorls 7 to 8. Nuclear whorls 3, glassy, smooth, translu- cent tan and well-rounded. Postnuclear whorl microscopically reticulate. Second, third and fourth postnuclear whorls bear 4 to 5 incised spiral lines, numerous, weak axial nodules and 3 to 5 old, whit- ish, swollen varices. Body whorl bears 3 to 6 blunt nodules at the shoulder, the last two being the larg- est and placed well back from the outer lip. Suture moderately indented and bordered below by a single, strong rounded cord which is rarely spirally striate. Color of outer shell variable but usually a cream to white background overlaid with brown and/or yellowish reticulations, maculations, suf- Plate 47. Geographical distribution of Strombus micro- urceus (Kira). fused spiral bands and/or minute stripes or bars. Outside of base of shell typically with 2 irregular patches or blue-brown. An all yellow or orange phase occurs rarely. Aperture narrow above, broad below. Columella slightly concave, smooth and yel- lowish orange along the outer border, but within the aperture it is dark purple-brown. The latter area bears 22 to 32 narrow, spiral flames or streaks of yellowish orange which on the lower and upper part of the columella bear raised, rounded teeth. Inside of outer lip with a wide band of dark purple- brown over which are numerous, irregular, raised spiral lirae or teeth. Deep interior of aperture dirty white to yellowish. Stromboid notch on lower outer lip relatively narrow and well-developed. Base of shell with 11 to 14 strong, rounded, spiral threads. Periostracum very thin ( peels off when dry ) , not smooth and light-tan. Operculum stromboid, one third the length of the shell and with 7 serrations. Radula ribbon 2 mm. in length, with 29 rows and similar to mutabilis. Its formula is: 2-1-2; 1-2 (plus peg); 5; 7. Preserved animals cream with sparse orange spots. Verge half the length of the shell, long, slender, with weak, lemon mottlings and a large lamellated pad at the distal end (Upolu Id., Samoa ) . Measurements (mm.)— length width no. whorls 27.0 14.0 7 ( large; Okinawa Id. ) 22.0 10.5 7 ( average; Palau Ids. ) 14.0 7.0 6 ( small; Mindanao Id. ) [09 - 8991 72 Canarium R. T. Abbott S trombidae Synonymy— 1955 Canarium microurceum “Kuroda, MS” Kira, Coloured Illust. Shells Japan, Osaka, ed. 1, pi. 15, fig. 5 ( nude name ) . 1959 Canarium microurceum “Kuroda, MS” Kira, Coloured Illust. Shells Japan, Osaka, ed. 2, p. 37, pi. 15, fig. 5 (Honshu southward). Types— A general geographic range was given by Kira ( central Honshu and southward), and we hereby restrict the type locality to Shirahama, Wakayama Pref., Honshu Id., Japan (ANSP no. 234923). The type figured by Kira is presumably in the collection of Mr. Tetuaki Kira of Japan. Nomenclature— Although Kuroda is given as the author of this species, it was a manuscript name and must now bear Kira as the author. Selected records (see accompanying map, pi. 47; solid dots: specimens examined; open circles: from the literature) —JAPAN: Shirahama, Wakayama Pref., Honshu (T. Habe, ANSP). PHILIPPINES: shore reefs, Cape Santiago, Ba- tangas Prov., Luzon Id. ( Dayrit and Abbott, ANSP); Gig- moto, Catanduanes Id. (ANSP). Also Panay, Mindoro, Mindanao Ids. (USNM). INDONESIA: Keledjitan, Bantam, Java. AUSTRALIA: Green Id., near Cairns, Queensland (MCZ). LORD HOWE ISLAND: ( H. L. Clark, MCZ). MARIANAS: Agrighan Id. (Robert R. Hill, ANSP); Agat Bay, Guam Id. (A. J. Ostheimer, NSF). CAROLINES: Ponape Id. (V. Wertley, ANSP); Yap and Elato Atoll (both USNM). FIJI: Suva, Viti Levu Id. (H. S. Ladd, USNM). SAMOA: east side of Vailele Bay, Upolu Id. (A. J. and Ruth Ostheimer, 1955, NSF). Fossil records— None reported. Strombus mutabilis Swainson, 1821 (PI. 20, figs. 15, 16, 17) Range— The entire tropical Indo-Pacific exclusive of Hawaii, the Line Islands, Marquesas and Easter Island. Remarks— This is a very widely distributed spe- cies which is common on reef flats. It is character- ized by its somewhat quadrate shape and rose- tinted aperture. The columella is pink, rose or faint brownish rose over which are 30 to 40 broad, raised, sometimes bifurcating, white, spiral lirae. Where specimens have been heavily covered with algal growths, an abnormal glaze may also cover them (as in some New Caledonia specimens). S. mutabilis is apt to be confused with arceus ( which has a longer, purple-tipped siphonal canal ) , with maculatus (which has a flattish spire and a white, smooth columella), with labiatus (which has axial riblets on the whorls in the spire) and with microurceus (which has the inner half of the colu- mella blackish purple ) . Strombus mutabilis, despite its variations in color and size among individuals, is a fairly constant spe- cies as a whole throughout its range. We recognize one weak subspecies in the Mauritius area— a golden-throated, white-columella shell, mutabilis ochroglottis Abbott. It has not appeared in Mada- gascan or East African collections, and one old specimen from P. P. Carpenter in the USNM sup- posedly from Aqaba, Red Sea, we look upon with suspicion. Abbott ( 1949, pp. 59-60 ) has demonstrated the existence of sexual dimorphism in the shell length of mutabilis. In the colony of 462 specimens from Agana Bay, Guam Island, Marianas, the mode of the shell length of the entire colony was 23 mm., that of the 210 males was 20.8 mm. and that of the 252 females was 24.1 mm. The range in size of the two sexes, however, was such that the character of the shell length cannot Ire used as an indication of sex, except for the generalization that the largest specimens are female and the very smallest are males. The rare color form, zebriolatus Adam and Leloup, (see pi. 20, fig. 17) possesses long, dark-brown to chestnut, axial, narrow, zigzag color stripes. It was originally found in Manokwari, Dutch New Guinea, and the Academy collection contains specimens from Catanduanes, Philippines; Okinawa Id., Ryu- kyus; Babelthuap Id., Palaus; and Ponape Id., Caro- lines. We have not seen this color form in Indian Ocean specimens. A pure orange color form occasionally occurs in some colonies. We have seen specimens from Zan- zibar, Ryukyu Islands, Luzon Island (Philippines), Choiseul Island (Solomons), Padaido Islands (Dutch New Guinea) and Upolu Island in West- ern Samoa. Habitat— Found in colonies from the intertidal zone to a depth of 10 fathoms, although it more Plate 48. Living animal of male Strombus mutabilis Swain- son. (from Quoy and Gaimard, 1833, pi. 51, fig. 12). [09 - 900] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 73 commonly is found just below the low tide line. Found where there is a mixture of coral sand or sandy mud and algae, whether these conditions are on outer reef flats, lagoons or sheltered shallow bays. The water may be clear or turbid. It is often associated with Strombus gibberulus, but not, to my knowledge, found intermingled with S. macu- latus. Description— Adult shell 16 to 40 mm. ( about 'll to 1/2 inches) in length, solid, smoothish, vari- colored, somewhat quadrate in shape, and with a pinkish aperture and striate columella. Whorls 8 to 9. Nuclear whorls 3, glassy, smooth, translucent white and well-rounded. First postnuclear whorl with 5 to 7 microscopic, incised, spiral lines which become obsolete in the penultimate or last whorl. By the second postnuclear whorl, a strong, rounded cord appears just below the indented suture but which may again disappear on the last fourth of the body whorl. Spire with 1 or 2 to as many as 15 swollen, whitish, sometimes broad, former axial varices. Whorls in spire usually shouldered, smooth- ish or axially ribbed or with small, weak nodules. Last whorl with 3 or 4 ill-formed, swollen, nodules at the shoulder which are located % to I2 of a whorl back from the swollen outer lip. Base of shell with about a dozen small, spiral threads. Parietal wall evenly rounded and smooth. Columella enameled, pinkish brown to mauve-brown over which are 30 to 40 rather broad, raised, whitish, rarely bifurcat- ing, spiral lirae which are strongest at the base of the columella. Inside of the body whorl usually pink, sometimes whitish or mottled with brownish, and bearing numerous, fine, raised, whitish, spiral lirae which usually run out to the edge of the outer lip. Stromboid notch poorly to well-developed. Up- per end of the aperture squarish and bounded on the right by a swollen portion of the outer lip. Siphonal canal short. Color of outer shell variable but usually bright: maculated, flecked, banded and spotted with either browns, yellows, blacks or orange. Rarely unicolor; rarely with long, zigzag axial flames or streaks of chestnut-brown (form ze- briolata Adam and Leloup). Periostracum thin, smoothish and translucent yellowish. Operculum stromboid, barely arching, dark- to light-brown, with 8 to 12 small, sharp serrations, and with a con- cave, longitudinal channel down the middle of the outer surface. Radula ribbon delicate, 3 to 4 mm. in length, with 36 to 42 rows of teeth. Formula: 2-1-2; 1-2 (plus peg); 4 or 5; 5 or 6. Verge slender, simple, with an elongate, lamellated, distal pad. Its base is green with white, embedded granules and orange Plate 49. Figs. 1-5, radula and verge of S trombus mutabilis Swainson, Luzon Island, Philippines. 6, verge of S. micro- urceus (Kira), Upolu Island, Western Samoa. 7 and 8, verge and operculum of S. terebellatus afrobellatus Abbott, Zanzibar. spots. The distal half is mottled in green. Posterior mantle filament green or orange and 2 mm. in length. See plate 49, figs. 1-5. M easurem ents (m m .)— length width no. whorls 40.1 22.9 8 ( large; Kenya ) 33.5 17.3 8 ( average; Samar Id. 15.8 8.3 7 (small; Samoa) Synonymy— 1821 Strombus mutabilis Swainson, Zoological Illustrations, London, series 1, vol. 2, pi. 71 (East Indies). Also refers to figures of Seba, pi. 61, figs. 26, 27, etc,; Conchyl.-Cab., vol. 3, figs. 799 and 807; Knorr, pt. 2, pi. 14, fig. 3. 1822 Strombus floridus Lamarck, Anim. sans Vert., vol. 7, p. 21 1 ( Ocean indien et des Moluques ) . Refers to Conchyl.-Cab., vol. 3, figs. 807-809; Lister, pi. 848, fig. 3; and others. 1840 S trombidea mutabilis Swainson, Treatise on Malacol- ogy, London, p. 310 (no locality). Refers to Con- chy!.-Cab., vol. 3, fig. 807. 1844 Strombus epimellus Duclos, in Chenu, Illustr. Conch., vol. 4, p. 3, pi. 16, figs. 11, 12; pi. 22, figs. 5, 6. 1852 Strombus flosculosus “Martini” Morch, Cat. Conehyl. Yoldi, vol. 1, p. 63 (Indies orientalis). 1876 Strombus flammeus Link, Issel and T.-Canefri, Annali Mus. Civico Storia Nat. Genova, vol. 8, p. 346. Also of other authors, Morch, 1852; Iredale, 1931; Ab- bott, 1949 (Nautilus, vol. 63, no. 2, p. 58), 1950; Dodge, 1946. Not Link, 1807. 1938 Strombus floridus Lamarck form zebriolata “Dautzen- berg” Adam and Leloup, Mem. Mus. Roy. Hist. Nat. Belgique, special series, vol. 2, fasc. 19, p. 117, pi. 1, fig. 7 (Manokwari, Dutch New Guinea). [09-901] 74 Canarium R. T. Abbott Strombidae Plate 50. Geographical distribution of Strombus mutabilis Swainson, and its subspecies ochroglottis Abbott in Mauri- tius. Types— The whereabouts of Swainson’s type of mutabilis is unknown. His type locality was “East Indies” which we further restrict to Cebu City, Cebu Island, Philippines. Nomenclature— Morch seems to be the earliest author responsible for associating the name fiam- meus Link, 1807 with the species. We believe this to be entirely unjustified, since Link’s name is based upon figure 799, of vol. 3, Conchylien-Cabinet. Mar- tini’s description, synonymy and figure are surely the young of Strombus decorus Roding of the In- dian Ocean. Lamarck’s well-known name, floridus, has been abandoned for many years because it was published a year later than Swainson’s. Selected records (see accompanying map, pi. 50; solid dots: specimens examined)— NATAL: Durban Bay (USNM). MOZAMBIQUE: Mozambique City ( K. Grosch, ANSP). KENYA: Malindi (USNM). RED SEA: Aqaba ( MCZ, ANSP); Jidda (USNM); Geb Zebara, Egypt (ANSP). ZANZIBAR: Paje; Chumbe Id.; Ras Nungwe; Fumba; Man- gapwani; Chwaka; Mazizini (all NSF, 1957). MADAGAS: CAR: Nossi-be (A. Chavane, ANSP). SEYCHELLES: Frig- ate Id.; Beau Vallon, Mahe Id. (both Yale Peabody Mus. ) MALDIVES: Hulule Id., N. Male Atoll; Gan Id., Addu Atoll (both Yale Peabody Mus.) CEYLON: Hikkaduwa (G. and M. Kline, 1956, ANSP). COGOS-KEELING ATOLL: (USNM). JAPAN: Shirahama, Wakayama Pref., Honshu Id. ( T. Habe, ANSP); Hachijo Id., 275 south of Tokyo (A. R. Calm, ANSP). RYUKYUS: Oshima, Osumi (MCZ); Okinawa Id. (Mrs. A. A. Scott, ANSP). PHILIPPINES: common throughout the islands: Luzon, Cebu, Samar, Ca- tanduanes, Bohol, Negros, Marinduque, Cuyo, Camiguin, Mindanao, Sulu Archipelago (ANSP, USNM and/or ANSP). INDONESIA: Tjilaoet Eureun, south Java (USNM); Am- boina (MCZ). AUSTRALIA: Western Australia: Yallingup Brook; Abrolhos Id., off Geraldton (J. A. Grigg, USNM). Queensland: Green Id., near Cairns (MCZ); Low Isles (Tony Marsh, ANSP); Wilson Id., Capricorn Group (MCZ). New South Wales: The Entrance, and Woolgoolga (both W. E. Old, Jr., ANSP). LORD HOWE ISLAND: (Tomlin, MCZ). CAROLINES: Yap Id. (C. O. Kile, ANSP). MAR- SHALL ISLANDS: common at Bikini, Eniwetok, Rongelap, and Wotho (USNM). AUSTRAL ISLANDS: Rurutu Id. (Aubert de la Rue, ANSP). Tubuai Id. ( H. J. Klein, ANSP). TUAMOTUS: Takume Atoll; Ngarumaoa Id., Raroia Atoll (both J. P. E. Morrison, USNM). [Record for Hawaii in Hawaiian Shell News, vol. 8, no. 7, p. 1, 1960 appears to be maculatus Sowerby.] Fossil records— Fossil mutabilis have been reported in the literature from the Pleistocene of Hawaii ( J. M. Ostergaard, 1928, p. 27), the Red Sea ( R. B. Newton, 1900), Viti Levu Id., Fiji (Ladd, 1934). Somalia (Abrard, 1941, p. 63), Plio- cene of New Hebrides (Abrard, 1946, p. 62), and Indo- nesia, but 1 have not seen specimens to verify these records. Abrard’s 1942 (vol. 18, p. 63, pi. 6, fig. 38) record is good for the Pleistocene of Khor Anghar, French Somalia. Strombus mutabilis subspecies oebroglottis new subspecies (PI. 20, figs. 9, 10) Range— Known only from Mauritius, Indian Ocean. Remarks and description— Colonies of this species from Mauritius are so uniformly different in their apertural coloration and in some sculptural details from colonies elsewhere in the Indo-Pacific that we [09 - 902] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 75 have allotted them to a subspecific rank. However, the differences are not always entirely consistent. The new subspecies ochroglottis differs in having the greater part of the columella pure white, while deep inside the aperture, both along the columella and the inner part of the body whorl, there is a chrome-yellow taint. In most specimens the colu- mella is smooth along the central portion. Most, but not all, specimens are more coarsely noduled and with a higher spire than most typical mutabilis. Also, the weak, small splotches of green found be- low the suture in the last whorl are absent in Mau- ritius specimens. Measurements (mm.)— length width no. whorls 28.0 16.0 7 ( holotype, ANSP ) 35.9 20.0 7 + ( large, paratype ) 27.0 15.0 7 ( average, paratype ) 16.1 9.0 6 ( small, paratype ) Habitat— Unknown; presumably shallow-water or intertidal. Synonymy— None; nor have we located any pre- Linnaean figures. Types— The type locality is Mauritius, Indian Ocean. Holotype, ANSP no. 250187; paratypes, ANSP no. 185466; USNM no. 26652a and 465685; and in the MCZ. Locality records— Known only from Mauritius. [A single Red Sea record in the USNM is probably an erroneous locality]. Strombus mutabilis subspecies ostergaardi Pilsbry, 1921 (Plate 51) Range— Pleistocene of Oahu Island, Hawaii. Remarks and description— This subspecies is known only from the Pleistocene. It resembles both mutabilis and maculatus, but I associate it with the former because of the nature of the sculpturing in the spire and on the columella. The shell is some- what pear-shaped with well-rounded shoulders. The whorls in the spire bear 14 to 18 small, but dis- tinct, shouldered axial riblets or elongate beads which disappear in the penultimate and last whorl. The columella bears about 40 small, but distinct, spiral lirae. The inside of the outer lip bears numer- ous fine spiral lirae. Stromboid notch moderately developed. Ostergaard ( 1960, Hawaiian Shell News, vol. 8, no. 7, p. 3) reports live specimens from Kwajalein Atoll, Okinawa Id., and New Hebrides, but 1 have not seen these specimens. If verified, this may be considered a full species. Plate 51. Strombus mutabilis ostergaardi Pilsbry. Pleisto- cene of Oahu Id., Hawaii. Left, holotype; right, paratype. Both x 2. Measurements ( rn in.)— length width no. whorls 24.7 12.2 9 ( holotype, Honolulu ) 31.2 16.1 7 + ( paratype, Honolulu ) 21.6 11.9 8 ( paratype, Kailua ) Synonymy- 1921 Strombus ostergaardi Pilsbry, Proc. Acad. Natural Sci- ences Philadelphia for 1920, vol. 72, p. 320, pi. 12, figs. 27, 28 (Honolulu Harbor); 1928, Ostergaard, Bull. 51, B. P. Bishop Mus., p. 25, pi. 1, fig. B. Types— The type locality is “from the dredger dump on the Harbor side of Sand Island, Honolulu Harbor,” Oahu Island, Hawaii. J. M. Ostergaard, collector. Holotype in ANSP no. 74549; paratypes in ANSP nos. 247760 and 74550 (Kailua). Records— Pleistocene: Oahu Island: Honolulu Harbor (ANSP); Wailupe Quarry no. 2, vicinity of Nanakuli Station; Mokapu Point; Kailua Coast (all J. M. Ostergaard, 1928, p. 25). Strombus maculatus Sowerby, 1842 (PI. 20, figs. 13, 14) Range— Hawaiian Chain, Micronesia and eastern Polynesia to Easter Island. Remarks— This species is abundant throughout the Hawaiian Chain and the Marshall Islands, but becomes progressively uncommon eastward towards Easter Island. Despite intensive collecting and our examining many hundreds of Strombus lots from the Marianas, Palaus, Carolines, Solomons and Samoa, we have seen only four specimens from those areas— one live specimen each from Kayangel, [09 - 903] 76 Canarium R. T. Abbott S trombidae Plate 52. Geographical distribution of S trombus macu- latus Sowerby. The species is rare in the Caroline and Palaus (NSF station 395, 1955), Elato Atoll and Satawal, Carolines, and Guam, Marianas. I have no reason to doubt the authenticity of these four rec- ords which occur a little over 1000 miles to the lee- ward and west of the densely populated Marshalls. This species also occurs in emerged beaches of pre- sumed Pleistocene age in the Hawaiians. It is pos- sible that Polynesia is the rather recent center of origin of this species and that it is, at this moment, spreading westward. An equally speculative ex- planation is that maculatus once occupied a much larger part of the Indo-Pacific and that mutabilis is moving eastward and, in some unknown manner, replacing or “crowding out” maculatus. Disease or genetic weakness can also explain the paucity of individuals in certain parts of the range of a species. S trombus maculatus is characterized by its pale color, relatively heavy shell, smoothish body whorl, rounded shoulder, the flattish whorls in its spire, its white aperture and by the smooth central por- tion of its glossy-white columella. Easter Island specimens are very richly colored with axial flames of chestnut-brown over a light- brown and white spotted background. Subsequent collecting of fresh specimens from Easter and the eastern Tuamotu Islands may substantiate the pres- ence of a subspecies of maculatus. This odd colora- tion is reminiscent of the S. mutabilis color form zebriolatus Adam and Leloup ( and incidentally the color pattern of Conus retifer Menke). Dautzenberg and Bouge’s variety depauperatus from Tongareva and the northwest Tuamotus is described as white or yellow with five transverse (axial) bands of small, very bright spots of reddish brown or russet. Should a subspecies exist in this area, this name might be applicable. Mariana Islands. Ostergaard ( 1950, p. 95 ) describes the egg mass as an entangled, single gelatinous, yellowish tube, 0.25 mm. in diameter, containing a single row of ova, each 0.10 mm. in diameter. They were laid on March 16, 1922. The smallest adult specimen of any Strombus I have seen was the 8-mm.-long maculatus collected by Dr. J. P. E. Morrison on the reef on the oceanic side of Enyu Island, Bikini Atoll, Marshall Islands. Description— Adult shell 8 to 36 mm. ( about M to 1/2 inches) in length, solid, moderately heavy, with smoothish, rounded, somewhat swollen shoulders, with a smooth central portion to the columella and with an enamel-white aperture. Color of outer shell whitish with weak maculations and fine nettings of light-yellow, light orange-brown or grayish brown. Rarely yellow with a white central band bearing irregular, small brownish spots. Very rarely with axial flames of reddish brown. Whorls 8 to 9. Nu- clear whorls 212, bulimoid, glassy, smooth, translu- cent whitish. First postnuclear whorl with 10 to 11 very fine but deep incised spiral lines becoming re- duced to 8 in the next whorl and gradually disap- pearing on later whorls. The first two or three post- nuclear whorls have a small subsutural cord, but this becomes weak in later whorls. The same whorls may have a spiral row of very small and weak pe- ripheral nodules or beads (12 to 14 per whorl). Other whorls in spire are smoothish and slightly convex. Former varices up to 14 and large. Shoulder of body whorl rounded, smooth, and may bear 1 to 3 very weak swellings. Surface of body whorl with numerous microscopic spiral threads. Base of shell with about a dozen weak spiral cords, either flattish or slightly rounded. Columella enamel-white, smooth at the center and usually with 3 to 7 spiral lirae at the top and 1 to 10 dentations near the base. [09 - 904] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strornbus 77 Inside of body whorl with 20 to 40 fine, raised spiral lirae. Stromboid notch usually quite shallow and poorly developed. Periostracum moderately thick, smoothish and translucent yellow. Opercu- lum stromboid, light-brown, relatively broad, thin, and with about 10 very small serrations. Soft parts not examined. Measurements ( mm.)— length width no. whorls 36.1 19.1 9 ( large; Hilo, Hawaii ) 26.8 14.0 9 ( average; Oahu Id. ) 12.9 6.3 8 (small; Marshalls) 8.0 4.2 8 (dwarf; Bikini Id.) Synonymy— 1839 Strornbus maculatus “Nuttall Jay, Catalog of Shells, N. Y., ed. 3, p. 82. Nude mime. 1842 Strornbus maculatus “Nuttall” Sowerby, Thesaurus Conchyliorum, London, vol. 1, Strornbus, p. 30, pi. 7, fig. 53 (Sandwich Islands); 1850, Reeve, Conch. Icon., vol. 6, Strornbus, sp. and fig. 23; 1957, De- mond, Pacific Science, vol. 11, p. 296. 1933 Strornbus floridus var. depauperata Dautzenberg and Bouge, Jour, de Conchyl., vol. 77, p. 296 ( Makatea [here selected as the type locality!, Apataki and Motutunga, Tuamotu Ids.). 1950 Strornbus maculatus Nuttall, Ostergaard, Pacific Sci- ence, vol. 4, p. 95, fig. 17 (egg mass). Types— Sowerby’s type of maculatus is presum- ably in the British Museum of Natural History in London. We restrict the original type locality of “Sandwich Islands” to Kawailoa, Oahu Id., Hawaii. Nomenclature— Although sometimes credited to Nuttall or Jay, this species was first described by Sowerby in 1842. Records (see accompanying map, pi. 52)— MARIANAS: Guam Id., 1 live specimen (F. J. Flatt, 1948, USNM). PALAUS: north side of Ngariungs Id., Kayangel (NSF sta- tion 395, ANSP, 1 live specimen. CAROLINES: Elato Atoll, near Lamotrek; Satawal Atoll (both USNM, but only 1 specimen). WAKE ATOLL: southeast end of Wake Island (du Pont- Academy Expect, 1958, ANSP). MARSHALLS: abundant on reefs on oceanic side throughout Eniwetok, Bi- kini, Rongelap, Uterik, Rongerik, Taka, Ailuk, Kwajalein and Lae Atolls (all USNM). GILBERTS: Apamama (USNM). PHOENIX IDS: Canton let; Hull Id. (both USNM). HA- WAIIAN CHAIN: Midway (Phil Spicer, ANSP). Tern Id., French Frigate Shoals (ANSP). Kauai Id. (A. Garrett, MCZ). Maui Id.: Honokowai (USNM). Oahu let: Kahaluu; Honolulu Harbor; Kawailoa; Waikiki; Waianae (all H. A. Pilsbry, ANSP). Hawaii Id.: Hilo (A. J. Ostheimer, 3rd, ANSP); Napoopoo (ANSP). LINE ISLANDS: Palmyra Id. (MCZ); Flint Id. (ANSP). COOK IDS.: Aitutake (J. P. E. Morrison, USNM); Rarotonga Id. (USNM). SOCIETY IDS.: outer reef, Moorea Id. ( H. A. Rehder, USNM). TUA- MOTUS: Reao ( Clermont-Tonnere) Id. (Titian R. Peale, 1824, ANSP); Takaroa Id. (A. de la Rue, ANSP); outer- reef, Raroia Atoll; outer reef flats, Takume Atoll (both J. P. E. Morrison, USNM); Fakarava Id. (USNM). EASTER ISLAND: (ANSP and USNM). [A. Garrett’s record of “Fiji’ in ANSP collection open to question.] Fossil records— HAWAIIAN CHAIN: Kauaiu Stream, Lanai Id.; Lanikai, Oahu Id.; Mokapu Peninsula, Oahu Id. (all USNM, all emerged beach of presumed Pleistocene age). LINE ISLAND: Flint Id. (1 post-Pleistocene fossil, ANSP). Fossil Relatives of the urceus- mutabilis-labiatus Group It would be unsatisfactory to make any decisions concerning many of the fossil forms in this group without examining good series of specimens which, however, are not available to us at this time. A few are sufficiently illustrated and described to assign them to a position close to either Strornbus urceus, mutabilis, labiatus, microurceus, maculatus or erythrinus. The records quoted for S. labiatus Rod- ing (as plicatus Lamarck) by Altena (1942, vol. 12, p. 55 ) from the Tertiary of the Indo-Pacific are un- doubtedly based upon a wide mixture of these spe- cies. S. genclinganensis Martin is treated as a sub- species under labiatus Roding. Plate 53. Strornbus ( Canarium ) unifasciatus Martin. Mang- kalihat, East Borneo, Indonesia. Lower Miocene. ( from Beets, 1941, pi. 3, figs. 119-122). Natural size. Strornbus unifasciatus Martin, 1884 Range— Lower and Upper Miocene of Java, In- donesia. Remarks— This species is very close to both mu- tabilis Swainson and maculatus Sowerby. Its stri- ated columella suggests a closer affinity to the former. Martin’s figures 436 and 436a are more like smooth erythrinus. The species was first founded upon an immature specimen. The figures given by Beets (1941, p. 65, pi. 3, figs. 119-122 and called unifasciatus Martin from the Lower Miocene of East Borneo appear to represent two species, one more similar to the mutabilis group, the other (figs. 121, 122) more like labiatus. Synonymy- 1884 Strornbus (Canarium?) unifasciatus K. Martin, Samml. geol. Reichs-Mus. Leiden, vol. 3, p. 143, pi. 8, fig. 142 ;ibid., 1899, n. Folge, vol. 1, pt. 1, p. 187, pi. 30, figs. 434, 434a, 435 (Tjilintung, Java); 1884, K. Martin, Jaarbock Mijnwegen in Nederland. Oost- Indie, Amsterdam, vol. 13, p. 176, pi. 8, fig. 142. Strornbus spolongensis Martin, 1916 Range— Lower Miocene ( West-Progo beds), Java, Indonesia. Remarks— This is closely related to Strornbus uni- [09 - 905] 78 Canarium R. T. Abbott Strombidae fasciatus Martin, but without examining specimens I would hesitate to suggest its closest affinities, ex- cept to say that it resembles the Recent mciculatus- mutabilis-microurceus group. Synonymy— 1916 Strombus (Canarium) spolongensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 2, pt. 6, p. 245, pi. 2, fig. 46 (Gunung Spolong, Java, Upper Miocene). Strombus fusiformis Sowerby, 1842 (PI. 20, fig. 30) Range— Red Sea and the western Indian Ocean. Remarks— Although rarely seen in collections, this solid, fusiform Strombus is locally common in many parts of its range. It is readily recognized by its rather heavy, fusiform shell, by its relatively long posterior siphonal canal, and by the two “humps” on the shoulder of the last whorl. It differs from the similarly shaped terebellatus in having a thick body wall and strong spiral lirae within the aperature. Although the shell has a superficial resemblance to the vittatus-campbelli group, the radula indicates that it is probably not related. Habitat— Occurs on sandy coral bottoms from 1 to 34 fathoms. It is rarely found in very shallow water, but has been occasionally washed ashore. The intestine contains white coral sand and fora- minifera. Description— Shell 26 to 45 mm. (1 to 1/i inches) in length, solid, heavy, fusiform, smoothish and with yellow-brown maculations. Whorls 9 to 10. Angle of spire 40°. Nuclear whorls 3/2, bulimoid, rapidly increasing in size, glossy, smooth, transparent and slightly rounded. Postnuclear whorls smooth, ex- cept for a single small cord below the suture which may persist to the last whorl. The spire has about 14 irregularly-spaced, low, rounded, small, whitish, former varices. Shoulder of body whorl with 2 ( rarely 3 ) rounded, low knobs, the anterior one be- ing smaller. Base of shell with about a dozen small, squarish, spiral cords which may also be present in the apertural varix. Aperture elongate, the posterior siphonal canal narrow, and extending straight up to or a little beyond the first suture. Inside of outer lip with 18 to 20 small but strong spiral lirae. Columella well-defined, straight, glossy, white to brownish pink, and with weak spiral lirae which, however, may be obsolete in the center portion. Stromboid notch usually poorly developed. Color of shell whitish cream, the early postnuclear whorls sometimes rose, the remainder of the shell mottled and speckled with light to dark yellowish brown. There may be 4 or 5 very narrow, spiral, brown- dotted bands of white on the last few whorls. Colu- mella and aperture white, rarely flushed with weak brownish pink. Periostracum very thin, smooth and translucent tan. Operculum M to )) the length of the shell, stromboid, light-brown, one edge with 9 sharp, small dentitions, the other edge straight, and with a strong central rib on the attachment side. Radular ribbon very delicate, 3 mm. long (from 33 mm. Zanzibar female shell), and with 40 rows of teeth. Formula: 2-1-2; 1-4 (plus peg) or rarely 1-3 (plus peg); 6; 8. Preserved animal cream with dark-green cobwebs and specklings on the foot, proboscis and body. Eye peduncles and tentacles yellow-chrome with white spots. Edge of mantle with a row of yellow squares. Posterior mantle fila- ment short (3 mm.). Verge short, 10 mm. in length, simple with a rather large lamellated pad at the distal end. M easurements (rn m .)— length width no. whorls 44.5 18.1 9 (large; Zanzibar) 36.5 15.0 9 (average; Zanzibar) 26.5 12.0 9 (small; Madagascar) Synonymy — 1842 Strombus fusiformis Sowerby, Thesaurus Conchyli- orum, London, vol. 1, p. 31, pf 9, figs. 91, 92 (no locality); 1843, Kiener, Coquilles Vivantes, Paris, vol. 4, Strombus, pi. 28, fig. 2; 1850, Reeve, Conch. Icon., vol. 6, Strombus, pi. 5, sp. 7. [09 - 906] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 79 Types— The holotype is in the British Museum of Natural History in London. We hereby designate Chumbi Island, west Zanzibar, as the type locality. Records (see accompanying map, pi. 55)— RED SEA: ( ANSP); Gulf of Aqaba; Jubal Id. (Issel and Canefri, f876, p. 354). SAUDI ARABIA: Shaikh Shuaib Id., Persian Gulf, 7-10 fms. ( D. Thaanum Coli n no. 4899); near Muscat in 15 fms. (Melville and Standen, 1901, p. 380); Aden (E. A. Smith, 1891, p. 419); Berbera, Gulf of Aden (USNM). ZANZIBAR: south side of Pwakuu Id., 11-18 fms. Mnazi Moja, intertidal; 1 mi. W.N.W. of Ras. Mbweni, 7 fms.; Chumbi Id., intertidal; 3 mi. w. of Ras Chukwani, 22 fms.; 2 mi. w. of Chango Ich, 15 fms.; h mi. w. of Ukombi Id., 6-9 fms. (all NSF, 1957). MOZAMBIQUE: Mozambique City (ANSP). MADAGASCAR: Pointe d’Ankify ( Dautzen- berg, 1929, p. 467); Nossi-be (A. Chavane, ANSP). IN- DIAN OCEAN IDS.: Amirantes, “Sealark” Stations F2, F7, F8, 31-34 fms. ( Melvill, 1909, p. 93). AUSTRALIA: Reeve’s 1850 record for this continent is probably erroneous. Fossil records— EGYPT: Pleistocene: beach, 50 ft. alt., Gemsah ( R. B. Newton, 1900, p. 508). Plate 55. Geographical distribution of Strombus fusiformis Sowerby. Strombus erythrinus subspecies erythrinus Dillwyn, 1817 (PI. 20, figs. 1-5) Range— For the species as a whole: Red Sea and East Africa to south Japan and to Hawaii and El- lice Islands. The subspecies rugosus Sowerby re- places the typical race in the Fiji and Ellice Islands. Remarks— This is a wide-spread species of coral waters and is usually uncommon, except in New Caledonia where it is common. It has had a con- fused synonymy and is not always correctly identi- fied in collections. It is somewhat similar to S. labiatus but is immediately distinguished by the purple-brown aperture and a columella which is white or yellowish and smooth on the outer half and purple-brown with weak yellowish teeth on the inner half. The apertural varix is usually crossed by numerous, small, rough, spiral cords. The subspecies rugosus is limited to Fiji and the Ellice Islands, but it may occur further eastward into Polynesia. It differs from erythrinus in having a shorter spire, in having 3 or 4 very large knobs on the shoulder of the dorsum, a second spiral row of much smaller knobs below these, and in having an entirely white columella and aperture. Intergrades occur in the Gilbert Islands. We cannot see any justification in considering ruppelli Reeve as a subspecies. We have Red Sea specimens before us that are inseparable from Pa- cific Ocean specimens. Dwarf and less colorful specimens have been dredged in the Philippines, Palau Islands and Hawaii. In the Palaus, the dwarf form may have a lavender or whitish nucleus and yellowish patches on the body whorl. In New Caledonia, the shells are large and darkly banded. This species is evi- dently in a state of flux. Until large series are ob- tained from many localities, it will be imperfectly understood. Habitat— This species has been dredged from 2 to 30 fathoms. In some areas, such as New Caledonia, it occurs in silty sand and algae areas just below low-water mark. In the Central Pacific it occurs either in the lagoons or outside the atolls in sparse numbers from 10 to 30 fathoms, and has been re- corded from outer reefs. Description— ( of typical erythrinus) . Shell 12 to 48 (usually about 32) mm. (/2 to % inches) in length, solid, elongate, somewhat rugose, with a purplish brown aperture and inner columella, and its exterior either whitish or marbled and banded with various shades of brown. Nuclear whorls 3, [09- 907 J 80 Canarium R. T. Abbott Strombidae the first one very small, all smooth, translucent, usually whitish, sometimes tan or brown and rarely lavender. Postnuclear whorls moderately rounded, usually with 5 to 10 incised spiral lines which may become small, rough spiral cords in later whorls; rarely smooth. After the first two postnuclear whorls, axial riblets become dominant ( 12 to 17 per whorl) and may form small rounded knobs at the slightly angular mid-sutural periphery. Spire with 4 to 10 whitish, rounded, axial, former varices. Last whorl with numerous, fine, crowded, rough spiral cords or threads. There is usually a weak, rounded, weakly beaded subsutural thread present. Shoulder of last whorl rounded and bearing about 9 to 11 small, axially lengthened knobs, those on the parie- tal wall being smallest and longest, followed by 5 or 6 rather large ones, and the two nearest the aper- tural varix being again small. Apertural varix swol- len, rust-brown, banded or whitish, and bearing numerous spiral threads. Below the row of shoulder knobs and above the base of the shell, there may be a very weak suggestion of a secondary spiral row. Color of shell very variable. Usually cream with light or dark suffusions of brown; sometimes whitish; sometimes with two narrow chocolate bands on the middle of the body whorl. Columella with about 24 indistinct spiral lirae or teeth which are usually very weak or absent in the middle por- tion. Base of columella usually brown. Inner half of columella brown or purple-brown and may have yellow spiral streaks. Outer half of columella glossy, raised, narrow, whitish or yellowish. Deep inside the aperture, the body wall is whitish to purple- brown, the latter becoming darker towards the edge of the outer lip and into the stromboid notch. There may be a white band along the inner edge of the outer lip. 20-28 fine spiral, yellowish brown, spiral threads are on the inside of the body whorl. Periostracum moderately thin, opaque, light-tan and moderately smooth. Operculum stromboid, light-brown, not arching, about % the length of the shell, with a weak central rib on the attachment side, and with 7 to 9 well-developed, sharp denti- tions on one edge. Radular ribbon delicate, similar to that of labia- tus, 3 mm. in length, amber -brown and with about 34 rows. Formula in New Caledonia specimens: 2-1-2 (rarely 3-1-3); 1-3 (plus peg); 5; 6. Preserved animal whitish with orange cobwebs and macula- tions on foot and body. Eye with one orange ring. Mantle edge with a line of tiny orange dots. Ten- tacles short and arising near the end of the eye peduncle. Verge 20 mm. in length in male shell 29 mm., long, slender, simple. Measurements ( mm.)— length width no. whorls 12.0 5.5 7.0 ( dwarf; Palau Ids. ) 23.1 10.5 8.0 ( small; Palau Ids. ) 29.3 13.9 9.0 ( average; Gilbert Ids. ) 42.8 17.3 10.0 (large; New Caledonia 48.5 19.5 10.0 (giant; New Caledonia Synonymy— 1795 Strombus erythrinus Chemnitz, Conchyl.-Cab., vol. It. pp. 146-147, figs. 1874-1875 (Red Sea). Non-bi- nomial. 1817 Strombus erythrinus “Chemnitz” Dillwyn, Descriptive Cat. Recent Shells, London, vol. 2, p. 673; 1818, Wood, Index Testae., p. 118; 1844, Duclos, Illustr. Conchy!., vol. 4, Strombus, pi. 5, figs. 16-19 [erytrhinus, errore typ.]. 1842 Strombus elegans Sowerby, Thesaurus Conchyl., vol. 1, p. 30, pi. 7, figs. 43, 48 (no locality). 1844 Strombus radians Duclos, in Chenu’s Illustr. Conchyl., vol. 4, Strombus, p. 5, pi. 4, figs. 15, 16 (les mers du Japon). 1850 Strombus ruppelli Reeve, Conch. Iconica, vol. 6, Strombus, pi. 8, figs. 13a, b (Red Sea). 1888 Strombidea erythrinus Dill., Jousseaume, Mem. Soc. Zool. France pour 1888, vol. 1, p. 174, no. 26. 1900 Canarium dentatum L., var. erythrynum Chemnitz, B Newton, Geol. Magazine, London, new series, dec- ade 4, vol. 7, pp. 508-509. 1912 Rostellaria rubicunda Perry, Matthews and Iredale, Victorian Naturalist, vol. 29, p. 10. Non Perry, 1810. 1925 Strombus (Canarium) plicatus Lamarck, Oostingh, Mededeel. Landbouw. Wageningen, vol. 29, pt. 1, pp. 59-69 (in part). ?1932 Strombus rugosus Sow., Risbec, Bull. Soc. Zool. France, vol. 57, p. 359, figs, (egg mass). ?1946 Strombus (Canarium) haemastoma Sowerby, Abrard, Ann. de Paleontologie, Paris, vol. 32, p. 62, pi. 4, fig. 29 (Pliocene, Malakula, New Hebrides). 1953 Strombus rugosus Sowerby, Dietrich and Morris, Nauti- lus, vol. 67, no. 1, pi. 4, fig. 24 (Kwajalein). Non Sowerby. Types— Dillwyn's erythrinus is based upon figures 1874 and 1875, vol. 11 of the Conchylien-Cabinet. The specimen is from the Red Sea which we desig- nate as the type locality. Sowerby’s types of elegans are in the British Museum in London. One small specimen with a lavender nucleus and smoother surface resembles our specimens from the Palaus. A large cotype, more rugose and less colorful, re- sembles our specimens from New Caledonia. N omenclature— This species has appeared in the literature most frequently as elegans Sowerby which, however, is ante-dated by Dillwyn’s valida- tion of the name erythrinus in 1817. Matthews and Iredale in 1912 suggested that Rostellaria ruhi- cunda Perry 1810 (Arcana, London, vol. 1, pi. 2, Conchology, fig. 2 from Amboyna ) was this species, but the extremely poor figure probably represents some sort of buccinid. Records (see accompanying map, pi. 56)— ZANZIBAR: off Pwakuu Id., 11-18 fms., sand, coral and sponge (NSF, 1957). SOMALIA: Djibouti (ANSP). RED SEA: Port [09 - 908] November 23, 1960 1NDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 81 Plate 56. Geographical distribution of Strombus erythrinus erythrinus Dillwyn and its eastern subspecies, rugosus Sudan (ANSP, MCZ). Aqaba, Israel (MCZ); Eilat, Gulf of Aqaba (A. Hadar, USNM). MADAGASCAR: Nossi-Fali near Nossi-be (Oostingh, 1925, p. 67). MAURITIUS: Bark- ley Id. (ANSP). SEYCHELLES: (Oostingh, 1925, p. 67). ANDAMANS: Port Blair (W. N. Carpenter, USNM). RYU- KYU ISLANDS: (Kuroda in MSS). TAIWAN: Kurun ( Ku- roda, 1941, p. 97). PHILIPPINES: off Tawi Tawi, 18 fms.; off Tinakta Id., Sulu Arch., 10 fms. (both USNM). Sisiman Cove, Bataan, 8 fms., and off Corregidor Id.; Tabaco, Luzon Id., 6-10 fms.; (all du Pont- Academy Exped., 1958, ANSP). Calapan, Mindoro Id. (Yale Peabody Mus. ). INDONESIA: Rotti Id. (Schepman, 1909, Siboga, p. 151, no. 16). ALIS- TRALIA: Queensland: Hinchinbrook Id. ( H. A. Pilsbry, ANSP); Green Id., near Cairns (MCZ). DUTCH NEW GUINEA: off Roemwakon, Aoeri Ids., Geelvink Bay, 20-25 fms. ( NSF, 1956). NEW CALEDONIA: Noumea Harbor, 1-2 fms.; Magenta, near Noumea, 4 ft.; Plage de Poe, Bou- rail, l fm.; Baie de l’Orphelinat, near Noumea; Baie de Citron, 1 fm. (all G. and M. Kline, 1958, NSF). Charron Id. (ANSP). MARIANAS: Saipan Id. (USNM). PALAU ISLANDS: Kossol Passage, 15-20 fms.; Babelthuap Id.; Yoo Passage, Eil Malk (all NSF, 1955). CAROLINES: Mog Mog, Ulithi; lagoon, Ifaluk (both USNM). MARSHALLS: Bikini lagoon, 25-30 fms.; Eniwetok; Rongerik, outer reefs; Kwajelein; 2 mi. west of Rongelap Atoll, 20 fms. ( all USNM). Ebon (MCZ). GILBERTS: Apaiang (A. Garrett, 1859, MCZ); Apamama (USNM). HAWAIIAN CHAIN: off Pokoi Bay, Oahu Id., 20 fms., coral and rubble bottom ( C. M. Burgess, coll'n.). Fossil records— EGYPT: Pleistocene: beach, 20 ft. alt., Gharib Lighthouse; beach, 50 ft. alt., Gemsah (both R. B. Newton, 1900, p. 508). FRENCH SOMALIA: Pleistocene: d'Obock and d’Hacoulta (Abrard, 1942, vol. 18, p. 63, pi. 6, fig. 37). Abrard’s (1946, p. 62) record of haemastoma Sowerby from the Pliocene of New Hebrides may be a small specimen of this species. Strombus erythrinus subspecies rugosus Sowerby, 1825 (PI. 20, figs. 11, 12) Range— Known only from Fiji, the Ellice, Samoan and Tonga Islands. Sowerby. Remarks and description— We have seen speci- mens only from Fiji and “Polynesia." Iledley re- ported it to be abundant alive in the Funafuti lagoon in the Ellice Islands, and if Schmeltz’s iden- tifications are correct, it occurs in Samoa and Tonga. This subspecies differs from the typical race in hav- ing a stouter shell, with a lower spire, in being pro- portionally broader, with an entirely white colu- mella and aperture, in having its 4 or 5 shoulder knobs on the body whorl much larger and more pointed, and in having a fairly well-pronounced second spiral row of beads around the middle of the last whorl. The outer lip is more squarely shouldered at the top. Color of shell whitish to mottled in brown, rarely banded. The spiral threads on the outer shell are well-developed. Columella usually smooth at the center. The locality of “Korea” given by Adams and Reeve in 1848 is probably erroneous. M easurements (mm.)— length width no. whorls 23.5 12.5 9 (small; Polynesia) 30.5 15.0 9 (average; Fiji) 43.0 20.1 7 + (large; Fiji) Synonymy— 1825 Strombus rugosus Sowerby, Catalogue Shells Tanker- ville, London, appendix, p. 20, no. 1791 (East In- dies); 1842, Thesaurus Conchyl., vol. 1, p. 30, pi. 7, figs. 58, 60. 1828 Strombus jugosus Wood, Index Testaceoh, Suppl., Lon- don, p. 13, pi. 4, fig. 4 (Indian Ocean). 1848 Strombus corrugatus A. Adams and Reeve, Zoology Voyage H.M.S. Samarang, London, Mollusca, p. 35, pi. 10, fig. 19 (Korea). 1899 Strombus dentatus, var. rugosus Sowerby, Hedley, Mem. Australian Mus., vol. 3, pt. 7, p. 428. Habitat— Unknown, except that Hedley reports it from the lagoon at Funafuti. [09 - 909] 82 Canarium R. T. Abbott S trombidae Plate 57. Geographical distribution of S trombus haema- stoma Sowerby and Strombus belli Kiener. S trombus scalari- formis Duclos is known only from two literature records. Types— The type of rugosus may be in the British Museum of Natural History in London, or sold at auction before reaching there. The locality of “East Indies” is probably erroneous. Records (see map, pi. 56)— ELLICE ISLANDS: Funafuti (Hedley, 1899, p. 428). FIJI ISLANDS: (A. Garrett, MCZ, ANSP); Yasawa Islands, off Viti Levu (Major Raven-Hart, USNM). SAMOA and TONGA: (Schmeltz, 1874, Mus Go- deffroy, cat. 5, p. 142). [Philippine records by Cuming are probably erroneous.! Fossil records— None reported. Strombus haemastoma Sowerby, 1842 (PI. 20, figs. 6) Range— Western Indian Ocean and Micronesia. Remarks— This is a rather rare species, which may account for the very incomplete record of its dis- tribution. There are six records from several pur- portedly reliable sources from the Indian Ocean in the collections of the ANSP, MCZ and USNM. The Marshall and Palau Island records are undisputed. S. haemastoma is very similar to belli, but differs in being more slender, in having its spire equal or greater than the length of the aperture, in having a columella which is less swollen, narrower, and with only 15 to 18 ( instead of about 24 ) weaker spiral lirae, in having a brownish purple aperture (in- stead of violet), and (in Marshall Island speci- mens ) in having a purplish apex. It is possible that S. scalariformis Duclos, 1833, is a malformed speci- men of this species. Habitat— Unknown. It has been collected dead on reefs and beaches. Description — Shell 16 to 21 mm. (M to /i inches) in length, solid, ovate-elongate, rugose, whitish with yellow maculations, and brownish to reddish violet around the aperture. Whorls 7 to 8. Nuclear whorls 3, smooth, glassy and translucent violet to whitish. Postnuclear whorls with 8 to 11 fine spiral threads which later ride over the numerous, small axial riblets ( about 20 to 25 per whorl ) . Spire with 12 to 14 small, axial, swollen, rounded, whitish former varices, the last 3 or 4 occurring in the pe- nultimate whorl. Subsutural thread weak or absent. Spiral threads on last whorl strong and 24 to 26 in number. Shoulder of whorls slightly angular. Last third of body whorl bearing 4 or 5 prominent, axially pinched knobs at the shoulder below which is a second spiral row of 4 to 6 much smaller knobs. Parietal wall with 8 to 10 neat axial ribs. Varix be- hind outer lip strongly swollen and with spiral threads. Columella relatively narrow, thickened red- to pink-brown, slightly concave and with 14 to 16 small spiral, lighter-colored lirae which are weakest at the center of the columella. Depths of aperture smooth and white. Inside of outer lip and siphonal canal with 14 to 16 strong, short spiral lirae and a band of purplish brown which is strongest on the shallow stromboid notch. Posterior anal canal is a short V-shaped channel. Periostracum, operculum and soft parts unknown. Measurements (mm.)— length width no. whorls 21.0 11.0 8 (large; Wotho Atoll) 16.0 8.0 7 ( small; Mauritius) Synonymy- 1842 Strombus haemastoma Sowerby, Thesaurus Conchyli- orum, London, vol. 1, Strombus, p. 31, no. 26, pi. 7, fig. 51 (no locality); 1850, Reeve, Conch. Icon., vol. 6, Strombus, pi. 5, figs. 5a, b; 1 844, Duclos, in Chenu’s Illustr. Conch j'l., vol. 4, pi. 4, figs. 17, 18. Types— The type is presumably in the British Museum of Natural History, London. Until exam- ined and compared with better specimens from both the Indian and Pacific Oceans, it is best that a type locality not be designated. Records (see map, pi. 57) -INDIAN OCEAN ISLANDS: Mauritius (ANSP, purchased from Sowerby post 1850; USNM, ex N. Pike and Quadras Collection; MCZ from C. [09-910] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 83 Decort). Reunion ( Deshayes, 1863, p. 114, no. o78). AUS- TRALIA: Bird Island (Queensland?) (USNM). ELLICE ISLANDS: Ujae Atoll; Wotho Atoll; Namu Id., Bikini Atoll, 3 mi. west of lieroni Id., Eniwetok Atoll (all dead, all USNM). COOK ISLANDS: Rarotonga, rare alive (A. Gar- rett, MCZ). Fossil records- Abrard’s (1946, p. 62) record from the Pliocene of New Hebrides is probably erythrinus Dillwyn and not this species. Strombus scalariformis Duclos, 1833 (PI. 58, figs. 1, 2) Range— “China Seas and Philippines. Remarks—1 To our knowledge, this shell has been collected three times, and, on the basis of its char- acters and the not uncommon occurrence of "sca- lariform” monstrosities in the family Strombidae, I am inclined to suspect that this species may be a form of haemastoma. I have not seen specimens, and only additional specimens would solve the mystery. Habitat— Unknown. Plate 58. Strombus scalariformis Duclos. China Seas. Length 22 mm. (from Kiener, 1843, pi. 14, fig. 3). Description— (irom the literature). Shell 22 mm. in length, solid, somewhat spindle-shaped, rugose, with a high spire, channeled suture and reddish violet aperture. Whorls 7 to 8. Nuclear whorls un- known. Spire tinged with violet. Possibly 8 to 12 whitish, swollen former varices on the spire. Suture well-indented and channeled. Shoulder of apical whorls angular. Spiral threads on body whorl about 24. Axial ribs prominent, possibly 14 on the last whorl. Color of shell yellowish. Columella narrow, raised, bearing fine spiral lirae and reddish violet. Inside of outer lip with stronger, fewer lirae. Strom- boid notch well developed. Soft parts unknown. Synonymy— 1833 Strombus scalariformis Duclos, Magasin de Zoologie, Paris, vol. 3, class 5, pi. 28 (les mers de la Chine); 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 14, fig. 3; 1844, Duclos, in Chenu’s Illustr. Conchyl., vol. 4, p. 2, pi. 4, figs. 9, 10. Types— The holotype was in Duclos’ collection and is probably in the Museum d’Histoire naturelle de Geneve. Type locality “seas of China.” Records— PHILIPPINES: Cebu Id.; Samar Id. (Elera, 1896, p. 253) [unconfirmed]. The collections at the Santo Thomas University in Manila were destroyed in 1945. Fossil records— None reported. Strombus belli Kiener, 1843 (PI. 20, figs. 7, 8) Range— Limited to the Hawaiian Chain. Remarks— This is one of the most attractive and distinctive Strombus of the Indo-Pacific. It is rare in most collections but is evidently not uncommon from 10 to 30 fathoms off the Hawaiian Islands. It is the only endemic full species in Hawaii, but is closely related to the rare S. haemastoma Sowerby from elsewhere in the Indian and Pacific Oceans. S. helli has been present in Hawaii since the Pleis- tocene. It is characterized by its small size (one inch or less), rotund shape, axially plicate and ean- cellate sculpture, its swollen, strongly lirate colu- mella and its bright lavender or violet aperture. Adults are quite variable in size. Specimens in the Museum of Comparative Zool- ogy from Rarotonga labelled “ helli" by Andrew Garrett are haemastoma. Habitat— This species has been dredged alive on coral sand and coral rubble bottom from 6 to 66 fathoms, and in dead condition as deep as 240 fathoms. Description— Shell 13.5 to 27.0 mm. (M to 1 inch ) in length, solid, rotund, ovate, cancellate and with a rugose, violet or lavender aperture. Whorls 9 to 10. Nuclear whorls 4, bulimoid, rounded, glossy, translucent white. First postnuclear whorl weakly malleated, the remainder bearing small, crowded axial riblets ( about 25 to 30 per whorl ) between which is a series of weak spiral threads. These be- come small cords in later whorls and number 25 to 30 on the last whorl. The axial riblets persist over the length of the parietal wall, and on the shoulder of the last third of the body whorl they form 4 or 5 moderately large, axially pinched knobs. The apex bears about 15 smoothish, white, small, swollen former varices, some of which are lined up one under the other. Suture minutely indented and bounded below by a strong, weakly beaded cord. Color of apical whorls tan with a broad spiral band of suffused brownish purple. Outer shell variable in color; usually whitish with heavy mottlings of [09-911] 84 Canarium R. T. Abbott Strombidae various shades of brown; rarely yellow-spotted. Rarely with an indistinct white band below the periphery. The outer portion of the outer lip may show violet streaks. Columella well-calloused, com- pressed, lavender, and bearing about two dozen, lighter-colored, slightly wavy, raised, rather strong, rarely bifurcating, spiral lirae. The upper 5 or 6 lirae extend into the aperture over a flattened to concave shelf. Posterior anal canal a narrow, verti- cal slit. Outer lip thin but strong, serrated. Strorn- boid notch well formed and flanked above by a small and below by a large and flaring tongue-like flange or tooth. Inside of outer lip constricted and bearing about two dozen strong lirae which may be broken into numerous, raised, small beads and bars. Depths of aperture white. Inside of outer lip witli violet and with a brown and white border. Perio- stracum thin, translucent tan, flaking off when dry. Operculum stromboid, with small serrations, and light yellow-brown. Animal whitish (preserved). In a shell 22 mm. long, the simple verge was 6 mm., the radula 3 mm. with 36 rows and a formula of 3- 1-3; 1-3 (plus peg); 6; 7. Measurem ents ( in m .)— length width no. whorls 27.0 16.3 9 + 21.5 12.0 11 13.2 0.7 8 Synonymy— 1843 Strombus hellii “Rousseau” Kiener, Coquilles Vivantes, Paris, vol. 4, Strombus, p. 59, pi. 13, fig. 2 ( mer des Indies, les cotes de file Zanzibar). 1860 S trombus cancellatus Pease, Proc. Zool. Soc. London, for 1860, p. 398 ( Sandwich Islands = Hawaiian Is- lands). Non Lamarck, 1816. Nomenclature— To our knowledge Rousseau did not publish a description of this species, and the first author is Kiener. Types— The holotype is presumably in the Mu- seum d’Histoire naturelle de Geneve. The type lo- cality of Zanzibar is probably erroneous. There is a probable cotype of S. cancellatus Pease in ANSP no. 247097. Records ( see map, pi. 57 ) ( all Hawaiian Chain ) — FRENCH FRIGATE SHOALS: Tern Id. (ANSP). NIIHAU ISLAND: (D. Thaanum, ANSP). MAUI: off Olowalu, 6-9 fms. (D. Thaanum, BPBM, USNM); off Mt. Lihau, 4-12 fms.; off Malu Bay, 4-12 fms. (both Albatross, USNM). MOLOKAI: off south coast, 43-66 fms. (Albatross Sta. 3850, USNM). OAHU: entrance to Honolulu Harbor, 6-8 fms. (D. B. Langford, BPBM): off Waikiki, 20-30 fms. ( D. Thaanum, BPBM); off Waikiki, 25-50 fms. (D. B. Kuhns, ANSP). Waianai Bay, 18 fms. (C. S. Weaver coll’n, Pele Exped. ); Keehe Lagoon, 30 fms. (C. S. Weaver, ANSP). HAWAII: off Rani Id., 235-240 fms., dead, Albatross Sta. 3982 (USNM). Fossil records— Probably Pleistocene, Kauain Stream, La- nai Id., Hawaiian Ids., 550 ft. altitude (USNM). Strombus dentatus Linne, 1758 (PI. 14, fig. 23) Range— East Africa to Polynesia. Remarks— This is one of the most widely distrib- uted and most attractive species in the Indo-Pacific, although it is never abundant and, in some areas, is considered uncommon or even rare. Its shell is highly glossed, axially plicate, rather heavy for its 1)2 inch size, and characterized by its fore-shortened outer lip which bears 3 or 4 tooth-like, white spines at the base. The columella is thick and white; the inner wall of the last whorl purplish brown with fine, white lirae; and the apex is usually lavender. It is quite variable in color, size and ribbing, and we can see no evidence of geographical subspeciation. Habitat— This is a shallow-water, coral sand- dwelling species found from the low tide mark to a depth of 20 fathoms. It is usually found near or on coral reefs, but not on the sandy-mud shores of large islands. It has been dredged dead down to 50 fathoms. Description— Shell 26 to 56 mm. ( 1 to 2 inches) in length, very solid, glossy, axially plicate, macu- lated with browns and white and with 3 or 4 white tooth-like projections at the base of the outer lip. Whorls 8 to 10; nuclear whorls 3, translucent laven- der (rarely purplish tan), glossy, smooth, well- rounded. First postnuclear whorl (and rarely the second) with 6 to 8 neatly incised, spiral lines. Re- maining whorls without spiral lines, but with micro- scopic, crowded, axial scratches. The last 3 or 4 whorls bear 6 to 14, slanting, rounded, smooth, short axial folds or ribs on the upper third of each whorl. Each of the first 3 or 4 postnuclear whorls bear 3 whitish, rounded, small, but distinct, axial, former varices which may or may not be lined up one under the other. Color of outer shell cream with irregular cloudings, maculations and flecks of chest- nut- or yellow-brown and white. Rase of siphonal canal with a diffused splotch of bluish brown. Colu- mella thick, glossy, smooth, except for 3 or 4 white lirae at the top and 3 or 4 brown lirae on the inner, lower end. Interior of aperture yellow or rose; inner wall of last whorl with a broad band of purple- brown over which are about 30 to 40 small, distinct, even, raised, white spiral lirae. Outer lip thickened, bearing below 3 or 4 distinct, tooth-like, white pro- jections. Stromboid notch obscured by these projec- tions. Ease of shell with about a dozen very weak spiral threads. Periostracum very thin and usually worn off most of the shiny shell. Operculum strom- ( large; off Oahu Id. ) ( average; off Oahu Id. ) ( small; off Oahu Id. ) [09 - 912] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 85 Plate 59. Geographical distribution of Strombus dentatus Linne. boid, light-brown, not arching, and witli about 12 small serrations. Animal and verge similar to those of arceiis. Radula ribbon amber and rose, 5 mm. in length, and with about 46 rows. Formula of Zanzi- bar specimen: 2-1-2; 1-3 (plus peg); 6; 6. Posterior mantle filament long. Measurements (mm.)— length width no. whorls 56.5 23.1 10 ( large; Palau Ids. ) 43.3 17.5 10 ( average; Okinawa Id. ) 26.0 11.5 8 ( small; Mindoro Id. ) Synonymy— 1758 Strombus dentatus Linne, Systema Naturae, ed. 10, p. 745, no. 0 (no locality); 1767, ed. 12, p. 1213, no. 513; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 285-288. 1791 Strombus tridentatus Gmelin, Systema Naturae, ed. 13, p. 3519, no. 30 (mari indico). Refers to Lister, pi. 858, fig. 14; Conchyl.-Cab., vol. 3, figs. 810-814; 1843, Kiener, Coq. Viv., vol. 4, pi. 26, fig. 2. 1798 Lambis dentata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 62, no. 790. Refers to Conchyl.-Cab., vol. 10, figs. 1501-02. 1817 Strombus samar “Chemnitz” Dillwyn, Descript. Cata- log Recent Shells, vol. 2, p. 674 (Amboina); 1823, Dillwyn, Index to Hist. Conchyl. Lister, ed. 3, p. 39. Refers to Lister, pi. 858, fig. 14. 1851 Strombus samarensis Reeve, Conch. Iconica, London, vol. 6, pi. 19, sp. and fig. 53a, b ( Philippine Islands ) . Types— According to Dodge, 1956, p. 288, there was no Linnaean type. Nor was a type locality given. Nomenclature— We are following Hanley, 1855, p. 276, Issel and T.-Canefri, 1876, p. 352, Dodge ( loc. cit. ) and most subsequent authors in apply- ing the name dentatus Linne to this species. Recent Japanese authors (Kira, Kuroda, etc.) have used tridentatus Gmelin for this species, and, erroneously we believe, applied the name dentatus to the pli- cate form of labiatus Roding. Literature records of dentatus Linne are too confused to be reliable, un- less the shell was illustrated or mention was made in whose sense the name was employed. Selected records ( see accompanying map, pi. 59; solid dots: specimens examined; open circles: literature records) — MOZAMBIQUE: Mozambique City (K. Grosch, ANSP). ZANZIBAR: Chumbe Id., 6 ft. ( NSF, 1957). RED SEA: ?Baia di Annesley (Issel and T.-Canefri, 1876, p. 353). MADAGASCAR: Nossi-be (A. Chavane, ANSP). MAU- RITIUS: ( MCZ). CEYLON: ?(Yale Peabody Mus.). JA- PAN: Osumi Ids.; Kyushu (MCZ). RYUKYU IDS.: Okinawa Id. (Mrs. A. A. Scott, ANSP). PHILIPPINES: Luzon Id.; Mindoro Id. (ANSP); Mindanao Id.; Balabac (both USNM). INDONESIA: Banda and Rotti Ids. ( Schepman, 1909, p. 153); Amboina (MCZ). CAROLINES: Yap Id. (C. O. Kile, ANSP); Truk Id. (Mrs. R. T. Gallemore, ANSP); Ponape (MCZ); Ifaluk (USNM). MARIANAS: Laulau Bahia, Sai- pan Id. (R. Sutcliffe, ANSP). MARSHALL IDS.: Bikini; Eniwetok; Rongerik; Kwajalein; Wotlio; Lae; Ujae (all USNM). HAWAII: off Waikiki, Oahu Id., 35-50 fms.; off Luniupoko Camp, Maui Id.; Keaukaha, Hilo, Hawaii Id. (all D. Thaanum, USNM); off Honolulu Harbor, Oahu Id. (D. Thaanum, ANSP). SOCIETY IDS.: Tahiti Id.: Puna- 109 - 913] 86 Canarium R. T. Abbott Strombidae auia and Atiue ( R. Robertson, 1952, ANSP). TUAMOTU ID.: Tekatikati Id.; Raroia Atoll (USNM); Marutea (Lord Hood Id. ) ( Dautzenberg and Bouge, 1932, p. 295 ) . Fossil records- ZANZIBAR: L. R. Cox (1927, p. 86) re- ports and figures (pi. 18, fig. 5) this species from the Aza- nian limestone from the base of a well near Makunduchi of Pleistocene age. It occurs in probable Pleistocene deposits, 550 feet altitude, Kawaiu Stream, Lanai Id., Hawaii (USNM). KENYA: Pleistocene: Mombasa Harbour (L. R. Cox, 1930, p. 138). Strombus fragilis (Reeling, 1798) (PI. 14, figs. 30) Range— Southwest Pacific: Ryukyus and Indo- nesia to Hawaii and Samoa. Remarks— This species is not uncommon in the central Pacific, but moderately rare to the west. It is characterized by its rather thin, but strong, elon- gate, smooth shell, and by its smooth, chocolate to red-brown columella. The dorsum of the body whorl rarely has one or two weak axial folds at the shoulder. Under a very high magnification, the sur- face of the outer shell may appear “silky” due to microscopic spiral and axial scratches. Habitat— It lives on a bottom of sand, broken coral, weed, and sponge from 2 to 25 fathoms both in lagoons and offshore ocean waters. Description— Shell 24 to 49 mm. ( 1 to 2 inches ) in length, oval-elongate, without ribs, smoothish, and moderately thin-shelled. Spire slightly concave and with smooth, well-rounded whorls. Whorls 9. Nuclear whorls 3, bulimoid, well-rounded, glossy, smooth and transparent tan. First half postnuclear whorl with 7 to 8 microscopic, spiral, incised lines, disappearing in later whorls. A small subsutural thread persists to the penultimate whorl. Surface of whorls in fresh specimens has a “silky” appearance due to microscopic axial and spiral scratches. Up- per part of spire with 5 to 11 very small, rounded, whitish former varices. Color of shell whitish to cream with large, irregular patches of dark- to orange-brown. Spire rarely tinged with light- lavender or rose. Columella solid-brown, smooth, except for 4 or 5 very weak spiral lirae at the base and rarely at the very top. Aperture light-brown within, with a dark, axial brown band near the outer lip, and with numerous, crowded, low, irreg- ular, fine, brown spiral lirae. Stromboid notch weak. Base of shell with about 20 fine, incised spiral lines. Periostracum moderately thick, translucent-brown and with a microscopically sculptured surface. Operculum stromboid, brown, broadly ovate at one end, sharply pointed at the other, not arching, with 7 well-developed dentitions, and with a well-devel- oped central rib on the attachment side. Radular ribbon 4 mm. in length, with 38 to 43 rows, brown-tinted or clear, and resembling the Plate 60. Geographical distribution of Strombus fragilis ( Roding ) . urceus and labiatus radulae. Formula from Guam and Palau Islands: 2-1-2; 1-3 (plus peg); 5; 5. Body, foot and verge cream with yellow macula- tions and spots. Proboscis tipped with brown in Palau specimen. Posterior mantle filament long or short. Verge long and slender, simple, 8 to 16 mm. in length. Measurements (mm.)— length width no. whorls 49.0 21.6 9 (large; Fiji) 38.0 15.5 9 ( average; Mindanao Id. ) 24.5 10.0 7+ ( small; Dutch New Guinea ) Synonymy— 1798 Lambis fragilis Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 62 (no locality). Refers to Conchyl.- Cab., vol. 10, fig. 1503. 1817 Strombus sainar ‘'Chemnitz” Dillwyn, Descript. Cata- log Recent Shells, London, vol. 2, p. 674 (in part by reference to Conchyl.-Cab., vol. 10, fig. 1503). See under our dentatus synonymy. ?1823 Strombus dubius Swainson, Philosophical Magazine, vol. 61, no. 301, p. 377 (no locality). Non Sowerby 1842. 1842 Strombus bulbulus Sowerby, Thesaurus Conch., vol. 1, p. 32, pi. 9, figs. 81-83. 1946 Strombus bullatus Sowerby, Dodge, Amer. Mus. Novi- tates, N. Y., no. 1314, p. 5 (error for bulbulus Sowerby, 1842). 1946 Strombus (Canarium) terebellatus Sowerby, Abrard, Annales de Paleontologie, Paris, vol. 32, p. 64, pi. 4, fig. 32 (Malekula Id., New Hebrides, Pliocene). Types— Roding’s name is based upon the Martini and Chemnitz, vol. 10, fig. 1503. The latter is based upon a specimen from the “East Indies”. We re- strict the type locality to Zamboanga, Mindanao Id., Philippines. There are three cotypes of bulbulus Sowerby in the British Museum of Natural History, [09 - 914] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 87 London. ANSP no. 39851 from the Philippines was purchased from Sowerby about 1850 and may be a cotvpe. I do not know the whereabouts of the type of dubius Swainson (non Sowerby). Nomenclature— This species until a few years ago was known as bulbuhis Sowerby, but must be known by its earlier name, fragilis (Roding). Records (see map, pi. 60)— RYUKYU IDS.: Okinawa Id. (uncommon, Mrs. A. A. Scott, ANSP). PHILIPPINES: Mindanao Id.: Zamboanga (ANSP), Samal Id., Davao Bay ( MCZ ) ; Basilan Id. (ANSP); Looc, Siburan, Negros Id. (USNM). INDONESIA: Pulo Kawassang, Paternoster Ids., n. of Sumbawa Id. (Schepman, 1909, p. 152). NEW GUINEA: off Sowek Id., Soepiori Ids.; 25-30 fms., 1 mi. east of Dauwi Id., E. Padaido Ids.; 20-25 fms., 1 mi. N.E. of Roemwakon, Aoeri Id., Geelvink Bay (all NSF, 1956). FIJI: Bega Id. (T. Dranga, ANSP); Suva Pt., Viti Levu (H. S. Ladd, USNM). MARIANAS: Piti Bay, Guam Id. (A. B. Bronson, ANSP); Managaha Id., Saipan (USNM); Apra Harbor, Guam Id. ( R. T. Abbott, USNM). PALAUS: Korak Id., Babelthuap; Gamudoko Id. (both NSF, 1955). CAROLINES: Yap Id. (USNM); Kwajalein Atoll (ANSP, USNM, Yale Peabody Mus., common); Ebon Id. (MCZ). ELLICE IDS.: Funafuti ( Hedley, 1899, p. 429). SAMOA: Pago Pago, Tutuila Id. (USNM). HAWAIIAN CHAIN: Honaunau, Hawaii Id., dead (ex J. Q. Burch, ANSP). Fossil records— Abrard, 1946, p. 64 reports two specimens (as terebellotus Sowerby) from the Pliocene from the banks of the Nua river, Malekula Id., central New Hebrides. Schepman (1907, p. 186) makes an unsubstantiated record of this ( as bulbuhis Shy.) in the post-Tertiary of the Celebes. A specimen closely resembling Recent fragilis was obtained by H. S. Ladd at Bikini Id., Bikini Atoll, Marshall Ids., drill hole 2 A, 925-935.5 ft. (Pleistocene?) (USNM). Strombus terebellatus Sowerby, 1842 (PI. 14, figs. 29; pi. 61, fig. 1) Range— East coast of Africa (subspecies afrobel- latus) to the Ryukyu Islands and to Fiji (subspe- cies terebellatus) . Remarks— This thin-shelled, smooth species is un- common on the east coast of Africa, evidently rare or absent in the central Indian Ocean and East Indies, but not uncommon in the Western Pacific. It has not been recorded from the Marshalls, Ha- waii or eastern Polynesia. Only further collecting will show whether or not this is a case of discon- tinuous distribution. The species is readily recog- nized by its smooth, slender, rather fragile shell, weak or absent columella callus and very shallow stromboid notch. It is likely to be confused with fragilis, but the latter is heavier, not as slender, with a darkly colored columella, and with fine spiral striae within the outer lip. The species may be divided into two geographi- cal races— the typical terebellatus terebellatus from the western Pacific whose last whorl descends con- siderably to produce a spire almost equal to half the entire length of the shell— and terebellatus afro- bellatus from the east coast of Africa whose spire is only one third the length of the entire shell. Habit at— Little is known of its habits. It probably lives in sand just off shore. Description— (terebellatus terebellatus). Shell 28 to 49 mm. ( 1 to 2 inches) in length, rather fragile, elongate, shiny-smooth, and mottled with browns. Whorls 9, smooth and slightly rounded. Nuclear whorls 3, smooth, glossy and translucent tan. First postnuclear whorl with 5 to 9 microscopic incised lines. Succeeding whorls smooth. There may be one or two very weak former, axial varices on the third and fourth whorls. Spire high; almost one half the length of the entire shell. Color of shell whitish to cream with heavy or weak, irregular mottlings of dark- or light-brown. In most specimens there are fine, irregular spiral lines of brown inside the aper- ture on the body whorl. Apical whorls sometimes suffused with purplish brown. Columella white with brownish flecks. Base of shell with about a dozen weak spiral incised lines. Outer lip thin, with a weak smooth varix, and a very shallow stromboid notch. Lirae absent within the aperture. Opercu- lum and soft parts unknown. Measurements (mm.)— total length spire length width no. whorls 49.2 23.0 18.0 10 45.0 20.2 16.3 9 39.6 17.5 13.5 9 36.5 14.0 13.0 8 Plate 61. Fig. 1, S trombus terebellatus terebellatus Sowerby, Mindanao Id., Philippines. Fig. 2, subspecies afrobellatus Abbott, holotype from Pange id., Zanzibar. Both x 2. [09 - 915] 88 Canarium R. T. Abbott Strombidae Plate 62. Geographical distribution of the Pacific Ocean subspecies, afrobellatus Abbott. S trombus terebellatus terebellatus Sowerby and its African Synonymy- 1842 Strornbus terebellatus Sowerby, Thesaurus Conchyl., London, vol. 1, Strornbus, p. 31, no. 30, pi. 9, figs. 84, 85 (no locality); 1843, Kiener, Coquilles Vi- vantes, vol. 4, Strornbus, p. 66, pi. 18, fig. 2, and errata p. 68 (mer des Indies). Also refers to Con- chyl.-Cab., vol. 3, fig. 810. 1843 Strornbus dentatus Wood, Kiener, loc. cit., pi. 18 ex- planation only. Non Linne 1758. Types— Sowerby 's cotypes are in the British Mu- seum of Natural History, London. We restrict the type locality to Viti Levu Island, Fiji. Records (see map, pi. 62) (for terebellatus terebellatus) — RYUKYU ISLANDS: Okinawa Id. (Mrs. A. A. Scott coll’n). PHILIPPINES: Basilan Id. (A. R. Cahn colln, ANSP); Lu- bang Id. (P. de Mesa, MCZ); Batangas, Luzon Id. (ANSP); Davao Bay, Mindanao Id. (MCZ, ANSP, USNM); Zambo- anga, Mindanao Id. (ANSP); Mambajas, Camiguin Id. (M. C. Quisumbing, ANSP); Lalay, Marinduque Id.; Cebu Id.; Jolo and Siasi Ids., Sulu Archipelago (all USNM). AUSTRALIA: Green Id., Queensland (Tony Marsh, ANSP). NEW GUINEA: Milne Bay (USNM). SOLOMONS: Treas- ury Id. (USNM, ANSP). NEW HEBRIDES: Espiritu Santo (C. M. Dumbauld, USNM). FIJI: (A. Garrett, ANSP). PALAU ISLANDS: Urukthapel Id. (NSF). CAROLINES: Truk Id. (Mrs. R. T. Gallemore, ANSP). ELLICE IS- LANDS: Funafuti, alive but uncommon ( Heclley, 1899, p. 428). Fossil records— R. Abrard’s 1946, p. 64, pi. 4, fig. 2 record of this species from the New Hebrides’ Neogene is errone- ous (see under S. fragilis). Schepman (1907, p. 186) makes an unsubstantiated report of it in the post-Tertiary of the Celebes. Strornbus terebellatus subspecies afrobellatus, new subspecies (PI. 61, fig. 2) Range— Western part of the Indian Ocean. Remarks— This subspecies differs from the typical western Pacific form in having a much shorter spire which is about one third the length of the entire shell and in lacking the weak, spiral lines of pale- brown within the aperture on the inside of the body whorl. Habitat— It has been collected on an intertidal reef in sand on Pange Island, on the west side of Zanzibar, East Africa. Description— Shell 29 to 39 mm. (1 to 1M inches) in length, rather fragile, smooth, glossy and colored cream with a heavy suffusion of light to dark brown mottlings and flecks. Similar to the typical terebel- latus (see above) but the apertural wall of the body whorl lacks the small, spiral brown color streaks, although the colors of the outer shell may show through. Columella slightly concave in the middle, slightly convex above. Stromboid notch very weak. Spire only one third the length of the entire shell. Periostracum very thin and smooth, and usually remaining only on the lower third of the shell. Operculum stromboid, slightly arching, light- brown, with 5 well-developed, sharp serrations. Ani- mal orange-yellow with white spots (preserved). Verge very long (18 mm.), narrow and simple. Radula ribbon 4 mm., delicate, and with a formula of 2-1-2; 1-3 (plus peg); 5; 5. Measurements (mm.)— total spire no. length length width whorls 30.0 8.0 11.0 9 ( holotype, ANSP ) 29.0 8.0 11.0 7 + ( paratype, ANSP ) 30.8 9.0 12.0 9 ( paratype, Coryndon Mus. ) 38.5 14.0 - - (paratype, USNM 604529) Types— The holotype is in ANSP no. 214295. The type locality is Pange Id., west side of Zanzibar, East Africa. Collected by Ostheimer, Orr and Tlior- ington, NSF station 601, in sand on intertidal reef, Jan. 20, 1957. Paratypes listed below. Records— MOZAMBIQUE: Mozambique City (K. Grotsch, ANSP 211423). TANGANYIKA: Mboa Magi, south of Dar- es-Salaam ( R. T. Abbott, LISNM). ZANZIBAR: Pange Id. (NSF, ANSP.) KENYA: Diani Beach ( H. Copley, Coryndon Mus. no. 1611). RED SEA: Jubal Id. (Issel and T.-Canefri, 1876, p. 354 [probably this subspecies]). [R. Abrard, 1946, p. 64, reports what is probably this subspecies as coming from Nossi-be, Madagascar.] Fossil records— None recorded. [09-916] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 89 Subgenus Dolomena Iredale, 1931 Type: Strombus plicatus pulchellus Reeve, 1851 Living species of this subgenus are limited to the Indo-Pacific, although fossil representatives are numerous in the Pliocene and Miocene of southeast Asia, and one species occurs in the Miocene of the Caribbean ( Strombus bifrons Sowerby, 1850; Wooding, 1928, p. 324, pi. 24, fig. 1, not pi. 23, figs. 3, 4). Members of the subgenus are absent in Ha- waii and Polynesia. The shells are usually small, with an expanded lip which has a slight posterior sinus, as well as a strong anterior stromboid notch. In many species the upper part of the lip forms a posterior canal which may be long, arching and attached to the spire. Most species have spiral lirae on the inner side of the outer lip, and some have a small amount of purple staining either on the columella or deep within the aperture. The penis has a “heel” or prong on the distal blade; the operculum is strongly ser- rated; the lateral radular tooth has a basal peg; and the inner marginal is usually much smaller than the outer marginal. Synonymy— 1931 Dolomena Iredale, Records Australian Mus., vol. 18, no. 4, p. 212. Type by monotypy: pulchella Reeve [= Strombus plicatus pulchellus Reeve]. Strombus plicatus subspecies plicatus (Roding, 1798) (PI. 18, fig. 12) Range— Red Sea. Remarks— The typical form is an uncommon spe- cies limited to the Red Sea, so far as we know. It is characterized by the full, rounded body whorl which bears 12 to 16 well-developed axial plica- tions over most of its length, by a few of the spiral plications on the columella being light-brown in color, by the absence of any color within the aperture or outer lip, and by the relatively large (2 inches) size of the shell. S. plicatus Roding appears to have four subspe- cies: 1) the nomenclatorially typical Red Sea sub- species. 2) the more common, widely distributed, Indian Ocean subspecies columba Lamarck. 3) the swollen, stunted subspecies (or possibly a form) sibbaldi Sowerby which ranges from the Gulf of Aden to Ceylon. 4) the small western Pacific sub- species pulchellus Reeve. Habitat— Unknown, but presumably in fairly shal- low water. Description— Adult shell 50 to 62 mm. (2 to 2M inches) in length, relatively thin, but strong, semi- glossy, axially plicate, and light-cream to brown- speckled in color. Spire elevated, rather acute, having an angle of about 50°. Whorls strongly shouldered, each bearing 17 to 20 small, but dis- tinct, axial ribs between which are about a dozen microscopic, spiral threads. Suture minutely in- dented, minutely waved and with a low, broad spiral cord just below. Color of shell whitish to cream with indistinct, broad spiral bands of yellow- brown flecks. The center of the last whorl may have a broad white band. Last whorl with 12 to 15 long, smooth, rounded axial ribs which are swollen at the shoulder and which become obsolescent near the outer lip. Spiral sculpture of fine, irregular threads which become larger and rounded on the base of the shell. Aperture white within. Outer body wall with about 40 spiral lirae which extend out almost to the edge of the outer lip. Columella almost straight, white, with irregular spiral, brown- colored lirae. Rase of columella with strong white lirae, and not extending much more than the lower part of the outer lip. Posterior siphonal canal short, deep, and extending up on to the next to the last whorl. Stromboid notch moderately developed. Periostracum thin, varnish-like, translucent. Oper- culum typical for the genus, % the length of the shell, light-brown, with about 10 saw-like teeth, and with a deep, narrow longitudinal trough on the outer surface. Animal and radula unknown. Plate 63. Fig. 1, S trombus plicatus columba Lamarck, Zanzibar. 2, S. plicatus pulchellus Reeve, Okinawa Island, Ryukyu Islands. Roth x 1.6. [09-927] 90 Dolomena R. T. Abbott Strombidae M east i rements (mm.)— length width no. whorls 64.0 34.5 11 ( large; Gulf of Suez) 57.0 31.0 10 ( average; Gulf of Suez 52.0 28.0 7 + ( small; Red Sea ) Synonymy- 1798 Lambis plicata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 65, no. 835 (no locality). Refers to Conchyl.-Cab., vol. 10, fig. 1496 (from the Red Sea). Not Strombus plicatus Lamarck, 1822. 1834 Strombus deformis “Gray” Griffith and Pidgeon, The Animal Kingdom— Cuvier, vol. 12, pi. 25 (facing p. 82), p. 600 (name and figure only); 1851, Reeve, Conch. Icon., vol. 6, Strombus, pi. 19, figs. 55, a, b; 1876, Issel and Canefri, Ann. Mus. Civico Storia Nat. Genova, vol. 8, p. 345. 1843 S trombus sibbaldi Sowerby, Kiener, Coquilles Vivantes, vol. 4, Strombus, p. 56, pi. 12, fig. 2. ( Not Sowerby, 1842.) 1908 Strombus plicatus Bolten, Hedley, Proc. Linn. Soc. New South Wales, vol. 33, pt. 3, p. 460. Types— Roding’s name is based upon figure 1496, vol. 10, of the Conchylien-Cabinet. Roding gave no type locality, so we designate the Red Sea from whence the Conchylien-Cabinet specimen came. Nomenclature— The name deformis has been generally used for this species. Issel and Canefri (1876, p. 346) point out that this Red Sea species was first figured by Chemnitz, but they failed to use Roding’s name. Records of deformis from Hong Kong and from Australia are evidently based upon misidentifications. Records of plicatus Roding from Australia probably were the subspecies pulchellus (see below). The plicatus Roding has nothing to do with plicatus Lamarck which was in general use a few years ago. Kiener ( 1843 ) evidently had deformis and sibbaldi interchanged. Records (see map, pi. 65)— EGYPT: Zafaran, 50 mi. south of Suez (USNM). Geb Zebara ( S. Vatakiotis, ANSP). Jubal Island (Issel and Canefri, 1876, p. 345). Fossil records— Abrard (1942, vol. 18, p. 63, pi. 6, fig. 36) records a very young “deformis Gray” from the Pleisto- cene of French Somalia. It may be the young of Strombus erythrinus Dillwyn. Strombus plicatus subspecies columba Lamarck, 1822 (PI. 18, figs. 1, 2; pi. 63, fig. 1) Range— Western half of the Indian Ocean. Remarks and Description- This is a moderately common subspecies along the East African coast, rare in Madagascar, but according to Melvill ( 1909, p. 93) is “one of the most abundant Gastropods in the regions traversed by the “Sealark” [Amirantes and Seychelles], It has not been authentically re- ported from Mauritius. The shell is brightly colored and nearly always between 1 and 1M inches in length. It differs from true plicatus in having a purple-brown blotch on the upper part of the colu- mella and an elongate, irregularly-sized blotch on the inside of the outer lip, in having the siphonal canal extend beyond the lowest part of the outer lip, and in lacking axial plications on the ventral side of the body whorl. The subspecies columba is apt to be conf used with the 'I- to 1 -inch-long sub- species pulchellus from the western Pacific. How- ever, in the latter there is always a tiny mauve blotch on the base of the columella (instead of pure white), the first 4 or 5 apical whorls lack spiral sculpturing, and the spiral lirae within the outer lip are coarser and almost always run out to the very edge of the outer lip. The apertural color blotches in columba are usually dark purple-brown (or even lavender in some Seychelles specimens), while in pulchellus they are frequently more dif- fused and more orange-brown. It is possible that this species will also turn up in the eastern Indian Ocean. In a 42 mm. Zanzibar specimen: operculum stromboid, one fourth the length of the shell, black- ish brown, not arching, with 7 to 8 well-developed serrations, and with a strong, median rib on the attachment side. Radula ribbon 3.5 mm., with 44 rows. The peg of the lateral is very small. Formula: 2-1-2; 1-4 (plus peg) or rarely 1-3 (plus peg); 5; 8. Body gray; tentacles and proboscis cream. Pos- terior mantle filament 3.5 mm. Verge 10 mm. in length, its distal blade with a well-developed “heel”. Plate 64. Fig. 1, radula of Strombus plicatus columba La- marck, Zanzibar. 2, verge. [09 - 928] November 23, 1960 INDO-PACIFIC MOLLUSC A, vol. 1, no. 2 S trombus 91 Plate 65. Geographical distribution of the subspecies of Strombus plicatus—plicatus (Roding), columba Lamarck, sibbaldi Sowerby and pulchellus Reeve. Habitat— Occurs in colonies on coral sand, grassy, broken shell and sponge bottom at a depth ranging from 3 to 44 fathoms. The “Sealark” (E. A. Smith, 1909, p. 93) obtained it on the Saya de Malha Banks at 123 fathoms, as well as at 21 to 44 fathoms. MADAGASCAR: Nossi-be (A. Chavane, ANSP); Ankify and Ambatoloaka ( Dautzenberg, 1929, p. 467). INDIAN OCEAN ISLANDS: Mahe, Seychelles (USNM); Amirantes, 32, 28, 33, 34 and 30 fms.; Saya de Malha Ranks, 123 and 47 fms.; Cargados Carajos, 30-32, 30 and 16-30 fms. (all Melvill, 1909, p. 93). [SLTMATRA: Tjalang (reported by Oostingh, 1929, no. 39, p. 2, but this may be pulchellus) . This is also possible of Hedley’s 1908, p. 460, record from Damley Island, Australia.] Fossil records— The vaguely similar Strombus deperditus J. de C. Sowerby, 1839 from the Miocene of India was re- ported under the name of columba Lamarck by Vredenburg ( 1928, p. 317) from the Upper Miocene of the Gaj of West Pakistan. I doubt if columba existed in the Miocene. Measurements (mm.)— length width no. whorls 47.0 24.0 10 ( large; Zanzibar) 41.5 23.5 10 (average; Zanzibar) 33.0 19.0 9 ( small; Zanzibar) Synonymy— 1822 Strombus columba Lamarck, Anim. sans Vert., Paris, vol. 7, p. 208 (Indes?); 1844, Duclos, in Chenu, Illust. Conchy]., vol. 4, Strombus, pi. 12, figs. 7, 8. 1823 Strombus tankervillii Swainson, Philosophical Maga- zine and Journal, London, vol. 61, no. 301, p. 377 ( no locality ) . Types— Lamarck’s type is presumably in the Museum d’Histoire naturelle de Geneve. His type locality was “Indies?”, which we now restrict to Ras Mungwe, Zanzibar. The whereabouts of Swain- son’s tankervillii is unknown, and he gave no type locality. Nomenclature— The name columba is a Latin feminine substantive noun meaning “dove”, and should not be changed to columbus while in the genus Strombus. Records (see map, pi. 65) -MOZAMBIQUE: Bazaruto Bay ( MCZ). ZANZIBAR: 1 mi. and 1% mi. W.S.W. of Ras Mungwe, 3-7 fms.; 2 mi. W. of Chango Id., 15 fms.; 2 mi. W. of Bawi Id., 15 fms.; 1 mi. N.N.E. of Pange Id., 7 fms.; 1 mi. S.W. of Nguruwe Id., 5-11 fms. (all NSF, 1957). Strombus plicatus subspecies sibbaldi Sowerby, 1842 (PI. 18, figs. 15, 16) Range— Gulf of Aden to Ceylon. Remarks and Description— This is a rare form of subspecies of plicatus. Much more material is needed to verify the suspicion that this is a mal- formation which sporadically appears in any col- ony. The shells resemble true plicatus in having a pure white interior to the apertune, but other- wise differ in having the last whorl more globose and less descending, thus giving the shell a stunted appearance and the spire a concave outline. The axial ribs are absent on the parietal portion of the last whorl, and in this respect resemble the sub- species columba. The spiral lirae on the columella are brownish purple in the three specimens we have seen, and not violet-purple as shown in Sowerby’s original illustration. American Museum of Natural History specimen no. 49426 from Ceylon is midway in character between plicatus and sib- baldi, but ANSP no. 39862 from Ceylon and pur- chased from Sowerby is of the typical sibbaldi shape. [09 - 929] 92 Dolomena R. T. Abbott Strombidae Measurements (mm.)— length width no. whorls 38.5 21.0 7 + (large; Ceylon, AMNH) 30.0 19.0 7+ (average; Ceylon, ANSP) Synonymy— 1842 Strombus sibbaldii Sowerby, Thesaurus Conchyl., Lon- don, vol. 1, p. 28, pi. 6, figs. 10, 11 (Ceylon). 1876 Strombus kieneri Issel and T.-Canefri, Ann. Mus. Civico Storia Nat. Genova, vol. 8, p. 346. New name for deformis Kiener, 1843, pi. 32, fig. 2, non Gray [Griffith and Pidgeon, 1834]. 1891 Strombus yerburyi E. A. Smith, Proc. Zool. Soc. Lon- don for 1891, p. 419, pi. 33, fig. 5 (Aden). 1843 Strombus deformis Gray, Kiener, Coquilles Vivantes, vol. 4, Strombus, pi. 32, fig. 2. Non Gray [Griffith and Pidgeon, 1834]; 1844, Duclos, Illustr. Conchyl., vol. 4, pi. 23, figs. 5-8. Types— The holotypes of sibbaldi and yerburyi are presumably in the British Museum in London. The type of kieneri would be the specimen which Kiener called deformis Gray and figured on his plate 32, fig. 2, and is presumably in the Museum d’Histoire naturelle de Geneve. The type locality for sibbaldi is "Ceylon”. Records (see map, pi. 65)— CEYLON: (ex Sowerby, ANSP; AMNH); off south coast, 34 fms. ( E. A. Smith, 1904, p. 469). ARABIA: Aden ( E. A. Smith, 1891, p. 418). INDIA: off Coromandel coast, 41 fms. (E. A. Smith, 1904, p. 469). I have not verified the record for Warrior Id., Torres Straits, Australia, 5.5 fms. (Melvill and Standen, 1899, p. 165). Strombus plicatus subspecies pulchellus Reeve, 1851 (PI. 18, figs. 3; pi. 63, fig. 2) Range— Southern Japan to Micronesia and Mela- nesia. Also Pliocene of New Hebrides. Remarks and Description— This small and attrac- tive Western Pacific subspecies is distinguished from typical plicatus and the Indian Ocean sub- species, columba, by its smaller size (usually 1 inch), absence of spiral sculpture on the first five whorls, smoothness of the central part of the colu- mella, diffused brownish orange coloring inside the aperture, and by the coarser, spiral lirae which usually fan out to the very edge of the outer lip. The spiral cord just below the suture is minute but very distinct. The base of the siphonal canal is tipped with a spot of black-brown. Rarely, the colu- mella may be all brown. Measurements (mm.)— length width no. whorls 38.5 19.0 10 ( large; Okinawa Id. ) 29.0 15.5 9 ( average; Luzon Id. ) 22.0 11.5 9 (small; Dutch New Guinea) Habitat-Dredged on coral-sand, sponge and weed bottom in depths from 8 to 50 fathoms. Synonymy- 1851 Strombus pulchellus Reeve, Conchologica Iconics, vol. 6, Strombus, sp. and fig. 52 ( Island of Ticao, Philip- pines). 1946 Strombus (Gallinula) malekulensis Abrard, Ann. Pale- ontol., Paris, vol. 32, p. 59, pi. 4, figs. 24-25 (Plio- cene, Malekula, Nua River, New Hebrides). ?1946 Strombus (Gallinula) minimus Linne, var. minor Abrard, loc. cit., p. 60, pi. 4, fig. 26 (Pliocene, Malekula, Nua River, New Hebrides). Types— The holotype of pulchellus is presumably in the British Museum in London. The type locality is Ticao Island, Philippines. Records (see map, pi. 65)— JAPAN: off Isshiki, Aichi Pref., Honshu Id., 50 fms. (T. Habe, ANSP). RYUKYU ISLANDS: between Naha and Roman, Okinawa Id., 40-50 fms. (Langford and Thaanum, ANSP); Motobu, Okinawa Id. (A. A. Scott coll’n ). TAIWAN: Taihoku-syu and Takao (Kuroda, 1941, p. 97). PHILIPPINES: east end of Corregi- dor Id., 6-10 fms.; south side of Corregidor Id., 11 fms.; cove west of Cochinos Point, 9 fms.; and east end of Sisiman Bay, all Luzon Id. (all dupont- Academy Exped., 1958, ANSP); Puerto Princessa, Palawan Id.; Cebu Id.; Sindagan Bay, Mindanao; 17 mi. northeast of Balabac Id., 44 fms., Albatross Station 5355; off Bantayan Id., 32 fms., Albatross Station 5192 (all USNM). Zamboanga, Mindanao Id. (MCZ). NEW GUINEA: 2 mi. north of Matas, Aoeri Ids.. f8-20 fms.; 1 mi. east of Dauwi, East Padaido Ids., 25-50 fms.; 1 mi. S.E. of Cape Dgarwawoffi, Japen Id. (all NSF, 1956). PALAU ISLANDS: Eil Malk (NSF, 1955). SOLO- MONS: Rabaul, New Britain Id. (USNM). MARIANAS: Pagan Id. (USNM). FIJI: off Rukua, Mbenga Id., 3-12 fms. ( Dranga and Thaanum, 1940). Fossil records— Altena (1942, p. 55) reports a specimen from the Pliocene Upper Kalibeng layers of Java, Indonesia, but I have not seen this specimen. Strombus palabuanensis Martin, 1899 (Upper Miocene of Java) may be related to this group. Abrard (see above synonymy) reports the equivalent of pulchellus from the Nua River Pliocene on Malekula Id., New Hebrides. Strombus palabuanensis Martin, 1899 Range— Upper Miocene of Java, Indonesia. Remarks— Probably allied to the Recent S. plica- tus pulchellus Reeve. Upper Miocene according to van der Vlerk, 1931, p. 246. Synonymy- 1899 Strombus (s. str.) palabuanensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 185, pi. 30, fig. 430 (Kampong Tjiodeng, Pala- buan, Java, Pliocene). Strombus deperditus J. de C. Sowerby, 1839 Range— Miocene of West Pakistan. Remarks— Although probably related to Strombus plicatus columba Lamarck, I do not agree with Vredenburg (1928, p. 317) that deperditus is a synonym of columba. [09 - 930] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 93 Synonymy— 1839 Strombus deperditus |. de C. Sowerby, Trans. Geol. Soc. London, 2nd series, vol. 5, pi. 26, fig. 19 ( Soom- row, Tertiary, India); 1854, Sowerby, d’Archiac and Haime, Deser. an. foss. gr. minim. Inde, p. 316, pi. 30, fig. 19. 1839 Strombus nodosus J. de C. Sowerby, loc. cit., pi. 26, fig. 20 ( Soomrow, Cuteh India, Tertiary). Non Bor- son, 1820. 1893 S trombus exnodosus Sacco, Molluscbi terreni Terziarii Piemonte e Lig., pt. 14, p. 5. New name for nodosus Sowerby, non Borson. 1904 Strombus sowerbyi Cossmann, Essai de Paleocon- chologie Comparee, book 6, p. 7, footnote. New name for nodosus Sowerby, non Borson. Strombus dilatatus subspecies dilatatus Swainson, 1821 (PI. 14, figs. 24, 25; pi. 66, fig. 2) Range— Singapore to the Solomon Islands; Philip- pines to Queensland, Australia. Remarks— This uncommon species is character- ized by its flaring, “tongue-like” outer lip, its smooth white columella and by the peculiar purple-brown patch of color within the somewhat constricted throat of the aperture. The spiral lirae within the apertural wall are white and usually bifurcating. The posterior siphonal canal varies in its position, rising directly up on to the spire in the typical form, but curving over far to the left in the form orosminus Duclos (fig. 24). Intergrades are not in- frequent in Philippine specimens. Old specimens have an aluminum-like glaze on the columella and the thickened outer lip. The color pattern in some specimens is rather like that of dark epidromis and lightly pigmented marginatus Septimus, to which this species is probably closely related. Habitat— Dredged in few numbers at depths of 4 to 39 fathoms on sandy-mud bottoms. Rarely cast ashore. Not associated with coral atolls. Description— Adult shell 33 to 58 mm. ( 111 to 211 inches ) in length, moderately heavy, slightly dorso- ventrally compressed, and with a smoothly rounded, flaring, “tongue-shaped” outer lip. Spire elevated, acute and with an angle of about 45°. Whorls 10 to 11, the early ones well-rounded, the last 2 to 3 usually shouldered. Nuclear whorls 3, bulimoid, rapidly increasing in size, glassy-smooth and opaque white. Remaining apical whorls purplish or tan, and finely sculptured with numerous, spiral, incised lines and numerous ( about 26 per whorl ) axial rib- lets. Body whorl with 3 to 7 low, axially lengthened knobs at the shoulder. Apex with 6 to 12 swollen, whitish, irregularly-spaced former varices. Suture finely indented, minutely waved and commonly bordered below by one or two small spiral threads. Color of outer shell whitish to tan with sparse to rarely heavy mottlings and speckles of dark-tan to yellowish brown. 4 weakly defined, white spiral bands are sometimes evident on the last whorl. Aperture somewhat constricted within, and bearing numerous, well-developed, sometimes bifurcating, spiral white rugae, and having within an elongate brownish purple patch. Outer lip broadly flaring and “tongue-like”, white and smooth along its inner border. Posterior siphonal canal long, narrow, and usually extending up on to 2 or 3 of the whorls of the spire. Occasionally, the posterior siphonal canal is curved over to form a hood-like ledge (form oros- Plate 66. Fig. 1, Strombus dilatatus swainsoni Reeve, East Indies. 2, S. dilatatus dilatatus Swainson, Loyalty Islands. 3, S. dilatatus dilatatus form orosminus Duclos, Luzon Id., Philippines. All x 1.5. [09-931] 94 Dolomena R. T. Abbott S trombidae 00° ’ l?n° 140° lftO° Plate 67. Geographical distribution of S trombus dilatatus Swainson. minus). Columella white, smooth, glossy, and with a swollen callus which runs backward on to the whorls above to border the posterior siphonal canal. Base of columella with 5 to 7 very weak teeth fac- ing the short anterior siphonal canal. Stromboid notch well-developed. Periostracum very thin, var- nish-like, smooth and translucent tan. Operculum and radula unknown. Measurements (mm.)— length width no. whorls 53.0 31.5 9 ( Cebu Id., form orosminus 51.0 29.0 10 ( large; Mindanao Id. ) 48.5 29.0 10 ( average; Mindanao Id. ) 35.5 21.0 9 ( small; form orosminus ) Synonymy— 1821 S trombus dilatatus Swainson, Zoological Illustrations, series 1, vol. 2, pi. 71 (no locality). October. Non Lamarck, 1822. 1844 S trombus orosminus Duclos, Illustr. Conchyl., vol. 2, p. 6, pi. 10, figs. 10, 11 (locality unknown). Types— The whereabouts of Swainson’s type of dilatatus is unknown. We hereby designate Upala Cay, Queensland, Australia, as the type locality. Duclos’ type of orosminus is presumably in the Mus. d’Hist. Nat. de Geneve. Records (see map, pi. 67 )— SINGAPORE: ( R. D. Pur- chon, ANSP). PHILIPPINES: Luzon Id.: San Nicolas Shoals, 7 and 10 fms.; off Corregidor Id., 6-11 fms.; Sisiman Cove, Bataan, 8 fms.; Lusong, on beach (all ANSP). Cebu Id.: near Cebu City (A. B. Franco, ANSP). Mindanao Id.: near Zamboanga (ANSP). Palawan Id. (MCZ). Panay Id.: off N.E. end, Albatross Station 5181, 26 fms. Negros Id.: off S.E. Bantayan Id., Albatross Sta. 5192, 32 fms. Sulu Archipelago: off Tataan Id., Albatross Sta. 5161, 16 fms. (all USNM). INDONESIA: off Taganar Id., Albatross Sta. 5358 (USNM). AUSTRALIA: Upala Cay, near Cairns, Queensland ( G. Sax, ANSP); Low Isles, Queensland (Tony Marsh, ANSP). SOLOMON ISLANDS: Rabaul, New Britain Id. (USNM). NEW CALEDONIA: Noumea, 4-12 fms. (G. and M. Kline, ANSP). LOYALTY ISLANDS: Mare Id. (MCZ). F ossil reco ids— U nknown . Strombus dilatatus subspecies swainsoni Reeve, 1850 (PI. 14, figs. 18, 19; pi. 66, fig. 1) Range— Unknown, but probably from Southeast Asia or the western part of Indonesia. Remarks and Description— We have seen less than a dozen specimens of this distinctive form which we believe will prove to be a good subspe- cies. It differs from the typical race in having a larger and heavier shell which rarely bears former varices in the spire and whose body whorl is cov- ered with numerous fine, spiral threads or incised lines. Adults vary in length from 54 to 64 mm. The lirae within the aperture are strong. Habitat— Unknown, but probably in muddy areas at a depth of 10 fathoms. Measurements ( mm.)— length width no. whorls 64.0 35.0 10 ( large; ANSP 39845, fig. 19 ) 58.0 30.0 10 ( average; MCZ, fig. 18) 53.0 25.0 10 (small; ANSP 39845) Synonymy- 1850 Strombus swainsoni Reeve, Conchologia Iconica, Lon- don, vol. 6, Strombus, sp. and figs. 28a, b ( no lo- cality; Cuming, coll.). Types— The type is presumably in the British Museum. There is no type locality, and until an authentic record is found one cannot be designated. ANSP 39845 (fig. 19) was purchased from Hugh Cuming about 1860, and may be from the type lot. Nomenclature— Reeve in 1850 figured this sub- species, thinking that it was the true dilatatus Swainson, and, because he erroneously thought that Swainson’s name was preoccupied by Lamarck’s 1822 dilatatus, he intimated that he was proposing the new name swainsoni. However, Reeve misiden- tified Swainson’s dilatatus, and the name should not have appeared in the synonymy under his valid de- scription and figure of swainsoni. Records— No accurate records exist. One speci- men from the Mus. Comp. Zool. is labelled “East Indies”. Fossil records— Unknown. [09 - 932] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 95 Strombus clilatatus subspecies taiwanicus Nomura, 1935 Strombus dilatatus subspecies fennemai Martin, 1899 Ra nge— Fossil (Pliocene) of Taiwan (Formosa) and ( ? ) Philippines. Remarks— This subspecies closely resembles the Recent subspecies swainsoni Reeve, and may well be its progenitor. It differs, however, in being nar- rower, in having a less flaring outer lip and in hav- ing fewer and more pronounced spiral incised lines on the body whorl. The tiny axial riblets are stronger and fewer in the early whorls. The length of specimens varies from 51.0 to 28.0 mm. Nomura records it from a number of localities in the Byoritu Beds of Taiwan: Hakusyaton; Wangwa; Rinsuikwa; Bosiho; Keiyukwa; Siko, etc. which he calls Plio- cene in origin. The type locality is 1000 meters east of Hakusyaton, station 20, Taiwan. Holotype, Reg. no. 53163, Tohoku Imperial Univ., Sendai. Dickerson’s (1921, Philippine Journal of Science, vol. 18, p. 5; also vol. 20, p. 202, pi. 5, fig. 6) reports what may be this subspecies ( as swainsoni ) from the Miocene [Pliocene?] of Bondoc Peninsula, Tayabas Prov., Luzon Island, Philippines. MacNeil (1960, pi. 12, figs. 14-15, 22-23) records it from the Nakoshi sand, the Naha limestone and Gabusoga, all Tertiary of west Okinawa Id. Synonymy— 1935 Strombus taiwanicus Nomura, Science Reports Tohoku Imperial Univ., Sendai, 2nd ser., vol. 18, no. 2, p. 177, pi. 8, figs. 15a, 15b, 16a, 16b (Byoritu Beds, Taiwain ) . 1960 Strombus (Labiostrombus) cf. japonicus Reeve, Mac- Neil, Tertiary and Quaternary Gastropods of Oki- nawa. U. S. Geol. Survey Prof. Paper 339, pi. 12, figs. 14, 15, 22, 23 (seen in MSS). Plate 68. Strombus fennemai K. Martin. Pliocene of Ceram Island, Indonesia, (from P. J. Fischer, 1927, pi. 212, figs. 24a, b). Natural size. Range— Pliocene of Java, Sumatra, and Timor, Indonesia. Remarks— The figures of the types are almost identical with specimens of S trombus plicatus co- Jumba Lamarck from the Indian Ocean, and not like the subspecies pulchellus Reeve which now lives in southwest Pacific area. S. fennemai differs from coliimba in having a slightly more rotund body whorl, in lacking a well-developed, lirated upper third of the columella, and in having a poorer development of axial riblets in the spire. Synonymy- 1899 Strombus (s. str.) fennemai K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 181, pi. 29, figs. 418-420 ( Sonde, Padasmalang, Java, Pliocene). 1942 Strombus (Labiostrombus) fennemai Martin, Altena, Leidsche Geolog. Mededeel., vol. 12, p. 50 ( Sema- rang, Java; Kendeng Beds, East Java, Pliocene; Atjeh, Sumatra, Pliocene; Poetjangan layers, Bareng beds, Bodjonegoro, Java). Strombus rembangensis Martin, 1899 Range— Lower Miocene of Java and Borneo. Remarks— This may be a malformed adult of one of the fossil subspecies of dilatatus. We have a similar appearing Recent specimen from New Cale- donia whose peculiar shape was caused by shell in- jury. Until more specimens are available, it would be difficult to assign rembangensis to its nearest relative. Synonymy- 1899 Strombus (s. str.) rembangensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 180, pi. 29, fig. 417 (Sedan in Rembang, Java). 1947 Strombus (Labiostrombus) rembangensis Martin, Beets, Geologie en Mijnbouw, 9th year, no. 3, p. 41 (Lower Miocene, Pulu Balang, East Borneo). Strombus labiosus Wood, 1828 (PI. 18, figs. 17, 18) Range— East Africa to the Ryukyus and the East Indies. Remarks— This species is rarely collected, except in dredge hauls when it appears to be relatively common. It is readily recognized by its rotund body whorl, by its flaring outer lip which is usually turned upward at the top, by the minutely cancel- late early whorls and by the white columella and outer lip. About half of the known specimens have a dozen or so brownish purple spiral lines deep [09 - 933] 96 Dolomena R. T. Abbott Strombidae £ > ' §. ilfc Plate 69. Figs. 1 and 2, Strombus labiosus Wood, off Taganak Id., Borneo. Both x 1.5. within the aperture, while others are colorless. Habitat— Dredged in 6 to 44 fathoms on muddy or sand and broken shell bottom. Description— Shell 25 to 51 mm. ( 1 to 2 inches) in length, solid, obese, with a flaring outer lip, and lead to brownish gray in color. Whorls 10 to 11. Nuclear whorls 2/2, small, elevated, rapidly increas- ing in size, transparent, glossy and smooth. First postnuclear whorl with about 10 microscopic spiral threads which in succeeding whorls cross about 2 dozen very small, rounded axial ribs, thus giving a microscopically cancellate appearance. In remain- ing whorls the axial ribs become increasingly larger and become knobbed at the periphery in the last whorl. Color of outer shell cream, tan, lead-gray or yellowish, sometimes splotched with a weak pur- plish brown and usually glistening. Columella and aperture enamel white. In some specimens there is a patch of a dozen or so purple-brown spiral lines deep within the aperture. Spiral sculpture absent on the ventral surface of the body whorl, but con- sists of numerous coarse threads on the last third of the whorl. Periphery of last whorl with 4 to 7 small rounded knobs. Outer lip wing-like and turned up- ward at the posterior end. Siphonal notch broadly U-shaped. Posterior siphonal notch short. Interior of outer body wall with numerous, white, spiral lirae which do not run all the way to the edge of the outer lip. Columella slightly concave, callus- like, with numerous spiral, fine lirae which are weak or absent on the middle of the columella. Lower third with about 10 fine spiral teeth. Perio- stracum very thin, smoothish, translucent tan. Soft parts not available for study. Measurements (mm.)— length width no. whorls 50.8 30.0 7+ ( large; “Philippines” ) 34.5 21.0 11 (average; Taganak Id., Borneo) 25.5 15.0 9 (small; Taganak Id., Borneo) Synonymy— 1828 Strombus labiosus Wood, Supplement to Index Tes- taceologicus, London, p. 54, pi. 4, fig. 3 ( name and figure only); 1842, Sowerby, Thes. Conchyl., vol. 1, Strombus, p. 27, pi. 6, figs. 15, 16 (no locality); 1843, Kiener, Coq. Vivantes, vol. 4, pi. 22, fig. 2. 1851 Strombus labiosus Gray, Reeve, Conchologica Iconica. London, vol. 6, pi. 18, fig. 50 (Cagayan, Island of Mindanao, Philippines, 25 fms). 1940 S trombus (Labiostrombus) labiosus Gray, Beets, Ge- ologie en Mijnbouw, 21 Jaargang, no. 2, pp. 17-25, fig. 1 ( bionomics of shell ) . 1947 Canarium ( Labiostrombus ) labiosus Wood, Wissema, Thesis, Leiden, p. 97 ( fide Cox, 1948, p. 28, 70). Not seen by us. Types— The holotype is presumably in the British Museum of Natural History in London. Wood gave Plate 70. Geographical distribution of Strombus labiosus specimens examined. Wood. Open circles indicate literature records; solid dots, [09 - 934] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 97 no type locality. We now select Bataan, Luzon Id., Philippines, as the type locality. Nomenclature— A number of authors have errone- ously attributed Gray as the author. Records (see map, pi. 70 )— MOZAMBIQUE: Port Amelia (USNM). ZANZIBAR: 2 mi. west of Chango Id., 15 fms. (NSF). MADAGASCAR: Tamatave ( Dautzenberg, 1929, p. 470). INDIAN OCEAN ISLANDS: Mulaku and South Nilandu Atolls, Maidive Islands (E. A. Smith, 1903, vol. 2, p. 612). RYUKYU ISLANDS: beach at Buckner Bay, Oki- nawa Id. (A. A. Scott, ANSP ) . PHILIPPINES: Luzon Id.: east end of Corregidor Id., 6-10 fms.; west side of Cochinos Pt., Bataan, 9 fms. (both du Pont -Academy Exped., 1958, ANSP); off Malavatuan Id., 18 fms., Albatross Station 5276. Cebu Id.: near Cebu City (USNM). Leyte Id.: off Tacbuc Pt., 48 fms.. Albatross Station 5477; west of Bucas Grande Id., 44 fms.. Albatross Station 5235 (both USNM). INDO- NESIA: Keledjitan, Bantam, Java Ich; off Taganak Id., Borneo Id., 39 fms., Albatross Station 5358 (both USNM). DUTCH NEW GUINEA: 1 mi. S.E. of Cape Dgarwawoffi, Japen Ich, 10-16 fms. (NSF, 1956). Fossil records— Cox (1948, p. 29) quotes Wissema’s 1947 thesis as recording typical labiosus as occurring in the Plio- Pleistocene of Nias Island (west side of Sumatra), Indo- nesia. See also C. Beets, 1940, pp. 17-25). Tesch (Palaeon- tologie von Timor, 1920, pi. 129, fig. 164 a, b) figures this species (as fennemai K. Martin) from the Pliocene of Timor. Plate 71. Strombus labiosus Wood. Holotype of var. teschi Cox from Dent Haven, British North Borneo. Pliocene. ( from Cox, 1948, pi. 2, figs. 8a, b). Natural size. Strombus labiosus subspecies teschi Cox, 1948 Range— Fossil (Pliocene), Java, Timor, Ceram, Philippines. Remarks— Of the several characters mentioned by Cox as distinguishing this shell from the Recent labiosus, only one seems to hold true: there are no tubercles at the shoulder-angle on the dorsal side of the last whorl. The other characters, such as the 48 degree angle of spire, conspicuous striations on the inner side of the wing and of the inner lip, and the details of spiral ornamentation are all repre- sented in living specimens. Wissema (1947, p. 97) notes that typical labiosus does exist in the Plio- Pleistocene of Nias Island (west side of Sumatra), Indonesia. Type locality: 7 km. inland from Dent Haven, Dent Peninsula, Borneo (Pliocene). Synonymy— 1948 Strombus ( Labiostrombus ) labiosus Wood var. teschi Cox, Schweizerische Palaeontologische Abhandl., vol. 66, p. 28, pi. 2, figs. 8a, b (Borneo, Pliocene). Strombus rutteni Altena, 1942 Range— Fossil (Pliocene) from Java, Indonesia. Remarks— In spire and columella characters this species most closely resembles Strombus labiosus Wood but its body whorl is not as rotund, nor the wing of the outer lip as high. The subsutural cord persists to the end of the last whorl, but this fea- ture may also occur in some living specimens of labiosus. Length of shell 43 mm., width 25 mm. Type locality: Padasmalang (Sheet 9313), Java: Upper Kalibeng layers, Pliocene. Paratypes from Doekoepengkol, Madioen, Java. Synonymy— 1942 Strombus (Labiostrombus) rutteni Altena, Leidsche Geologische Mededeelingen, vol. 12, p. 53, figs. 15a, 15b (Pliocene, Java). Plate 72. Strombus rutteni Altena. Holotype from the Plio- cene of Java, Indonesia, (from Altena, 1942, p. 53, figs. 15a, b). x 1.5. Strombus marginatus Linne, 1758 Until much more material is collected along the shores of Southeast Asia from India to Japan, this species will remain a puzzle with regards to the distribution of its forms and/or subspecies. True marginatus of Linne was the earliest name applied to this complex, but unfortunately is the peculiar form bearing a strong, sharp, smooth spiral keel on the shoulder of the last two whorls. This keeled form occurs in the coral-water areas of northern Ceylon, northwest Sumatra and central Burma. A subspecies, or possibly only an ecologic form, oc- curs in the non-coralline areas of the mainland of [09 - 935] 98 Dolomena R. T. Abbott Strombidae Plate 73. Geographical distribution of the races of S trom- bus marginatus Linne. Ceylon. This is the elongate, glossy form, lacking a peripheral keel and having a single rather well- developed nodule on the dorsum near the shoulder. Linnaeus named this succinctus in 1767. Ranging northward from the Gulf of Siam to at least Hong Kong, and reappearing in Japan proper, is the sub- species robustus Sowerby which is a heavier, more rotund shell with a tendency towards short plica- tions or several nodules on the shoulder. A third subspecies seems quite recognizable in the warm waters of the great Southwest Pacific Island arc running from the southern Ryukyus, the Philippines and New Guinea to New Caledonia. This is the small, somewhat laterally compressed, darkly-colored subspecies Septimus Duclos. We have kept the synonymies and treatment of these forms or subspecies separate, until more is understood of this species. Strombus marginatus subspecies marginatus Linne, 1758 (PL 18, figs. 6, 7; pi. 74, fig. 1) Range— Known only from the areas surrounding the Bay of Bengal. Remarks— This shell is characterized by the sharp, narrow, spiral carina or keel on the shoulder of the last 2 or 3 whorls. In most specimens the keel dis- appears in the area behind the outer lip. The body whorl may be smooth or entirely covered with nu- merous fine, but distinct spiral, incised lines. Inter- grades exist in Burma between this form and the unkeeled robustus. Habitat— Found below the low tide line in sandy silt and green algal bottoms not far from coral reefs. Von Martens (1887, p. 189) reports it on mud banks at low tide and at four fathoms in the Mergui Archipelago, off Burma. Description— Adult shell 39 to 57 mm. ( lfi to 2fi inches) in length, solid, rotund, with its narrowing posterior canal arching up on to the spire, and with a strong, angular, sharp spiral keel on the shoulder of the last two whorls. Color of shell whitish with broad and narrow, broken, spiral bands of light- to dark-brown. Nuclear whorls 3, small, elevated, smooth and translucent-tan. Next 3 or 4 postnuclear whorls flat to slightly convex, with numerous, long, crowded, axial riblets (27 to 31 per whorl). In the last 2 or 3 whorls these riblets disappear or are re- duced to tiny beads set on the shoulder keel just above the suture. In the last whorl the keel be- comes smooth. Spiral sculpture in the spire of 8 to 15 fine threads. Spire with 0 to 8 small, swollen, whitish, former varices. Base of shell with about a dozen sharply incised lines which may also be pres- ent over the entire body whorl. Columella white, nearly straight, slightly swollen, and smooth, except for about a dozen weak, broken, wavy, raised lirae at the top and 4 to 10 tiny dentitions at the base. Outer lip thin, incurled, sinuous in side view, and arching up and over to the left on to the spire [09 - 936] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 99 Plate 74. Immature shells of Strombus. Fig. 1, S. mar- ginatus marginatus Linne, Ceylon. Fig. 2, S. marginatus Septimus Duclos, Philippines. Fig. 3, S. minimus Linne, Philippines. All x 2. where it ends at the first ( rarely the second ) suture above. Stromboid notch very shallow, sometimes only an undulation. Inside of outer lip with about 3 dozen fine, irregular or broken, raised white lirae. Periostracum thin, smoothish and translucent yel- lowish. Operculum stromboid. Radular ribbon 4 mm. in length, wine-red, with 35 rows of rather delicate teeth. Formula of a Cey- lon specimen: 2-1-3; 1-3 (plus peg); 5; 6. Proboscis and eye peduncles brownish maroon witli white spots. Sides of foot weakly suffused with brown. Tentacles rather short. Verge 20 mm., stout, with a broad, “heeled” distal blade; its stem dusted with brownish orange and small white spots. Posterior mantle filament short (2 mm.). Measurements (mm.)— length width no. whorls 57.0 29.9 10 ( large; Northern Ceylon ) 45.2 27.5 9 ( average; Gulf of Manaar ) 39.0 26.0 7+ (small; “East Indies”) Synonymy— 1758 Strombus marginatus Linne, Systema Naturae, ed. 10, p. 744, no. 430 (no locality); 1767, ed. 12, p. 1209, no. 499; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ilf, art. 3, pp. 262-264; 1842, Sowerby, The- saurus Conch., vol. 1, pi. 6, fig. 17; 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 16, fig. 2; 1851, Reeve, Conch. Icon., vol. 6, Strombus, pi. 18, fig. 49. 1798 Lambis carinata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 62, no. 779. Refers to Conchy!. -Cab., vol. 10, figs. 1489, 1490 and vol. 3, fig. 816. Type— It is agreed by most workers (see Dodge, 1956, pp. 262-264) that Linnaeus did not have a type specimen, nor could he locate a published figure. Knorr’s pt. 3, pi. 13, fig. 4 could be either marginatus or succinctus. The earliest of the recog- nizable figures is what Schroter called marginatus Linne (Einleit. Conchylien. Linne, vol. 1, p. 431, pi. 2, fig. 10; 1783). Subsequent workers have agreed Plate 75. Fig. 1, radula of Strombus marginatus marginatus Linne, Ceylon. 2, verge. that Linne’s brief description does point to this spe- cies. No locality was given. The shell borrowed by Linne was probably from Ceylon, an island we now designate as the type locality. Records— INDIA: Pamban, Gulf of Manaar (E. Thurston, 1895, p. 125). CEYLON: Pearl Bank, Gulf of Manaar (Kline, ANSP); north tip of Eluvativu Id. (Kline, ANSP). BURMA: Maungmagon, north of Tavoy (MCZ). SLT- MATRA: Oedjoeng Batee Kapal; Lam Baro, Atjeh Head ( Oostingh, 1929, no. 39, p. 2). [Philippine records are un- confirmed.] Fossil records- None reported. Strombus marginatus subspecies succinctus Linne, 1767 (PI. 18, figs. 13, 14) Range— Ceylon to Madras, India. Remarks— This is a distinct shell limited to the non-coralline waters of Ceylon and the southeast end of India. Insufficient material is at hand to de- cide for certain whether this represents a very localized subspecies or an ecologic form. The shell is more elongate and paler in color, and is char- acterized by a single, rounded, distinct knob on the dorsum of the body whorl at the shoulder. The pale brownish yellow body whorl has 4 or 5 narrow white bands which are delicately flecked with brown. Flabitat— Below low water mark in muddy sand in a sheltered bay (G. and M. Kline in Ceylon); dredge in muddy sand off Madras in several feet of water (Crichton, 1940, p. 203). Description— Shell 40 to 53 mm. ( lfi to 2 inches) in length, smoothish, moderately elongate, and weakly patterned with soft yellow-brown and white. Whorls 10 to 12. Nuclear whorls 2%, small, elevated, glossy, translucent-white. Apical whorls with numerous (about 30) axial riblets which are crossed by about a dozen microscopic, spiral [09 - 937] 100 Dolomena R. T. Abbott Strombidae threads. The first 3 or 4 postnuclear whorls bear a total of 4 to 8 swollen, whitish, rounded, former varices. Suture minutely indented; in the early whorls it is bordered below by a distinct spiral, striated cord; in the third-to-last whorl the suture is commonly bordered above by small beads or nodules. Body whorl usually smooth, except for a few weak spiral threads at the top and about a dozen incised lines at the base. Ventral side of body whorl smooth and flattened; dorsal side, near the shoulder, with one small, but prominent, rounded, low nodule. Color of outer shell a light yellow-brown consisting of very fine reticulated and arrow-shaped lines. Body whorl bears 4 (occa- sionally a 5th at the base) spiral white bands which are sparsely overlaid with weak arrow- shaped brown lines. Aperture elongate, white and spirally striated within. Parietal callus white, slightly swollen and weakly wrinkled at the top. Outer lip sinuate, sharp, its edge curling inward slightly; posterior canal long, adhering to 2 or 3 whorls in the spire, and extending straight up. “Stromboid notch” weak. Periostracum thin, var- nish-like and transparent. Operculum and soft parts not known. Measurem cuts (mm.)— length width no. whorls 55.2 22.0 10 (large; “Indian Ocean”) 47.1 20.8 10 (average; Nilaveli, Ceylon) 38.3 18.1 9 (small; Trincomalee, Ceylon) Synonymy— 1767 Strombus succinctus Linne, Systema Naturae, ed. 12, p. 1212, no. 509 (In India); 1855, Hanley, Ipsa Linnaei Conchylia, London, p. 274; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 278-280. 1767 S trombus accinctus Linne, Systema Naturae, “13th ed.” in Vienna (reform. Holmiensen), p. 1212; 1958, Iredale, Proc. Royal Zool. Soc. New South Wales, for 1956-57, p. 61; 1778, Born, Index Rerum Nat. Mus. Caes. Vindob., pt. 1, p. 280. 1768 Strombus succinctus Linne, Systema Naturae, ed. 12, vol. 3, "Errata”, (for uccinctus ). 1777 Strombus succinctus Linne, Martini, Conchyl.-Cab., Nurnberg, vol. 3, p. 104, pi. 79, fig. 815 (but not 816); 1843, Kiener, Coquilles Vivantes, vol. 4, Strombus, pi. 10, fig. 2 (excellent); 1842, Sowerby, Thesaurus Conchyl., vol. 1, pi. 6, fig. 20. Types— The type locality is “In India” which we further restrict to Madras, India. Hanley ( 1855, p. 274) states that the type is in the Linnaean col- lection and implies that it is figured in Sowerby (1842, vol. 1, pi. 6, fig. 20). None of Linne’s figure references refer to this subspecies, all of them be- ing septimns Duclos, except for Seba, pi. 62, fig. 20 which is vittatus Linne. Records (see map, pi. 73)— INDIA: Madras (Crichton, 1940, p. 203). CEYLON: Kachcheri Bay, Trincomalee, Ceylon (G. and M. Kline, NSF); 12 miles north of Trin- comalee (W. E. Old, Jr., ANSP); Ara Pt., Nilaveli (H. G. Deignan. USNM); Pearl banks, Gulf of Manaar ( R. Tonk- lass, ANSP). Strombus marginatus subspecies robustus Sowerby, 1874 (PI. 18, figs. 8, 9; pi. 76) Range— South China Sea to southern half of Japan. Remarks— This is a quite variable shell, not only within a single colony, but also in certain char- acters which show a geographical cline. It is per- haps best characterized by its rotund shape (not elongate as in succinctus, and not laterally com- pressed, as in Septimus ), and varying number (1 to 12) of short plications or elongate nodules on the shoulder of the last whorl. In almost every speci- men the plication bordering the left side of the parietal wall is the largest and longest. The upper part of the aperture extends up over two sutures at least, sometimes straight up towards the apex, and rarely may arch over to the left. The apex may be pink, purplish or whitish. In Japan the species is abundant and the shell attaining its maximum size of 67 mm. It is also abundant in the Gulf of Siam where the length of the shell ranges from 26 to 50 mm. The MCZ contains two specimens from Sarawak, Borneo, which we refer to as the “ rotund form.” This shell (see pi. 76) is pyriform, smoothly Plate 76. Figs. 1 and 2, Strombus marginatus robustus Sowerby, bulbous form from Sarawak, Borneo. Both x 1.5. [09 - 938] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 101 rounded at the shoulder, with one weak plication bordering the left side of the parietal wall; finely and microscopically spirally striate and with its posterior canal arching far over to the left. This might prove someday to be a valid subspecies. Habitat— On sandy to sandy mud bottoms from 1 to 25 fathoms near shore. In Kyushu Island, Japan, it is abundant. Description— Shell 29 to 67 mm. (1 to 4 V> inches) in length, rotund and broadly elongate. Color yellow-brown with irregular white marking. Similar to marginatus succinctus and marginatus septimus, but the body whorl is more rotund; the shell gener- ally larger; commonly ( although not always ) with spiral threads over the entire last whorl; and with the long posterior siphonal canal arching up over 1 to 3 sutures. The left border of the ventral (pari- etal) side of the body whorl characteristically has a peculiar axial swelling. The shoulder on the dor- sal side may have one small node and on the ven- tral side may rarely have a series of short, small axial plications. In some Japanese specimens the interior may be weakly flushed with light-violet. Periostracum thin, translucent yellowish and smoothish. Operculum and soft parts not available for study. Measurements ( mm.)— length width no. whorls 67.1 37.2 8+ (large; Kyushu Id., Japan) 44.1 24.7 10 ( average; Hong Kong ) 29.0 16.0 9 ( small; Koh Chang, Thailand ) Synonymy — 1874 S trombus robustus Sowerby, Proc. Zool. Soc. London for 1874, p. 599, pi. 72, figs. 5, 5a (Hong Kong). 1899 Strombus septimus Duclos, Crosse and Fischer, Journ. de Conchyl., vol. 37, p. 287 (Annam). Not Septi- mus Duclos. 1959 Labiostrombus succinctus Linne, Kira, Coloured II- lustr. Shells of Japan, Osaka, 2nd ed., p. 36, pi. 15, fig. 14. Types— Sowerby’s holotype of robustus is in the British Museum of Natural History in London. The type locality is Hong Kong. Nomenclature— Workers on the Japanese fauna have in the past considered robustus Sowerby a synonym of succinctus Linne. We consider the lat- ter to be an Indian Ocean subspecies of margina- tus Linne, and robustus an eastern Asian sub- species of marginatus. Records (see map, pi. 73)— BURMA: Sandoway (MCZ). THAILAND: Koh Chang; Koh Samet; Khan Nu Paknam (all Gulf of Siam, USNM); Ban Ao Moo, Bang Ko Chai, Chantaburi Prov. (G. Moore, MCZ). HONG KONG: south of Lema Id., 25 fms. (A. J. Staple, ANSP). BORNEO: Sarawak (rotund form MCZ). JAPAN: Honshu Island: Sagami Bay, Kanagawa Pref. (A. R. Calm, ANSP); Kii, Wakayama Pref. (MCZ). Shikoku Island: Tosa Bay, Kochi Pref. (ANSP). Kyushu Island: Moeshima, Kagoshima Bay (T. Habe, ANSP); Miyazaki Pref. (T. Kuroda, 1935, p. 47, no. 142). Fossil records— Schepman (1907, p. 186) reports this sub- species from the post-Tertiary of the Celebes. Strombus marginatus subspecies septimus Duclos, 1844 (PI. 18, figs. 10, 11; pi. 74, fig. 2) Range— Ryukyu Islands southward through the Philippines, East Indies and New Guinea to New Caledonia. Remarks— The shells of this subspecies are more brightly colored and with a more flaring lip, which gives the shell a slightly flattened or dorso-ventral compression. The color is generally dark-brown with 4 to 6 narrow, white spiral bands which bear numerous arrow-shaped spots of dark-brown. The upper end of the outer lip, or posterior canal, usu- ally ascends only to the first suture above, and not up to the second or third suture, as in robustus. The dorsal hump, so prominent in succinctus, is reduced or absent in septimus. Habitat— This subspecies has been collected on coral sand and rubble bottom in 4 to 16 fathoms of water. It is commonly washed ashore after storms. We have not found it on coral reefs nor in pure mud areas. Abundant in the Philippines and the Solomons, but rare in the Ryukyus and southern Indonesia. Description— Shell 26 to 48 mm. ( IK to 2 inches) in length, broadly elongate and colored with dark- chestnut to rich yellow-brown. Similar to margina- tus succinctus, but is not as elongate; its lip is broader and more flaring; the posterior siphonal canal extends up to only the first suture; the shoulder of the body whorl lacks the single node but may have one to several very small nodules or beads. Color of shell dark-chestnut to light-brown with 5 to 7 narrower white spiral bands which are crossed by darker, more numerous, arrow-shaped bars of brown. Apex with 9 to 11 whitish former varices. Periostracum moderately thin, translucent- brown. Operculum stromboid, with 5 to 6 serra- tions. Verge maroon-brown with white dots and a “heel” on the distal blade. Radula 2 to 3 mm., with 37 to 42 rows of teeth. Formula variable: 2-1-2 (also 3-1-3); 1-3 (plus peg), also 1-5 (plus peg); 5 to 7; 6 to 9. Animal like marginatus. [09 - 939] 102 Dolomena R. T. Abbott Strombidae Measurements (mm.)— length width no. whorls 48.5 25.0 10 4- ( large; Dutch New Guinea ) 42.6 23.4 10 (average; Solomon Islands) 30.2 15.0 10 (small; Luzon Id.) Synonymy— 1844 Strombus Septimus Duclos, Illustr. Conchyl., vol. 4, p. 7, pi. 13, figs. 9, 10, pi. 15, fig. 11, pi. 26, fig. 2 ( locality unknown ) . 1885 Strombus succinctus var. Septimus Duclos, Tryon, Manual of Conchology, vol. 7, p. 117. Types— Duclos’ type is presumed to be in the Museum d’Histoire naturelle de Geneve. No local- ity was given, and we hereby designate Lusong, Bataan Peninsula, Luzon Island, Philippines, as the type locality. Nomenclature— Tryon (1885, p. 117) considered Septimus as a variety of succinctus. Most other workers have considered it to he merely a form and synonym, but with a series of accurately local- ized material we consider it to he of subspecific rank. The pre-Linnaean figures are mostly of this subspecies: Rumphius, pi. 37, fig. x; Gualtieri pi. 33, fig. B; Seba, pi. 61, fig. 15 (hut not 20). Records (see map, pi. 73)-RYUKYU ISLANDS: Buck- ner Bay, Okinawa Id., rare (Mrs. A. A. Scott, ANSP). PHILIPPINES: Luzon Id.: Sisiman Cove; Lusong; Bataan; east end of Corregidor Id.; San Nicolas Shoals Light, Ma- nila Bay (all du Pont- ANSP expedition, 1958). Masbate Id. (ANSP). Basilan Id.; Dupolog, Mindanao Id.; Puerto Princessa, Palawan Id. (USNM). INDONESIA: Amboina Id. ( MCZ ) . NEW GUINEA: Oro Bay (ANSP); off Cape Dgarwawoffi, and off Samberbaba, both Japen Id. (NSF). SOLOMONS: Kieta, Bougainville Id. (W. J. Eyerdam, ANSP). NEW CALEDONIA: Bourail ( Mme. Reverce, ANSP). Fossil records— None reported. Fossil Relatives of marginatus A number of Tertiary species have been de- scribed from the East Indies which are undoubt- edly closely related to and the possible progenitors of either Septimus and/or robustus. I have not had the opportunity to examine sufficient fossil material to hazard a scheme of relationships. One species from northeast Borneo (Pliocene) conld probably be considered as rather closely resembling the sub- species, septimus, i.e. togopiensis Cox, 1948. These fossil species, subspecies or forms are: Strombus togopiensis Cox, 1948 Range— Pliocene of northeast Borneo, Indonesia. Remarks— Probably closely related to S. margina- tus septimus Duclos. Synonymy— 1948 Strombus (Labiostrombus) togopiensis Cox, Schweizer- ische Palaontologische Abhandl., vol. 66, p. 27, pi. 2, figs. 10a, b (Dent Peninsula, Borneo, Pliocene). Plate 77. Strombus togopiensis Cox. Holotype from Dent Haven, British North Borneo. Pliocene, (from Cox, 1948, pi. 2, figs. 10a, 10b). Natural size. Strombus sedanensis Martin, 1899 Range— Lower Miocene of Java, Indonesia, and Lower Miocene of West Pakistan. Remarks— The characters of the spire and the carination of the shoulder in the area of the parie- tal wall suggest an affinity with the Recent mar- ginatus group. However, the rather thick, rounded outer lip and the strong tubercles on the shoulder of the body whorl are particularly curious. I would be inclined to accept this as a full species. I. van der Vlerk, 1931, p. 247 calls the Rembang beds Lower Miocene. Synonymy- 1899 Strombus (s. str.) sedanensis K. Martin, Samml. geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 180, pi. 29, figs. 416, 416a (Sedan in Rembang, Java); 1928, Vredenburg, Mem. Geol. Survey India, Cal- cutta, vol. 50, pt. 1, p. 313 (Gaj beds, near Karachi). Strombus javanus Martin, 1879 Range— Miocene of Java, Indonesia. Remarks— This species was based upon an in- completely preserved specimen, but from the characters of the outside of the outer lip and its size, it could be somewhat likened to the Recent Strombus marginatus septimus Duclos. Miocene ac- cording to van der Vlerk, 1931, p. 246. Smith’s sem- peri is possibly a synonym. Synonymy— 1879 Strombus javanus K. Martin, Die Tertiarschichten auf Java, Leiden, p. 47, pi. 9, fig. 2 (Java, Tertiary). Prior to Dec. 1879. 1879 Strombus sumatranus H. Woodward, Geol. Mag., Lon- don, new series, decade 2, vol. 6, p. 543, pi. 14, fig. 19 (Tertiary Clay-marl, West Coast of Sumatra). Dec. 1879. 1900 S trombus javanus var. semperi W. D. Smith, Philip- pine Jour. Science, vol. 1, pi. 3, fig. 3, p. 629 ( Upper Miocene, Loboo River, Batangas Prov., Lu- zon Id., Philippines). [09 - 940] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 103 Plate 78. Figs. 1 and 2, Strombus sedanensis Martin. Lower Miocene of Java, (from K. Martin, 1899, pi. 29, figs. 416, 416a). Fig. 3, Strombus javanus Martin. Type from the Miocene of Java (from K. Martin, 1879, pi. 9, fig. 2). All natural size. Strombus variabilis subspecies variabilis Swainson, 1820 (PI. 14, figs. 21, 22) Range— Sumatra and Thailand, eastward through Micronesia, Melanesia, northern Australia to Samoa. Remarks— This species is rather variable in size, shape and color pattern. Insufficient material is available to establish any clear-cut subspeciation, but we would point out the presence of three forms. Typical variabilis from the western part of the range is the largest ( usually about 2 inches in length), generally has a brown spot on the col- umella, has numerous, short, dark-brown, zigzag axial lines on the outer shell, and has well-shoul- dered apical whorls. At the east end of the range the supposed subspecies athenius is smaller (usu- ally about 1)1 inches in length), is lighter-colored, with a more rounded spire, much heavier shell, and rarely has a brown patch on the columella. A dark-banded color form (pi. 14, fig. 20) occurs sporadically in the middle of the range. Habitat— Just below the low tide mark in weedy, sandy mud to 25 fathoms in weed and coral sand. Description— (of the typical form) Shell 33 to 60 mm. (1)1 to 2)1 inches) in length, solid, mode- rately light, with a well-knobbed, shouldered spire, and winged outer lip. Color of shell enamel-white to cream with a heavy flecking and suffusion of dark- to yellow-brown, mainly consisting of numer- ous, crowded, short, zigzag lines. The last whorl has 5 narrow, indistinct, spiral, white bands. Col- umella and aperture smooth and milky white. Most specimens have a distinct, dark-brown, ob- long color patch on the center of the columella. Whorls 9 to 10. Nuclear whorls 3, the first being minute and elevated, the remaining being rounded, glossy, smooth, and translucent-tan. First three postnuclear whorls with 10 to 12 microscopic, squarish spiral threads. The fourth and later whorls bear 13 to 16 evenly-sized and evenly-distributed rounded knobs midway between the suture, this giving the periphery of the whorl an angular or shouldered appearance. Ventral side of body whorl smooth, slightly keeled above; dorsal side with 3 or 4 knobs, the next to last being largest, and the earlier ones being axially elongated. There are 8 to 9 whitish, rounded, swollen former varices in the first 4 postnuclear whorls. Base of shell with about IS indistinct spiral threads. Columella smooth, rarely with a few very weak plications at the base. Inner lip usually smooth, but rarely with a few lirae near the base and stromboid notch. Upper end of outer lip with a short posterior siphonal groove and a shallow notch. Periostracum rather thin, translucent-tan, usually worn away. Opercu- lum stromboid, strongly arching, light-brown, about one third the length of the shell, and with 7 strong serrations. Radula ribbon 4 mm., light-tan and with about 42 rows of teeth. Formula 2-1-2; 1-2 (plus peg); 4; 4. Verge with a prominent thumb-like append- age on the broad, distal blade. Measurem ends (mm.)— length width no. whorls 60.0 30.0 9 + ( large; Cebu Id. ) 49.0 25.0 10 ( average; Balabac Id. ) 33.0 18.5 9 (small; Balabac Id.) Synonymy— 1820 Strombus variabilis Swainson, Zoological Illustrations, series 1, vol. 1, pi. 10 (South Seas) (spotted variety from India ) ; 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 21, fig. 2; 1844, Duclos, in Chenu’s Illustr. Conchyl., vol. 4, pi. 11, figs. 9, 10; 1850, Reeve, Conch. Icon., vol. 6, Strombus, pi. 10, fig. 21c and d. 1829 Strombus lituratus Menke, Verzeichniss Conchyl. - Samml. Malsburg, Pyrmont, p. 58, no. 1205 ( no locality); 1871, Mtirch, Malakozool. Blatter, vol. 18, p. 127. Types— Swainson’s type is probably lost. He evi- dently had a mixture between the typical variabilis and the smaller athenius. The fact that he speaks of “numerous undulated short lines of darker col- our” and a shell “two inches and a quarter long” leads us to believe he meant the Philippine form which we have figured on plate 14, fig. 21 and 22. For similar figures, see those mentioned in the above synonymies. We restrict the type locality to Cebu Island, Philippines. [09-941] 104 Dolomena R. T. Abbott Strombidae Plate 79. Figs. 1 and 2, immature and adult S trombus variabilis variabilis Swainson, Luzon Id., Philippines. 3 and 4, adult and immature S. variabilis athenius Duclos, Dutch New Guinea. All x 1.5. Records (see map, pi. 80 )— THAILAND: Koh Samet; Koh Chang (USNM); Ban Pe, Rayong Prov. (MCZ). IN- DONESIA: Poelo We, Atjeh, w. Sumatra Id. (Oostingh, 1929, no. 39, p. 2); Bouro Id., Moluccas (MCZ); Woda Id., Halmahera (MCZ). MALAYA: Singapore (USNM). PHILIPPINES: Mindoro: San Jose; Tilic Bay, Lubang Id. (both MCZ); Luzon: Tabaco, Albay Prov. (du Pont -Acad- emy Exped., 1958); Legaspi Bay (ANSP); Palawan: Bala- bac and Cuyo Id. (ANSP); Cebu: Bantayan (ANSP); near Cebu City (A. B. Franco, ANSP); Bohol: off Jagoliao Id., 2 fms. (E. Zambo, ANSP); Mindanao: Zamboanga (ANSP); Sulu Archipelago: Sanga Sanga Id. (John Root, ANSP). CAROLINES: Peleliu Id., Palau Ids. ( NSF, 1955). AD- MIRALTY ISLANDS: 16-25 fms. ( R. B. Watson, 1886, “Challenger,” p. 420). SOLOMONS: Buin, Bougainville Id. (MCZ; ANSP). AUSTRALIA: Yirrkala, Arnhemland; Groote Eylandt, Gulf of Carpenteria (both USNM); Sweers Id., Gulf of Carpenteria (MCZ); Queensland: Green Id., Batt Reef, and Low Id. (MCZ; ANSP). [Melvill and Sykes, 1899, p. 44, record “Andamans”, but this needs verification.] Fossil records— Pliocene : Nias Id., off Sumatra, Indonesia ( Icke and Martin, 1907, pp. 214, 239, pi. 15, figs. 23, 23a (this is based upon a young specimen which only resembles an immature variabilis) . Pliocene: Upper Kalibeng layers, Java, Indonesia, according to Altena (1942, pp. 54, 55). Quaternary of East Borneo: Poeloe Boenjoe (Beets, 1950, vol. 15, p. 244). Strombus variabilis subspecies athenius Duclos, 1844 (PI. 14, fig. 20; pi. 79, figs. 3, 4) Range— Northern New Guinea, Marshalls to Samoa and New Caledonia. Remarks— Insufficient material is available for a more complete understanding of this subspecies. It is possible that it represents a stunted ecological form. It is characterized by its much heavier shell, more rounded apex, absence of numerous, small, axial lines of dark-brown, its lighter color (which is all-white in Samoa and the Gilberts ) , and by the more weakly shouldered apical whorls. Its body whorl usually has 5 very broad light orange-brown bands. 10 of 57 specimens examined have the small dark patch on the columella, while in the typical variabilis from the Philippines 90 per cent of sev- eral dozen specimens have the brown columellar patch. Its length is between 26 and 43 mm. Animal similar to that of variabilis. Verge with a less devel- oped thumb-like appendage. Radula formula: 2-1- 2; 1-3 (plus peg), rarely 1-2 (plus peg); 4 or 5; 4, 5, or 6. A third kind which appears to be a color form (our plate 14, fig. 20, p. [09-833] ) has turned up in the Palau Islands, the Solomons, New Caledonia and northeast Australia. It is characterized by 5 or 6 very dark, even, chocolate-brown spiral bands. Reeve (1850) figures it in his pi. 10, fig. 21a from Darnley’s Island, Australia. The sculpturing and the shape of the shell is more like true variabilis, but in color pattern it is more like athenius. Measurements (mm.)— length width no. whorls 43.0 24.0 8 + ( large; Dutch New Guinea ) 36.5 20.0 9 (average; Dutch New Guinea) 27.0 15.5 9 (small; Dutch New Guinea) Synonymy— 1843 Strombus variabilis Swainson, Kiener, Coquilles Vi- vantes, Paris, vol. 4, pi. 21, fig. 2a (not fig. 2); 1850, Reeve, Conch. Icon., vol. 6, pi. 10, fig. 21b (not others). 1844 Strombus athenius Duclos, in Chenu’s Illustr. Conchyl., vol. 4, p. 7, pi. 11, fig. 2 (probably not fig. 1). Lo- cality unknown. Types— We hereby designate as the lectoholotype the specimen figured on plate 11, fig. 2 in Duclos’ S trombus monograph of 1844. We restrict the type locality to Riak Island, Dutch New Guinea. N omenclature— Duclos described this species on the basis of several good characters, as mentioned above. However, he was not aware that some speci- mens may have a brown columellar patch, a fea- ture which he erroneously attributed solely to [09 - 942] Plate 80. Geographical distribution of S trombus variabilis variabilis Swainson and its eastern subspecies, athenius Duclos. variabilis. It is possible that his figure 2 is a true variabilis. Records (see map, pi. 80)— DUTCH NEW GUINEA: ofl Rani Island, Schouten Ids., 1-3 fms.; off Rouw Id., Aoeri Ids., 1 fm.; Matas Id., Aoeri Ids., intertidal; off Mios Wo- endi, Padaido Ids., 3-8 fms.; 2 mi. west of Korido, Soepiori Id., reef flat (all NSF, 1956). NEW CALEDONIA: Dge (Cockerell, ANSP); Laregnere Reef, E. of Noumea, 2 fms.; Noumea, 4-12 fms.; barrier reef, Touho Bay (all G. and M. Kline, NSF, 1959). FIJI: Suva (ANSP). MARSHALL IS- LANDS: Bikini; Rongelap; Majuro; Arno lagoon (all USNM); Kwajalein (Yale Peabody Mus.). GILBERTS: Apiang ( MCZ). SAMOA: Tutuila Id. (ANSP). CARO- LINES: Truk Id. (Mrs. R. T. Gallemore). Fossil records— None recorded. Strombus minimus Linne, 1771 (PI. 18, figs. 4, 5; pi. 74, fig. 3) Range— Ryuku Islands to Indonesia and east- ward through Melanesia to Fiji. Remarks— This species is locally abundant in the middle of its range, but rather uncommon to the north and to the east. It is evidently absent from small islands or coral waters. It is readily recog- nized by its small, thick, heavy shell, chrome- yellow aperture, and by the swollen columella and thick posterior siphonal canal. No subspecies have been recognized, except Abrard’s Pliocene minor which we believe is pnlchellus Reeve. Habitat— Lives in large colonies from the low tide mark down to 12 fathoms, sometimes on coral sand and weed bottom, at other times in muddy, dirty, shallow water. Its sparsity in the Ryukyu Islands, Fiji and New Caledonia probably means that it does not live in very shallow water in those areas. They are cast up on many Philippine beaches after storms. Description— Shell 14 to 40 mm. (usually about 30) in length (about 1 inch), very heavy and solid for its size, with a swollen columella, with the edges of its posterior siphonal canal swollen, and with a chrome-yellow aperture. Color of outer shell dark- to light- brown with minute flecks of cream. Last whorl with one, rarely two, spiral rows of white squares. Parietal wall usually cream to whitish. Whorls 9; nuclear whorls 3, the first very small and elevated, the remainder rounded, glossy, smooth and translucent brown to tan. Next 4 post- nuclear whorls with about 15 microscopic, squar- ish, spiral threads which cross over the 9 small, rounded, whitish former varices. Axial sculpture begins in the 5th whorl in the form of about 17 elongated, shouldered knobs. The smooth ventral wall of the body whorl is swollen on its left by an elongate axial ridge, and followed anteriorly by 2 or 3 small shoulder knobs. Posterior siphonal canal slightly S-shaped, extending up over 2 su- tures, and with 2 very thickened, flattened edges. Columella very swollen, white, smooth, but rarely with 2 to 4 weak lirae at the base. Outer lip smooth within, except for a dozen very weak, short lirae at the base. Interior of aperture chrome-yellow. Base of shell with about 10 weak spiral threads. Strom- boid notch very shallow. Periostracum moderately developed, translucent-tan. Operculum stromboid, slightly less than one third the length of the shell, and with 8 well-developed, sharp serrations. Radula ribbon delicate, translucent-tan, 2 mm. in length (shell 29 mm.), and with about 33 rows. Formula for one Luzon specimen: 3-1-3; 1-4 (plus peg); 5; 7. Another Luzon specimen: 2-1-2; 1-3 (plus peg); 5; 6. Tentacles and posterior mantle filament long. Animal with red-brown maculations and dustings of black. Verge unknown. [09 - 943] 106 Dolomena R. T. Abbott S trombidae Plate 81. Strombus minimus Linne, Luzon Island, Philip- pines. Fig. 1, dorsal view of head, showing eye peduncles and proboscis. 2, operculum. 3, posterior comer of mantle margin, showing 2 mm. -long, fleshy appendage. 4, 3-mm. terminal part of gelatinous egg mass removed from pre- served female. 5, radular teeth. Measurements (mm.)— length width no. whorls 14.0 9.0 5 + (small; Luzon Id.) 41.0 21.5 10 (large; Cebu Id.) 31.5 18.0 10 (average; Luzon Id.) Synonymy — 1771 Strornbus minimus Linne, Mantissa plantarum— regni anim. appendix, p. 549 (In India orientali); 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, p. 298-299; 1851, Reeve, Conch. Icon., vol. 6, pi. 18, fig. 47. 1798 Lambis minimus Gmelin, Roding, Museum Bolteni- anum, Hamburg, pt. 2, p. 65, no. 836. Refers to Conehyl.-Cab., vol. 10, figs. 1491, 1492. 1822 Strombus troglodytes Lamarck, Anim. sans Vert., vol. 7, p. 209 (Grandes Indes). Refers to S. minimus Linne; Conchyl.-Cab., vol. 10, figs. 1491 and 1492; and others. Plate 82. Geographical distribution of Strombus minimus Linne. 1843 Strombus troglodites Lam., Kiener, Coquilles Vivantes, vol. 4, pi. 31, fig. 2. Types— According to Dodge (1956, p. 298), Lin- naeus type is not in existence. We hereby restrict the type locality to Cebu City, Cebu Island, Philip- pines, from whence our figures 4 and 5 specimens come. Records (see map, pi. 82)— RYUKYU ISLANDS; Buchner Bay (A. A. Scott, ANSP). TAIWAN: Taihoku-syu ( Kuroda, 1941, p. 97). PHILIPPINES: abundant throughout the archipelago: Luzon; Mindanao; Cebu; Panay; Palawan; Mindoro; Negros; Camaguin; and Marinduque Ids. (ANSP; USNM; MCZ). INDONESIA: Amboina (MCZ); Isle of Dol, N. Loloda Ids. (MCZ); Riouw Ids.; Banka Id.; Cele- bes; Flores; Timor (all Oostingh, 1923, p. 83). NEW GUINEA: Dauwi Id., E. Padaido Ids. ( NSF, 1956); Finseh- hafen (MCZ); Samarai and Oro Bay (ANSP); Seleo Id., Aitape Id. (USNM). SOLOMONS: ' Rabaul, New Britain Id. (USNM); Guadalcanal Id. (MCZ). NEW HEBRIDES: Espiritu Santo Id. (MCZ). NEW CALEDONIA: Touho Bay (G. and M. Kline, NSF, 1959). FIJI: off Rukua, Bega Id., 3-12 fms. (D. Thaanum); Suva Pt., Viti Levu Id. (H. S Ladd, USNM). Fossil records— INDONESIA: Pliocene of Sonde, Java (K. Martin, 1899, p. 182, pi. 29, fig. 421). Abrard’s 1946, p. 60 record for the Pliocene of New Hebrides is probably pul- chellus Reeve. [09 - 944] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 107 Subgenus Labiostrombus Oostingh, 1925 Type: Strombus epidromis Linne, 1758 Because of anatomical and shell characters, we are considering Labiostrombus, with its sole living species, epidromis Linne, as a subgenus distinct from the many-species subgenus Dolomena Ire- dale. Labiostrombus, in the past, has been applied to species which we now consider to be members of Dolomena ( minimus , marginatus, pulchellus, etc. ) . Three fossil species of Labiostrombus are known: Strombus leurus Wooding, 1928, from the Miocene of Jamaica, West Indies; denti Cox, 1948 from the Pliocene of Borneo; and kemedjingensis Martin, 1916, from the Lower Miocene of Java. Conchologically, the shells are characterized by having the upper part of the outer lip rising evenly, without a posterior sinus, on to the whorl above. The shell is rather light-weight and thin, the col- umella smooth and axial sculpture usually poorly developed. The main cusps in the central and the elongate lateral sometimes have very tiny sub-den- ticles. The marginals are long and slender; the lat- eral has a small basal peg. In general, the radular teeth are very delicate and loosely attached to the odontophore. The very long prong-like penis has three peculiar divisions at the distal end. The eye stalks and tentacles are long, and the operculum has many small serrations. Synonymy— 1847 Gallinula “Klein” Herrmannsen, Indicis Generum Malacozoorum Primordia, vol. 1, p. 461. Not validly proposed (name only). Non Brisson, 1760. 1852 Gallinula “Klein” Morch, Cat. Conch, de Yokli, p. 61. Type hereby selected: S trombus epidromis Linne, 1758. 1854 Gallinula “Klein H. and A. Adams, Genera of Recent Mollusca, vol. 1, p. 259. (Invalid type designation by Kobelt, 1878, Illust. Conchyl., p. 105: s. Isabella Lamarck [not in original list by name]); [Tryon, 1885, vol. 7, p. 101 and Oostingh, 1925, p. 58 both gave the invalid type designation of succinctus Linne]. 1868 Gallinula H. and A. Adams, American Journal of Con- chology, vol. 4, pt. 3, p. 139. Type by listing first species, campbelli; invalid designation. 1904 Gallinula “Klein” Cossmann, Essai de Paleoconch. Compar., Paris, 6th book, p. 9. First valid designa- tion of type: Strombus epidromis Linne. 1925 Labiostrombus Oostingh, Mededeelingen van de Land- bouwhoogeschool Wageningen, vol. 29, pt. 1, p. 58. New name for Gallinula H. and A. Adams, non Brisson, 1760. Type [invalid] by original designa- tion: succinctus Linne. New type designation hereby made: Strombus epidromis Linne. Nomenclature— The first valid introduction of Gallinula for mollusks was made by Morch in 1852. Tryon’s type designation of succinctus Linne is in- valid, since that species is not listed by any previ- ous authors in Gallinula. Cossmann’s 1904 designa- tion of epidromis Linne is the first valid designa- tion. Oostingh’s Labiostrombus, a new name, will have to take this type also. Herrmannsen’s 1847 name is not defined nor contains any species by name. It is also a homonym of the bird genus Gallinula Brisson, 1760. Strombus epidromis Linne, 1758 (PI. 17, figs. 17; pi. 83, figs. 1, 2) Range— Ryukyu Islands and Singapore to Queens- land and New Caledonia. Remarks— This handsome species is sparsely dis- tributed throughout its moderately restricted range, but evidently occurs in large colonies in certain very localized areas. For its three-inch size, the shell is rather light, has a large, rounded, flaring outer lip, and is characterized by a smooth, enamel- white aperture and columella. Older specimens may have an aluminum-like glaze on the lips. The upper part of the outer lip may either just reach to the first suture above or extend up to the second suture. The radula and the verge are both very un- usual (see under description). Plate 83. Figs. 1 and 2, Strombus epidromis Linne (im- mature), Dutch New Guinea. Both x 2. [09 - 955 J 108 Labiostrombus R. T. Abbott Strombidae Although no fossil specimens have been found, closely related species are recorded from the Plio- cene and Upper Miocene of Indonesia. Habitat— Lives in muddy to sandy bottoms from 1 to 16 fathoms. Fine coral sand and foraminifera have been found in the intestine. The species has not been found on small coral atolls, but rather is associated with larger islands. Description— Shell 53 to 90 mm. (2 to 3/2 inches) in length, relatively thin-shelled but strong, with a smoothish last whorl, with a large, flaring, rounded outer lip, and with a smooth, white aperture. Whorls 10 to 11, rounded to shouldered in the apex and only slightly convex in the last. Angle of spire about 50° and with a dozen whitish former varices. Nuclear whorls 2, rounded, elevated, glossy, smooth, translucent and either whitish or lavender. First 4 postnuelear whorls with about 10 squarish, minute, spiral cords which cross about 20 small, crowded, rounded axial riblets. The spiral cords disappear on the lower whorls, and the axial riblets are reduced to a spiral series of small, even, knobs on the now squarish shoulder of the whorl. Last whorl glossy, smooth, except for 3 or 4 long, weak, axial plications. One or two of the latter usually have a small, low, rounded knob. Suture even, minutely indented, and, in the apex, bounded below by a weak spiral cord. Aperture ample, smooth and enamel-white within. Columellar callus weak, smooth and white. Posterior siphonal canal short; stromboid notch broadly U-shaped. Base of shell with about a dozen, weak spiral threads. Outer shell whitish to cream with weak mottlings, speckles and rarely with flames of yel- low-brown. Spire rarely lavender. Periostracum rather thin, rather smooth and grayish to trans- lucent-tan. Operculum stromboid, chestnut to black-brown, arching about % the length of the shell; attachment side with a strong, central rib; convex edge with 12 to 14 small, narrow, curved serrations. Radular ribbon proportionately small (5.5 to 6.0 mm. in length and 1.5 mm. in width), with weakly attached, tan to amber teeth in 55 to 60 rows. Cen- tral ovoid to quadrate with a simple or tricuspid central cusp flanked on each side by 2 or rarely 3 smaller cusps. Lateral thin, quadrate, with a small basal peg, with its largest cusps usually bearing 1 or 2 tiny dentitions. Outer marginal long and bent. Formula: 2-1-2 or 3-1-3; 1-4 (plus peg); 5; 6 (New Caledonia and Dutch New Guinea speci- mens examined). Body stromboid; eye peduncles with purplish brown background and white spots; proboscis with Plate 84. Strombus epidromis Linne, Dutch New Guinea. Fig. 1, central. 2, lateral tooth. 3, inner marginal. 4, outer marginal. 5, verge, showing enlargement of distal end. 6, operculum. dark, narrow, transverse color lines. Tentacles long. Verge long and narrow (30 mm. in length; shell of male, 68 mm.). Terminal end three- pronged. Measurements (mm.)— length width no. whorls 89.0 49.5 6 + ( large; Cebu Id. ) 74.0 42.0 10 (average; Dutch New Guinea) 53.5 32.1 9 ( small; New Caledonia) Synonymy— 1758 Strombus epidromis Linne, Systema Naturae, ed. 10, p. 745, no. 436 (In O. Asiae); 1767, ed. 12, p. 1211, no. 506; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 273-275. 1798 Lambis epidromis Gmelin, Roding, Museum Bolteni- anum, Hamburg, pt. 2, p. 65, no. 834. Refers to Conchyl.-Cab., vol. 3, fig. 821. 1885 Strombus expansa “Martini”, Tryon, Manual of Con- chology, Pliila., vol. 7, p. 138. Types— “ The specimen marked for epidromis in the Linnaean collection in [Linnaean Society of London] London is the epidromis of all authors and may therefore be accepted as the type specimen.” (Dodge, 1956, p. 274). We hereby restrict the type locality to Amboina, Indonesia. Records (see map, pi. 85)— SINGAPORE: (R. D. Pur- chon, ANSP). RYUKYU ISLANDS: Buckner Bay, Okinawa Id. (A. A. Scott, ANSP). PHILIPPINES: off Corregidor Id., Luzon Id., 6-10 fms. (du Pont- Academy Exped., 1958). Cebu Id.: Bantayan (E. Zambo, ANSP); near Cebu City (A. B. Franco, ANSP). Bohol Id.: Jagoliao Id., 2 fms. (du Pont- Academy Exped., 1958). Cuyo Id., Palawan Prov. (ANSP). Mindanao Id.: Mindanao (ANSP). Sulu Archi- pelago: Bongao Channel. Sanga Sanga Id. (John Root, ANSP); Jolo Id. ( USNM ) ; Tataan Id., 16 fms., Albatross Station 5161 (USNM). INDONESIA: Amboina (MCZ). November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 109 Plate 85. Geographical distribution of Strombus epidromis Linne. DUTCH NEW GUINEA: Mios Woendi Isle, Padaido Ids. ( NSF, 1956). AUSTRALIA: Queensland: Cooktown (Tony Marsh, ANSP), Bundaberg (W. E. Old, Jr., ANSP). NEW CALEDONIA: Dge (Cockerell, ANSP); Baie de l’Orpheli- nat, 1 fm., mud; Bourail; Noumea, 4-12 fms. (all G. and M. Kline, NSF, 1959). Fossil records— None recorded. Strombus sp. may be the young of denti or an epidromis- like species. Synonymy— 1948 Strombus (Labiostrombus) denti Cox, Schweizerische Palaeontologische Abhandlungen, vol. 66, art. 2, p. 29, pi. 2, figs. 4a, b, 5 (Pliocene, Borneo). Plate 86. Strombus ( Labiostrombus ) denti Cox. Dent Haven, British North Borneo. Pliocene, (from Cox, 1948, pi. 2, figs. 4a, b, holotype, fig. 5, immature, paratype). Natural size. Strombus denti Cox, 1948 Range— Fossil (Pliocene) of northeast Borneo, In- donesia. Remarks— This species appears to be related to epidromis, but differs in having long axial riblets in the spire, lacking the angulation of the spire whorls, and in having a less expansive outer lip which reaches up to the second suture above. Whorls 12, length of shell 90.5 mm. Type locality: 7 km. in- land from Dent Haven, Dent Peninsula, northeast Borneo, Indonesia. Cox points out that Strombus deningeri Fischer, 1921, based upon an immature specimen from the Pliocene of Ceram, is suspi- ciously akin to young specimens of denti. The lat- ter may be a subspecies or synonym of deningeri Fischer. Dickerson’s (1922, vol. 20, pi. 5, fig. 5) Strombus kemedjingensis Martin, 1916 Range— Lower Miocene (West-Progo beds), Java, Indonesia. Remarks— Based upon a badly broken specimen. Martin likened it to vittatus Linne and Isabella Lam. [= canarium Linne], but I would suggest that it is allied to the denti Cox and epidromis Linne complex. Lower Miocene according to van der Vlerk, 1931, p. 247. Synonymy- 1916 Strombus (Gallinula) kemedjingensis K. Martin, Samml Geol. Reichs-Mus. Leiden, n. Folge, vol. 2, pt. 6, p. 246, pi. 2, figs. 47, 48 ( LTpper Miocene, Kali Kemedjing, Java); 1928, Leidsche Geol. Meded., vol. 3, pt. 2, p. 126. [09 - 957] 110 Labiostromhus R. T. Abbott Strombidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [09 - 958] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 111 Subgenus Doxander Iredale, 1931 Type: Strombus vittatus Linne, 1758 Whether one considers Strombus vittatus Linne, campbelli Gray, and japonicus Reeve as distinct species or subspecies of vittatus, the three make a rather small, compact and distinct subgenus which is characterized by shells with a high spire, a verge with a rather large accessory pad, and a radula whose marginals are very thick, roundish in cross- section and with very reduced or absent denticles. There are only 24 to 28 rows of teeth. The oper- culum is serrated along one edge. The subgenus is represented in the Recent Indo-Paeific only. There are three fossil species or subspecies in the Upper Miocene and Pliocene of Indonesia which are treated in this account. Synonymy- 1931 Doxander Iredale, Records Australian Mus., vol. 18, no. 4, p. 212. Type by original designation: vittatus Gmelin [= Strombus vittatus Linne], Nomenclature— Gabb in 1868 implied, but did not legally select, campbelli (a close relative to vitta- tus) as the type of the genus Gallinula H. and A. Adams. However, the latter name is a homonym in any event and would not precede Doxander ( see under synonymy of Labiostrombus Oostingh). Strombus vittatus Linne, 1758 Strombus vittatus is a polytypic and very vari- able species extending in range from Japan south- ward through the East Indies and Melanesia to northern Australia. There are three geographically distinct subspecies, probably of late Pliocene or early Pleistocene origin. One of these is the north- ern japonicus Reeve from Japan. The typical sub- species, vittatus, occupies the central area from southeastern Asia to Melanesia. The third subspe- cies, campbelli, is evidently confined to the Austral- ian mainland. Workers solely using the morpho- logical concept of species consider these as sepa- rate species. A synopsis of the key characters of the three sub- species is given below: vittatus— body whorl smoothish, its dorsum with one weak knob; 1 to 4 smooth spiral cords or in- cised lines just below the suture (rarely a weak channel ) . japonicus— body whorl strongly and spirally corded; subsutural threads 1 to 3 and weak. campbelli— body whorl smoothish, its dorsum al- ways with one fairly strong knob or swelling sub- sutural band well-beaded, especially in earlier whorls. Strombus vittatus subspecies vittatus Linne, 1758 (PI. 17, figs. 14) Range— Southern China to eastern Malaya to Melanesia and northern Australia. Plate 87. Geographical distribution of S trombus listen T. Linne, japonicus Reeve and campbelli Griffith and Pidgeon. Gray, and the subspecies of Strombus vittatus— vittatus [09 - 963] 1 12 Doxander R. T. Abbott S trombklae Plate 88. Radula of Strombus vittatus vittatus Linne, Hong Kong. Fig. 1, central. 2, lateral. 3, inner marginal, showing cross-sections. 4, outer marginal. Remarks— This subspecies is very variable in size, coloration, height of spiral and degree of axial pli- cation. It is moderately common offshore, but in- frequently cast up on beaches. Specimens with ex- tremely high spires and strong axial plications per- sisting to the parietal wall are referred to as form turritus Lamarck or australis Schroter. Intergrades with the shorter, smoother forms are not uncom- mon. The small hump or smooth knob on the dorsum of the body whorl (so common in camp- belli) appears in some specimens. Specimens from Hong Kong have barnacles attached to the “dorsal” surface. A peculiar specimen from China lacking axial sculpture and having a channeled suture was illu- strated by Chemnitz in the Conchyl.-Cab., vol. 11, pi. 195A, figs. 1870 and 1871. It received a valid name in 1802 ( sulcatus Holten) and in 1817 ( sul - catus Dillwyn). I have not seen specimens, and am inclined to believe, as have others, that it is a mal- formed vittatus. A similar malformation has been found in S. gigas of the West Indies (named cana- liculatus Burry, 1949). Description— (of the typical form). Shell 40 to 66 mm. (1 M to 3M inches) in length, rather thin but strong, with a moderately- to well-produced spire, a winged outer lip, white aperture and with its exterior colored a light yellow-brown to tan which may have 4 to 5 narrow, white and brown flecked spiral bands on the body whorl. Whorls 11 to 12. Nuclear whorls 4, translucent-white or laven- der, glossy, and with the first one very small. First postnuclear whorl glistening, first with 1 to 3 spiral incised scratches, then with glossy, crowded, rather neat, axial riblets ( 15 to 22 per whorl ) . In later whorls the spiral incised lines are usually limited to 3 to 6 on the lower half of the whorl. The shouldered axial riblets do not invade the area just below the suture where there may be a smooth concave area or a series of 1 to 4 spiral threads. Base of shell with 15 to 20 low, flat-topped, spiral cords. Center of body whorl usually smooth, but may have a single low knob on the dorsum just below the suture. The axial riblet may disappear on the last two whorls. Columella slightly arching, enamel-white, its left side sometimes bordered by a longitudinal chink, smooth in the center, but above with a dozen very weak spiral rugae and at the base sometimes with 2 or 3 very weak lirae. Posterior canal narrow and extending to the suture above. Wing of outer lip tongue-like and curled slightly inward. Interior of body whorl glossy white and with a series of tiny, irregular, low, white, spiral lirae which are dispersed along a band a slight way back from the edge of the outer lip. Stromboid notch usually weakly developed. Ante- rior siphonal canal short, not recurved nor twisted. Spire angle varying from 30 to 40 degrees, and with or without 4 or 5 small, swollen, whitish former vari- ces in the upper 6 whorls. Periostracum extremely tlii u and transparent, usually being worn off, but sometimes persisting in the form of axial fimbria- tions in the concave subsutural channel. Oper- culum stromboid, light-brown, slightly arching, moderately thin and in our worn specimens show- ing signs of about 7 weak serrations. Radula with moderately strong teeth, 9 mm. in length and with only 25 to 26 rows. The marginals are thick and with few denticles and the peg on the lateral is quite reduced. Formula: 3-1-3 (also 2-1-2); 1-2; 2 to 4; 5. Verge simple, with a well- developed laminated pad which is mottled. Poste- rior mantle filament short. Measurements (mm.)— length width no. whorls 86.5 35.0 10 + 72.3 31.2 10 + 40.0 20.8 8 + 36.1 18.1 11 Synonymy— 1758 Strombus vittatus Linne, Systema Naturae, ed. 10, p. 745, no. 439 (In O. Asiae); 1767, ed. 12, p. 1211, no. 508; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., voi. Ill, art. 3, pp. 276-278. 1798 Lamlns vittatus Gmelin, Roding, Museum Boltenianum, Hamburg, pt. 2, p. 66, no. 838. Refers to Conchyl.- Cab., vol. 10, figs. 1481, 1482. ?1802 Strombus sulcatus Holten, Enumeratio Systematica Conchyl., beat. Chemnitzii, p. 56, no. 735. Refers to Conchyl.-Cab., vol. 11, figs. 1870, 1871; 1823, Dillwyn, An Index to Hist. Conchyl. Lister, ed. 3, p. 38. Not G. Fischer, 1807; not Anton, 1839 which is S. alatus Gmelin. (large; Hong Kong) ( average; Mindanao Id. ) ( small; Luzon Id. ) ( dwarf; Palawan Id. ) [09 - 964] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 113 1805 Strombus australis Sclnoter, Wiedermann’s Archiv. fur Zool. und Zootomie, Braunschweig, vol. 4, p. 93 ( Sudlandcrn ). Refers to Conchy]. -Cab., vol. 10, figs. 1481, 1482. Non Gray, 1826, non Anton, 1839 ( The turreted form. ) 1822 Strombus turritus Lamarck, Anim. sans Vert., Paris, vol. 7, p. 212 (no locality). Refers to Conchyl.- Cab., vol. 10, figs. 1481, 1482. Non Link, 1807. ( The turreted form. ) 1950 Strombus (Doxander) vittatus Gmelin, Beets, Overdruk Leidse Geol. Mededeelingen, vol. 15, p. 245. (Tur- reted form from Quaternary of East Borneo. ) Types— Linne’s type of vittatus was present in the Linnaean collection at the Linnaean Society of London (according to Hanley, 1855, p. 273). We hereby restrict the type locality to Amboina, Indo- nesia. Nomenclature— The higher spired and more strongly plicate form was first named australis Sehroter in 1805, and has precedence over the form name turritus Lamarck, 1822. The latter is a secon- dary homonym of Lambis turrita boding, 1798 which is a synonym of g alius Linne from the Carib- bean. Records ( see map, pi. 87 ) ( for typical vittatus and its “turreted” forms)— HONG KONG: south of Aap Li Chaan (A. Staple, ANSP); beach at Stanley (Mme. cle Breuil, ANSP ). THAILAND: all Gulf of Siam: Maikhas Beach, Phuket (G. M. Moore, MCZ); Khan Nu Paknam; Bangbert Bay; Koh Samet; Singora (all USNM). PHILIPPINES: 7-10 fms. at San Nicolas Shoals, Manila Bay, Luzon Id. (clu Pont-Academy Exped., 1958, ANSP); Cuyo Id., Pala- wan Prov. (ANSP); 40 fms., Aborlan, Palawan (ANSP. MCZ); Iloilo, Panay Id. (LISNM); Cadiz and Santa Cruz, Negros Id. (USNM); north end of Cebu Id. (Dr. Lucerno, ANSP); Catbalogan, Samar Id. (R. T. Abbott, MCZ); Zamboanga, Mindanao Id. (MCZ); oft Tawi Tawi, Sulu Archipelago ( 18 fms., green mud. Albatross Sta. 5164 USNM). INDONESIA: .39 fms., oft Taganak Id., Borneo (USNM). Sarawak, Brit. Borneo (MCZ). Keledjitan and Tjiperwagaran, Bantam, Java (both USNM); Amboina, Celebes (MCZ); 1400 meters in the Madura Straits; 27-32 meters in the Macassar Straits; 13 meters, Pulu Jedan, Am Ids. (all “Siboga”, Schepman, 1909, p. 148). DUTCH NEW C.LTNEA: 10-16 fms., off Cape Dgarwawoffi, Japen Id. ( NSF, 1956). AUSTRALIA: Queensland: Brampton Reef, Bowen (ANSP); Cape Upstart (ANSP); sand flats at Port Douglas (Tony Marsh, ANSP). FIJI: 12 fms., off Levuka Id. (E. A. Smith, “Challenger”, 1886, p. 420). Fossil records— Reported, but unverified by me, from the Pliocene of Java (see van der Vlerk, 1931, P- 247). Prob- ably based upon other species. S. deningeri P. J. Fischer, 1921, is the young of a vittatus vittatus -like Pliocene Strom- bus from Ceram Island, Indonesia. Beets ( 1950, vol. 15, p. 245 ) reports two specimens of the “turreted” form from the Quaternary of Blitong, Java, and one Quaternary specimen from Poeloe Boenjoe, East Borneo, Indonesia. A Pliocene Java, Indonesia, form (S. triangulatus Martin) is extremely close to Recent specimens of vittatus and might well be con- sidered a subspecies of it. Strombus vittatus subspecies japonicus Reeve, 1851 (PI. 17, figs. 18) Range— Southern half of Honshu Island to south- ern Kyushu Island, Japan. Remarks and description— This common Japanese subspecies differs from the southern vittatus in be- ing more brightly colored, in having a lower spire (45 to 50 degree angle), and in having small but well-developed, crowded spiral cords over the en- tire body whorl. There are two or three very weak, sometimes obsolete, spiral threads just below the suture. The outer lip is generally thicker and the spiral lirae within the aperture are fewer ( 25 to 30), longer and stronger. The posterior siphonal canal is usually longer, running beyond the above suture, and sometimes being sinuate. The banded pattern on the body whorl is pronounced in most specimens, and the brown coloring is usually dark. We have seen an all-orange specimen. In some specimens the shoulder of the body whorl may bear 5 to 8 indistinct, white-blotched rather equal- sized nodules. Periostracum moderately thick, yel- low-brown, rough and flaking off when dry. M easurements (m m .)— length width no. whorls 66.0 32.5 10 (large; Nagasaki) 56.8 27.1 11 ( average; Hazu-gun 47.6 24.0 9 + (small; Hiroshima) Habitat— Common in colonies on mud or gravel and mud bottoms from 5 to 20 fathoms. Synonymy— 1851 Strombus japonicus Reeve, Conchologica Iconica, Lon- don, vol. 6, Strombus, pi. 17, fig. and sp. 42 (Japan). 1869 Strombus japonicus Reeve, Lischke, Japanische Meeres- Conchylien, suppl. 4, vol. 1, p.' 30, pi. 5, fig. 7 (Nagasaki). 1959 Labiostrombus japonicus Reeve, Kira, Colour. Illus. Shells of Japan, Osaka, 2nd ed., p. 13, pi. 15, fig. 13. Types— Reeve's type is presumably in the British Museum of Natural History in London. The type locality is “Japan which we further restrict to Nagasaki, Kyushu Island. Records (see map, pi. 87) (all Japan)— HONSHU: Nonai, Mutsu Bay, Aomori Pref. (Nomura and Hatai, 1931, p. 11); Oga Peninsula, Akita Pref. ( Nishimura and Watabe, 1943, no. 3, p. 67 ) . Yamagata Pref. ( Nomura and Zimbo, 1936, no. 30, p. 28, no. 102). Tateyama, near Tokyo Bay, Chiba Pref. (ANSP); Suruga Bay, Shizuoka Pref. ( K. Oyama, 1943, p. 16, no. 146). Shirako, Mie Pref. (T. Habe, ANSP). Ei, Awaji Ich, Hyogo Pref. ( T. Habe, ANSP). Hiroshima, Hiroshima Pref. (A. R. Cahn, ANSP). Isshiki, Hazu-gun, Aiehi Pref. ( T. Habe, ANSP); Wakasa Bay, Kyoto Pref. (ANSP). SHIKOKU: Tosa Bay (ANSP). KYUSHU: Chi- kuzen (K. Hatai, ANSP); Tomioka, Amakusa, Kumamoto Pref. (T. Habe, ANSP); Nagasaki, Nagasaki Pref. ( H. Loomis, ANSP: USNM). Iwakawa, pt. 2, 1905, p. 86 re- cords this species from Ogasawarajima [Bonin Ids.] but I have not been able to verify this. Fossil records— RYUKYU ISLANDS: Simaziri Beds, lower Pliocene, Gabusoga, Okinawa Id. ( Nomura and Zimbo, 1936, vol. 18, no. 3, p. 259, pi. 11, figs. 27a, b. ) JAPAN: Honshu Island: Otake, Shimosa in the Upper Musashino of the Upper Pliocene or later (Yokoyama, 1922, vol. 44, p. 70, pi. 3, fig. 12); the Oti Graben Pleistocene in Noto Pen- insula ( Y. Otuka, 1935, vol. 13, p. 366, pi. 54, fig. 99). [09 - 965] 114 Doxander R. T. Abbott Strombidae Strombus vittatus subspecies madiunensis Martin, 1899 Rouge— Pliocene of Java. Remarks— This species is closest in most char- acters to vittatus joponicus, differing only in having a slightly lower spire and in having slightly more angular whorls in the spire. Altena’s Atjeh, Sumatra specimen ( madiunensis Martin, Altena, 1942, p. 52, figure 14) is probably not this subspecies, but pos- sibly another species which, at least in shape, is more like S. marginatus succinctus Linne from the Bay of Bengal. Nomura and Zimbo’s joponicus (1936, p. 259, pi. 11, fig. 27) from the Lower Plio- cene of Okinawa may be this subspecies. Synonymy— 1899 S trombus (s. str.) madiunensis K. Martin, Samml. Geol. Reichs-Mus. Leiden, n. Folge, vol. 1, pt. 1, p. 183, pi. 29, figs. 422, 422a, 422b ( Sonde, Gendingan, Madiun Residency, Java, Pliocene). Plate 89. S trombus vittatus deningeri P. J. Fischer. Pliocene of Ceram Island, Indonesia. Types, (from Fischer, 1927, pi. 212, figs. 25a, b). x 2. Strombus vittatus subspecies deningeri P. J. Fischer, 1921 Range— Pliocene of Ceram, Indonesia. Remarks— This species was based upon an imma- ture specimen which closely resembles the young stage of the Recent vittatus. It is not allied to vari- abilis or denti Cox. Synonymy— 1921 S trombus deningeri P. J. Fischer, Centralblatt fiir Mineral., Geol. Palaont., vol. 22, p. 244 (Western Seran); 1927, P. J. Fischer, Palaontologie von Timor, Stuttgard, pt. 15, art. 25, p. 56, pi. 212, fig. 25 a, b. (Pliocene, Seran [Ceram Id., Indonesia]). Strombus vittatus subspecies campbelli Griffith and Pidgeon, 1834 (PI. 17, figs. 13) Range— The north half of Australia from West- ern Australia to New South Wales. Remarks and Description— This is a very distinc- tive geographical subspecies of vittatus and is, by some, considered a separate species. The whorls in the spire are usually flatter and the subsutural cord bears numerous, small, but well-developed, axially elongated beads. The latter may disappear in the last whorl. Nuclear whorls 3M, glossy, translucent white, rarely tinged with lavender. First postnu- clear whorl with a fine incised line just below the suture, then joined by numerous axial, glossy rib- lets. Spire with 1 or 2 to 14 small, swollen, rounded, white, axial former varices. Color of shell whitish cream with irregular mottlings of light- to dark- brown which under the microscope show numer- ous delicate zigzag streaks, flecks and arrowheads. Columella enamel-white, glossy, with a few weak rugae at the top inner end and rarely with 1 or 2 weak lirae near the base. Spiral lirae within the inside of the outer lip usually very weak. Parietal wall slightly flattened due to an axial swelling to its left. This is followed by 2 or 3 small, rounded axial swellings on the shoulder, and then on the dorsum by a rather pronounced rounded knob. Base of shell with about a dozen spiral incised lines. Periostracum moderately thin, translucent yellow-brown, somewhat rough, and flaking off when dry. Operculum stromboid, about one fourth the length of the shell, and with 7 serrations. Radula 5 mm. and with 27 rows of teeth. Formula: 3-1-3; 1-2 (plus peg) (also 1-3 (plus peg)); 1; 3 to 5. Animal and verge similar to that of vittatus vittatus. Measurements (mm.)— length width no. whorls 68.4 30.4 9 + ( Sandy Cape, Qld. ) 50.3 24.0 8 + ( Bowen, Queensland ) 32.2 15.5 10 (Western Australia) Synonymy- 1834 Strombus campbellii “Gray” Griffith and Pidgeon, The Animal Kingdom— Cuvier, London, vol. 12, p. 600, pi. 25, fig. 6 (no locality); 1842, Sowerby, Thesaurus Conchyl., vol. 1, pi. 6, figs. 22, 23; 1908, Hedley, Proc. Linn. Soc. New South Wales, vol. 33, pt. 3, p. 460. 1886 Alaba (Styliferina) sulcata Watson, Voyage of the Chal- lenger, Zoology, vol. 15, p. 570, pi. 42, fig. 7 (Cape York, Queensland); 1905, Hedley, Proc. Linn. Soc. New South Wales, pt. 4, p. 523. [09 - 966] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 115 Plate 90. Strombus vittatus campbelli Griffith and Pidgeon, Broome, Western Australia. Figs. 1-4, radulae. 5, verge. Types— The type of Griffith and Pidgeon appears to be lost. No locality was given. The drawing of the type (PI. 24, fig. 6, supra cit.) is a low-spired, well-pigmented specimen such as we have seen from Bowen, Queensland, Australia. Nomenclature— Gray is usually not considered to be the author of Griffith and Pidgeon’s “The Ani- mal Kingdom”, and since no adequate descriptions accompany the explanation to the plates the latter two gentlemen are considered the authors of camp- belli. Records (see map, pi. 87) (all Australia)— WESTERN AUSTRALIA: Eighty Mile Beach; Cape Bossut; La Grange Bay (all H. L. Clark, MCZ); Ridall’s Beach and Black Ledge and sand flat, 2% miles south of Broome ( all V. Orr, ANSP); Cape Leveque (FI. L. Clark, MCZ); Augustus Id. (B. Bardwell, ANSP). NORTHERN TERRITORY: East Point, near Darwin (H. L. Clark, ANSP); Van Dieman’s Gulf (Calvert’s coll’n., ANSP); Yirrkala, Arnhem Land (USNM); Gulf of Carpenteria (ex Roth, Hedley, 1908, p. 460). QUEENSLAND: oil Cape York, 3-12 fms., coral mud ( R. B. Watson, “Challenger”, 1886, p. 418); between Ham- mond and Wednesday Ids. (Melvill and Standen, 1899, p. 165); Bedford Beach, Cooktown (MCZ); Cairns (MCZ); Dunk Id. (H. A. Pilsbry, ANSP); Magnetic Id., off Towns- ville (MCZ); Queens Beach, Bowen (P. Coleman, ANSP); Keppel Id. (Tony Marsh, ANSP); Bustard Head (ANSP); Moreton Bay (B. R. Bales, ANSP). NEW SOUTH WALES: Port Stephens (fide Angas, 1877, p. 185). [We have speci- mens from the Solomon Islands, but feel that the locality data is erroneous.] Fossil records— None reported. Strombus triangulatus Martin, 1879 Range— Upper Miocene of Java, Indonesia. Remarks— This species is allied to the Recent S. vittatus Linne. I. van der Vlerk ( 1931, vol. 5, p. 291 ) considered this species to be one of the guide fossils for the Miocene of the East Indies, although I believe some Pliocene species could be confused Plate 91. Strombus triangulatus Martin. Upper Miocene of Java, Indonesia. Type, (from K. Martin, 1879, pi. 9, figs. 5, 5a). Natural size. with it. Martin’s 1899 figs. 431, 431a are immature and may be the young of varinginensis Martin. Synonymy— 1879 Strombus triangulatus K. Martin, Die Tertiarsehichten auf Java, Leiden, vol. 5, p. 49, pi. 9, fig. 5 (Java): 1899, ibid., n. Folge, vol. 1, pt. 1, p. 186, pi. 30, figs. 431, 431a (Java). Strombus listeri T. Gray, 1852 (Pi. 92) Range— Known only from the Bay of Bengal from Ceylon to Burma. Remarks— I am provisionally treating listeri as a good species, although it is possible that it is merely a large, gerontic form of vittatus. The pe- culiarly misshapen siphonal canal suggests the lat- ter. To date, we know of only three adult speci- mens—the holotype in the Hunterian Museum in Glasgow which probably is the shell figured by Lister in 1685 on plate 855, fig. 12a. Its length is 4/i inches. Sowerby’s holotype is a fresh specimen and is 57s inches in length. Langdon ( 1874, Quart. Jour. Conch., London, vol. 1, p. 74) reports another specimen from Ceylon. The color of the shell is similar to that of vittatus— light-brown and whitish overlaid with dark-chestnut axial stripes. There is no spiral incised line below the suture as in most specimens of vittatus. E. A. Smith ( 1904, Ann. Mag. Nat. Hist., ser. 7, vol. 13, p. 469) states that the Steamer “Investigator” obtained two young speci- mens in the Gulf of Martaban, Burma, in 67 fathoms. If a good species, it is one of the rarest and most desirable of the world’s Strombus. Synonymy- 1852 Strombus listeri T. Gray, Ann. Mag. Nat. Hist., series 2, vol. 10, p. 429 (no locality). 1870 Strombus mirabilis Sowerby, Proc. Zool. Soc. London for 1870, p. 257, pi. 21, fig. 4 (Ceylon). Fossil records— None reported. [09 - 967] 116 Doxander R. T. Abbott Strombidae Plate 92. Strombus listen T. Gray. Ceylon, (holotype of S trombus mirabilis Sowerby, 1870, pi. 21, fig. 4). Natural size. [These occasioned blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sec/nence.] [09 - 968] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 117 Subgenus Lentigo Jousseaume, 1886 Type: Strombus lentiginosus Linne, 1758 This appears to have been a fairly wide-spread and old group which first occurred in the Lower Miocene of Indonesia ( S trombus preoccupotus Fin- lay) and the Miocene of Florida ( Strombus ald- richi Dali ) . Today, the living remnants are widely separated geographically and conchologically. In the Indo-Pacific are lentiginosus Linne and pipits Roding. In the tropical Eastern Pacific is granu- latus Swainson, 1822. A probable member of the subgenus, Strombus latus Gmelin, 1791 (formerly bubonius Lamarck), occurs alone in West Africa. Strombus fasciatus Born, a rather aberrant form, is limited to the Red Sea area. The shells are moderately large, somewhat quad- rate in shape, and have three or four spiral rows of knobs on the last whorl. The upper row bears large, somewhat equal-sized, blunt spines. The penis in lentiginosus and pipits has an auxiliary prong, and there is no basal peg on the marginal radular tooth. The operculum is weakly serrated. Synonymy— 1886 Lentigo Jousseaume, Le Naturaliste, Paris, 1st series, vol. 3, 8th year, no. 28, p. 220. Type by monotypy: L. lentiginosus Linne. [Lentigo “Klein Moreh, 1868, Malak. Blatt., vol. 15, p. 21 is a nude name.] Strombus lentiginosus Linne, 1758 (PI. 17, figs. 11, 12; pi. 94, fig. 4) Range— East Africa to the Marshall and Tuamotu Islands. Remarks— This is a rather widely distributed, shallow-water and coral-water species which is re- markably constant in its sculpture, coloration and size. It is characterized by the cream and light orange aperture which is frequently overlaid by a silvery or aluminum-like glaze, by the 4 or 5 squar- ish blunt knobs on the shoulder of the body whorl, and by the peculiar, angular lobe at the base of the columella. This species is most likely to be con- fused with S. pipus (Roding). The latter has a purple-brown aperture bearing fine spiral lirae, and its parietal wall has a peculiar “feathery” zig- zag pattern of soft browns. S. lentiginosus is common wherever it has estab- lished itself. In Polynesia it occurs in relatively few areas but is moderately common when found. Habitat— This species occurs in moderate to large numbers from low tide mark to a depth of about 12 feet, usually on a coral sand bottom which may be rocky and with weeds. They occur on barrier, fringing or lagoon reefs, and usually where the water is relatively clear. The outer shell surface may bear green algae, bryozoans, Vermetid and Hipponicid gastropods ( Sabia conica ( Schu- macher ) ) . Description— Shell 55 to 104 mm. (2 to 4 inches) in length, solid, heavy, nodulose, with a pinkish cream aperture and glossy parietal wall. Color of outer shell whitish with heavy specklings and mot- tlings of greenish gray to grayish brown. Parietal wall cream with gray and brown, irregular mottlings over which is laid a clear to translucent, shiny glaze. The glaze sometimes extends up on to the spire as far as the apex. Interior of aperture smooth, and flushed with light orangish rose. Outer lip tan with 5 to 6 indistinct light-brown bars. Columella smooth and white within, and swollen and pro- jecting near the lower end. Outer lip wavy, the up- Plate 93. Fig. 1. Living animal of Strombus pipus ( Reel- ing), female. Fig. 2, Strombus lentiginosus Linne, male, (both from Quoy and Gaimard, 1833, pi. 50). [09 - 973] 118 Lentigo R. T. Abbott Strombidae per end with 2 or 3 tongue-like projections. Strom- boid notch deep and somewhat W-shaped. Whorls 9 to 10. Nuclear whorls unknown, but appear to he 2 or 3. Postnuclear whorls with 12 to 14 spiral threads, increasing to small cords in later whorls. Suture minutely indented and wavy. Early whorls with numerous fine axial riblets becoming nodulated in later whorls at the shoulder. Last whorl with 7 to 9 large, squarish knobs at the shoulder. Below this are 3, 4 or 5 spiral rows ol small rounded knobs. In front of most of these knobs is a chestnut-brown spot. The upper 4 whorls in the spire bear 9 to 12 small, rounded, whitish former varices, sometimes lined up one below the other. Siphonal canal short, thick-walled and twisted to the right and slightly upward. Periostracum very thin, translucent-tan, and flaky when dry. Operculum stromboid, black- ish brown, strongly arched, with 2 or 3 worn, in- distinct or smoothish serrations, and about % the length of the shell. Radula ribbon strong, wine-red to amber, 7 to 8 mm. in length, and with about 45 to 47 rows of teeth. Lateral without a peg. Formula: 2-1-2; 1-3; 6; 7. One Palau specimen with marginal formula of 5; 5 and 6; 6. Verge 24 to 28 mm., with a strong thumb-like appendage on the distal blade. Poste- rior mantle filament proportionately short (3 to 4 mm.). Quoy and Gaimard ( 1833, Atlas of the Voyage de 1’ Astrolabe, pi. 50) depict the living animal as being mottled with green, with a yellow margined mantle edge, and the yellow eye being bordered with red ring. Measurements (mm.)— length width no. whorls 105.0 67.0 8 + ( large; “East Indies” ) 73.0 48.2 9 + ( average; Schouten Islands ) 54.5 37.0 8 + ( small; Palau Islands ) Synonymy— 1758 Strombus lentiginosus Linne, Systema Naturae, ed. 10, p. 743, no. 427 (In O. Asiae); 1767, ed. 12, p. 1208, no. 495; 1956, Dodge, Amer. Mus. Nat. Hist., vol. Ill, art. 3, p. 253-254; 1842, Sowerby, The- saurus Conchy!, vol. 1, Strombus, pi. 8, fig. 79; 1843, Kiener, Coq. Vivantes, vol. 4, pi. 18, fig. 1. 1798 Lambis rana Roding, Museum Boltenianum, Hamburg, pt. 2, p. 63, no. 802. Substitute name for lentigi- nosus L. 1834 S trombus rana Oken, Isis, Leipzig, vol. 27, pt. 4, pi. 6, fig. 3; 1836, loc. cit., vo! 29, pt. 1, p. 43. Refers to Quoy and Gaimard, 1833, Voy. Astrolabe, pi. 50, figs. 3-5. Types— According to Dodge (1956, p. 254) the type of lentiginosus is in the Linnaean collection in the Linnaean Society of London. Linne’s local- / Plate 94. Figs. 1-3, immature and adults of Strombus pipus (Roding), Mindanao Id., Philippines. 4, immature S. lenti- ginosus Linne, Philippines. All natural size. Plate 95. Strombus lentiginosus Linne, Palau Islands. Figs. 1-4, radulae. 5, verge. 6, operculum. 109 - 974] PHILIPPINE November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 119 ity was “O. Asiae” which we further restrict to Am- boina, Indonesia. Selected records (see accompanying map, pi. 96; solid dots: specimens examined; open circles: literature records) — TANGANYIKA: Inner Sinda Id. ( R. T. Abbott, USNM). KENYA: Diani Beach and Mombasa (R. T. Abbott, MCZ). ZANZIBAR: Common around the island (NSF). MADA- GASCAR: Nossi-be; Nossi Andrano; Tulear; Saradrana; Sainte Marie (all Dautzenberg, 1929, p. 467). MALDIVES: north Male Atoll (Y'ale Seychelles Exped., Peabody Mus. ). RYUKYU ISLANDS: Okinawa Id. (Mrs. A. A. Scott, ANSP). TAIWAN: Hoko ( Kuroda, 1941, p. 97). INDO- NESIA: Obi Major; Nusa Laut; Taam; Rotti; Kur Id. (all Schepman, 1909, p. 147). ELLICE ISLANDS: Funafuti (Hedley, 1899, p. 428). CAROLINES: Yap; Eawujlik; Ifa- lik (all USNM). MARSHALLS: Eniwetok; Bikini; Ron- gelap; Rongerik (all USNM). SOCIETY ISLANDS: Outer reef, Moorea Id. (H. A. Rehder, USNM). Ative, Punaavia. Tahiti Id. (R. Robertson, ANSP). TUAMOTUS: (Dautzen- berg and Bouge, 1933, p. 299). Fossil records— None reported. Strombus pipus (Roding, 1798) (PI. 17, figs. 9, 10; pi. 94, figs. 1-3) Range— East Africa to the Society Islands. Remarks—' This attractive, purple-mouthed Strom bus is rather uncommon, although it has a range as wide as that of its abundant relative, S. lenti- ginosus. In some areas, it is considered a rarity al- though this may be due to the fact that it lives well off shore. Our records show two areas where it is not uncommon— the western portion of the Indian Ocean and the Philippines. It has not been re- corded, as yet, from Micronesia or Hawaii. S. pipits is characterized by the purple-brown aperture which has a series of fine spiral lirae along the inside of the outer lip, and by the “feathery" mosaic of soft-brown maculations on the glossy parietal wall. The shell is smaller than most lenti- ginosus, and the shoulder knobs on the body whorl vary from 8 or 9 large to 15 very small ones. Inter- grades exist in the same colony, and there is no correlation between darkness of aperture and de- gree of sculpturing. The parietal glaze does not extend up on to the apex, as in lentiginosus. The young of pipits have a large brown spot on the upper third of the columella, a feature absent in lentiginosus. Habitat— Lives on bottom of coral sand and algae from 8 to 39 fathoms. Description— Shell 38 to 70 mm. ( lM to 2/i inches ) in length, solid, roughly sculptured, roundly quad- Plate 96. Geographical distribution of Strombus lentiginosus Linne. [09 - 975] 120 Lentigo R. T. Abbott Strombiclae Plate 97. Radula of Strombus pipus (Roding). 1, central. 2, lateral. 3, inner marginal. 4, outer marginal. rate, and with a rugose, purplish-black aperture. Color of outer shell white with light or heavy, dif- fused light-brown mottlings and numerous, very fine axial lines of brown which give a “feathery " mosaic appearance. Whorls 10 to 11. Nuclear whorls 2M, whitish, smooth, and glossy. First 2 or 3 postnuclear whorls with numerous, small, but strong, smoothish axial riblets ( about 24 per whorl ) between which are about 10 spiral threads. The latter cross the surface of the riblets after the third whorl. Penultimate whorl with 8 to 14 small knobs. Shoulder of body whorl with 8 or 9 rather large squarish, axially pinched knobs or with 9 to 15 small rounded knobs. Below these are 3 or 4 spiral rows of much smaller, rounded knobs or swollen beads. Apex with 8 to 12 whitish, swollen former varices, the last 2 or 3 sometimes being quite broad and low. Parietal wall glazed with the shell color showing through. Columella thick and with a cream, smooth glaze over the lower two thirds. Upper portion tainted with brown or purplish. Deep interior of aperture whitish lavender becom- ing dark purple-brown or purple-black and spirally rugose towards the thickened outer lip. Top of outer lip extends slightly upward and with a broad sulcus which has 2 small white, rounded projec- tions. Central portion of lip edge cream with 5 to 6 pairs of brown bands. Stromboid notch U-shaped and sometimes with 3 to 5 small white teeth at the edge. Base of outer lip projecting and with 3 to 5 small white teeth. Interior of siphonal canal purpl- ish brown. Periostracum thin, microscopically stri- ate, translucent-tan. Operculum stromboid, light- brown, rather narrow, arching with about 7 very poorly developed (or badly worn) serrations, and with a strong narrow rib on the attachment side. Radula ribbon 7 mm. in length, its denticles long. Formula: 2-1-2; 1-5 or 1-6; 9 or 10; 10 or 11. Poster- ior mantle filament very short. Quoy and Gaimard (1833, Atlas of the Voyage de I Astrolabe, pi. 50, fig. 1,2) depict the living ani- mal as being delicately mottled and flecked with light-brown, and with the yellow mantle edge hav- ing large spots of brown. The eye rings of white, black, bluish and red. See our plate 11, p. 33. Measurements (mm.)— length width no. whorls 70.0 42.5 9 + ( large; Mauritius ) 38.0 28.0 8 + (small; Philippines) 57.5 36.5 10 (average; Zamboanga) Synonymy— 1798 Lambis pipa Roding, Museum Boltenianum, Hamburg, pt. 2, p. 63 (no locality). Refers to Conchyl.-Cab., vol. 3, figs. 825, 826. 1817 Strombus papilio “Chemnitz” Dillwyn, Descr. Cat. Recent Shells, London, vol. 2, p. 661 (East Indian Seas). Refers to Conchyl.-Cab., vol. 10, figs. 1510, 1511; 1818, Wood, Index Test., London, p. 116 East Indies); 1843, Kiener, Coq. Vivantes, vol. 4, pi. 17, figs. 1, 2. Plate 98. Geographical distribution of Strombus pipus (Roding), showing the discontinuity of its range. [09 - 976 J November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 121 1822 Strombus exustus Swainson, Zoological Illustrations, London, series 1, vol. 3, pi. 134 ( Haynam Id., East Indies); Berge, Conchylienbuch, Stuttgart, p. 231, pi. 38, fig. 4. 1851 Strombus adustus “Swainson” Reeve, Conch. Icon., vol. 6, Strombus, pi. 13, sp. 29. Non Gray, 1826. Type— Roding’s pipus is based upon figures 825 and 826 of volume 3 of the Conchylien-Cabinet which, in turn, are based upon a specimen in Bol- ton's collection, according to Martini. That speci- men is the type, but is probably lost. We restrict the type locality to Jolo Id., southern Philippines. Swainson’s type of exustus may be at Cambridge University, England. N omenclature— The more familiar name of papi- lio Dillwyn, 1817, has already been replaced by many recent authors with Roding’s earlier pipus. S. exustus Swainson is a synonym based upon a color form, and adustus Reeve was a misquote of Swain- son’s exustus. Records (see map, pi. 98)— ZANZIBAR: 13a mi. W.S.W. of Ras Mungwe, 8 fms. (NSF, 1957). INDIAN OCEAN ISLANDS: Cargador Carajor, 30 fms.; Amirantes, 30-39 fms.; Praslin Id., Seychelles, 34 fms. (all Melvill, “Sealark”, 1909, p. 92). Mauritius (ANSP). RYUKYU ISLANDS: Okinawa Id. (A. A. Scott and A. R. Calm, ANSP). TAI- WAN: Taihokusyu (Kuroda, 1941, P- 97). PHILIPPINES: Gubat, Sorsogon Prov., Luzon Id. ( duPont-Academy Exped., 1958); Cebu City and Olango Id., Cebu Id. (A. B. Franco, ANSP); Jolo Id., and Siasi Id., Sulu Archipelago (MCZ); oil Simonar Id., 28 fms.; of! Lapac Id., Tapul Id., 10 fms., Albatross Sta. 5149 (both USNM). INDONESIA: Banda, Timor and Rotti Ids.. 9-45 meters ( Schepman, 1909, p. 147). AUSTRALIA: Green Id., Queensland (Tony Marsh, in litt. ) . SOLOMONS: Florida Id. (USNM). SOCIETY ISLANDS: Punaruu, Punaauia, Tahiti ( R. Robertson, 1952, ANSP); Mataiea, Tahiti (J. Jaquenin, USNM). Fossil records— None reported. Strombus fasciatus Born, 1778 (PI. 14, figs. 16, 17) Range— Limited to the Red Sea. Remarks— This is one of the most distinctive and most restricted in its range of all the Indo-Pacific S trombus. It has no close living relatives, although it shows some features of decorus Roding. It is moderately common wherever it lives. The shell is readily recognized by its somewhat conic shape, by its large, somewhat evenly-sized shoulder spines, by the orange-yellow aperture and by the 5 to 7 broken, narrow, spiral bands of black-brown on the body whorl. Habitat— It is presumably moderately common in shallow water in sand or muddy areas. Description— Shell 32 to 50 ( 1)1 to 2 inches ) in length, solid, somewhat conic, spined at the angular shoulder, with a yellow-orange aperture, and the body whorl with 5 to 7 broken, narrow, black- brown bands. Whorls 9. Nuclear whorls 3(?), glossy translucent whitish. Postnuclear whorls pink- ish or whitish, slightly rounded, may have 3 to 5 small, whitish former varices, and may or may not have a single, spiral incised line below the suture. Knobs develop on the penultimate whorl, and be- come 8 to 12 smooth, pyramidal spines on the angu- lar shoulder of the last whorl. Central part of last whorl may have 1 or 2 low, rounded, broad spiral cords. Columella straight, weakly calloused, smoothish, except for microscopic spiral striae near the edge in some specimens. Columella whitish to translucent-yellow. Inner aperture smooth, glossy, flushed with yellow-orange. Siphonal canal very short. Stromboid notch U-shaped. Outer shell cream with fine, light-brown flecks over which are 5 to 8 broken, black-brown, narrow, spiral color bands. Periostracum thin, smoothish and translu- cent-tan. Operculum, radula soft parts not observed. Measurements ( mm.)— length widtli no. whorls 50.0 26.0 9 (large; Red Sea) 42.1 24.0 9 ( average; Red Se: 32.1 19.1 7 + (small; Red Sea) Synonymy— 1778 Strombus fasciatus Born, Index Rerum Natur. Mus. Caesarei Vindobonensis, p. 274 (no locality); refers to Chemnitz Conchyl.-Cab., vol. 3, figs. 800-802. Non Gmelin 1791, non Roding 1798. 1788 S trombus subalata Herbst, Natur. Abbild. der merkw. Wiirmer, Berlin, vol. 9, p. 204, no. 15, pi. 48, fig. 3 ( Red Sea ) . 1798 Lambis elegantissima Roding, Museum Boltenianum, Hamburg, pt. 2, p. 61, no. 773; 1807, Link, Beschr. Natur. -Samml. LTniv. Rostock, pt. 2, p. 108 (no locality ) . 1817 Strombus polyfasciatus Dillwyn, Descriptive Cat. Re- cent Shells, vol. 2, p. 662 (Red Sea). 1822 Strombus lineatus Lamarck, Anim. sans Vert., vol. 7, p. 211, no. 29 (Ocean indien?). Refers to Conchyl.- Cab., vol. 3, figs. 800-802; 1843, Kiener, Coquilles Vivantes, vol. 4, pi. 30, figs. 1, la. 1828 Strombus lineolatus Wood, Index Testaceol., Suppl., p. 13, pi. 24, fig. 11 (no locality). 1844 Strombus subulatus Herbst, Duclos, Illustr. Conchy!., vol. 2, p. 7. 1885 Strombus flavigula Meuschen, Tryon, Manual Conch., Phila., vol. 7, p. 120. Types— Born’s type is probably in the museum at Vienna. No locality was given, and we hereby des- ignate the Red Sea as the type locality. Records— RED SEA: Port Sudan, Anglo-Egyptian Sudan (ANSP); Jobal Straits, Egypt (ANSP); Massaua, Eritrea (ANSP); Aqaba, Gulf of Suez (A. Hadar); Jidda Harbor, Saudi Arabia (USNM); Sharm Ubhar, and Genaba Bay, Farasan Kebir, Saudi Arabia (both USNM); 10 km. north of Jidda, Saudi Arabia (C. Aslakson, ANSP). Fossil records— Port Sudan, raised coral reef. Pleistocene ( Hall and Standen, 1907, p. 67 ) . EGYPT : Pleistocene; beach, 50 ft. alt., Gemsah; beach, 80 ft. alt., Wadi Gueh, west of Kosseir (both R. B. Newton, 1900, p. 508). [09 - 977] 122 Lentigo R. T. Abbott S trombidae Strombus latus Gmelin, 1791 (PI. 99, figs. 1 and 2) Range— West coast of Africa from Spanish West Africa to Angola, including the Cape Verde Islands. Remarks— This is not an Indo-Pacific species, but is included here, since it is the only living Strombus in the Eastern Atlantic. It is recognized by its some- what quadrate shape, rather evenly-sized knobs on the shoulder, the peculiar, raised spiral cord or Plate 99. Figs. 1 and 2. Strombus latus Gmelin (formerly bubonius Lamarck), Tropical West Africa. Both natural size. ridge running back from the stromboid notch, and by its light-brown, white-flecked and rose-striped exterior. The periostracum is usually heavy and brownish. In general shape and coloration, it stands midway between lentiginosus of the Indo-Pacific and granulatus of the Panamic Pacific Province. Unfortunately, there have been four earlier names applied to this shell which, until Dodge’s 1956 re- vision, had been known as bubonius Lamarck. Duclos employed the name latus Gmelin in 1844 in his monograph. Description— Adult shell 90 to 156 mm. in length, solid, moderately heavy, somewhat quadrate in shape, well-spired, and maculated with orange- brown, rose and white. Whorls 10. Nuclear whorls not seen. Spire slightly convex, the early whorls somewhat distorted by 2 to 5 raher broad former varices. Spiral sculpture of 8 to 12 fine cords which disappear in the last two whorls. Suture well-in- dented, wavy because it half-covers the numerous, evenly-sized, rounded knobs in the whorls. 8 to 10 rounded knobs on the shoulder of each whorl. In the last whorl, the last three knobs are the largest. Below these, and midway on the whorl, is a second spiral row of much smaller, rounded knobs. Near the base of the shell, and in line with the stromboid notch is a single, raised, strong cord or ridge. Outer lip slightly expanded, its edge reflected, smooth, glossy and with 3 or 4 broad bands of light-brown. Columella and parietal wall smooth, glossy, cream- ish tan. Interior of aperture white. Stromboid notch deep. Base of columella constricted. Periostracum moderately thick, light-brown, and flakes off when dry. Operculum and soft parts unknown. Measurements ( mm.)— length width no. whorls 156.0 102.0 10 + ( large; West Africa) 106.0 68.0 10 + ( average; Cape Verde Ids. ) 88.0 51.0 10 + ( small; West Africa) Types— The whereabouts of the types is unknown. We hereby designate Sierra Leone, West Africa, as the type locality. Synonymy— 1791 Strombus latus Gmelin, Systema Naturae, ed. 13, p. 3520 (no locality). Refers to Seba, pi. 63, figs. 4, 5; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, p. 272, footnote 1. 1791 S trombus fasciatus Gmelin, Systema Naturae, ed. 13, p. 3510, no. 9 (in Africa); refers to Conehyl.-Cab., vol. 3, figs. 833-834. Non Born 1778. 1795 Strobus auratus Spalowsky, Prodromus Systema His- toricum Testaceorum, Vienna, pp. 43-44, pi. 6, fig. 9 (as S trombus) (India orientals? ) . [09 - 978] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 123 Plate 100. S trombus ( Lentigo ) preoccupatus Finlay. A Borneo, Indonesia, (from Beets, 1941, pi. 3, figs. 123-144). series of Lower Miocene specimens from Mangkalihat, East Natural size. 1798 Lambis cornea Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 63, no. 808 (no locality); refers to Conchyl.-Cab., vol. 3, figs. 833-834. 1798 Lambis carnaria Roding, loe. cit., p. 64, no. 809 (re- fers to Conchyl.-Cab., vol. 3, fig. 893, a young specimen ) . 1807 Lambis fasciatus Gmelin, Link, Beschr. Natur.-Samml. Univ. Rostock, pt. 2, p. 109. 1822 Strombus bubonius Lamarck, Anim. sans Vert., vol. 7, p. 203 (Antilles [erroneous]); 1950, Nickles, Man- uels Ouest-Africains, vol. 2, p. 76, fig. 106. 1822 Strombus dilatatus Lamarck, Anim. sans Vert., vol. 7, p. 203, no. 8 (no locality). Refers to Seba, vol. 3, pi. 63, figs. 4 and 5 [which might also be a smooth S. costatus Gmelin]; Kiener, 1843, Coquilles Vi- vantes, vol. 4, pi. 5. Non Swainson 1821. 1827 Strombus adansoni Defiance, in Blainville’s Diet. Sci. Nat., vol. 51, P- 115. Refers to Adanson, 1757, pi. 9, fig. 30 ( L’ile Goree). 1833 Strombus bubo Lamarck, Deshayes, in Exped. Scientif. de Moree, Mollusques, vol. 3, pt. 1, p. 192. Non Roding, 1798. 1844 Strombus lotus Gmelin, Duelos, in Chenu, Illust. Con- chy]., vol. 4, Strombus, pi. 13, figs. 5-7. Records— SPANISH WEST AFRICA: Rio de Oro (M. Nickles, 1950, p. 77. CAPE VERDE ISLANDS: (ANSP, MCZ). SIERRA LEONE: Turtle Island, off Sherbro Is- lands; off Freetown (both MCZ). SPANISH GUINEA: Adje, 20 miles south of Benito; Corisco Island (both MCZ). Principe Island (MCZ). San Thome Island (Hoyle, 1887, p. 340 and MCZ). ANGOLA: (M. Nickles, 1950, p. 77). Strombus preoccupatus Finlay, 1927 Range— Lower and Upper Miocene of Java and Borneo, Indonesia. Remarks— Beets (1941, p. 67, pi. 3) has given an excellent series of figures of this interesting species which has no Recent counterpart in the Indo- Pacifie. S. fasciatus (Red Sea) may belong to this stock. The Recent S. g ranulatus Swainson from the tropical Eastern Pacific is extremely similar to pre- occupatus. Strombus nodosus (Borson, 1820) from the Italian Tertiary belongs to the same group. Synonymy— 1881 Strombus spinosus K. Martin, Samml. geol. Reichs- Mus., Leiden, 1st ser., vol. 1, p. 122, pi. 7, figs. 3, 4 (Podjok; Djokdjokarta; and Wirosari, Java; non Linne, 1767, p. 1212; 1899, K. Martin, loc. cit., part 45, p. 176, pi. 28, figs. 408-409; 1921, loc. cit., n. Folge, vol. 1, pt. 2, p. 468, pi. 59, fig. 56 (Tji Talahab and Tji Angsana, Java). 1927 Strombus preoccupatus Finlay, Trans, and Proc. New Zealand Inst., vol. 57, p. 502 (new name for spi- nosus Martin, non Linne); 1941, Beets, Overdruk Verhand. Geolog. Mijnbowk. Genoot. Nederl. Kolo- nien, Geol. series, vol. 13, p. 67, pi. 3, figs. 123-144 ( East Borneo ) . [09 - 979] 124 Lentigo R. T. Abbott S trombidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [09 - 980] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 125 Subgenus Euprotomus Gill, 1870 Type: Strombus aurisdianae Linne, 1758 This group is limited to the Indo-Pacific, one species extending from Melanesia to East Africa, the other five mainly centered in the Western Pa- cific. The fossil record goes back only to the Pliocene of the Western Pacific. The shells are medium- sized for the genus and are characterized by a high spire, a high prong-like extension of the posterior end of the outer lip, and a smooth columella. The inside of the outer lip may be smooth or with strong, spiral lirae. The penis is “keeled", that is bearing an accessory projection of the distal blade. The tentacles are usually short and located quite near the end of the eye peduncle. The marginal radular teeth are very broad and bear 4 to 6 large, triangular denticles. There is no basal peg on the lateral tooth. The operculum bears 6 to 8 small serrations. Synonymy— 1847 Monodactylus “Klein” Herrmannsen, Indices Generum Malacozoorum Primordia, vol. 2, p. 53 [invalid]; 1852, “Klein" Mdrch, Cat. Conchyl. de Yoldi, p. 62; 1854, “Klein" H. and A. Adams, The Genera of Recent Mollusca, vol. 1, p. 259; 1868, H. and A. Adams, Gabb, American Journal of Conchology, vol. 4, pt. 3, p. 138. Type by subsequent designation: Strombus adustus = aurisdianae aratrum Roding, 1798; 1884, “Klein" P. Fischer, Manuel de Con- ehyliologie, Paris, pt. 7, p. 670. Type by subsequent designation: Strombus pacificus Swainson = vomer Roding, 1798; 1904, “Klein” Cossmann, Essai de Paleoconch. Comparee, Paris, 6th book, p. 7. Type by subsequent designation: Strombus gallus Linne [invalid]. Non Monodactylus Lacepede, 1800. 1870 Euprotomus Gill, American Journal of Conchology, vol. 5, pt. 3, p. 131, footnote. Type by monotypy: Strombus aurisdianae Linne. Strombus aurisdianae subspecies aurisdianae Linne, 1758 (PI. 14, figs. 3, 4; pi. 101, figs. 1-4) Range— Central East Africa to the Solomon Is- lands, and the Ryukyu Islands south to Queensland, Australia. Remarks— This is a well-known and moderately common shallow-water species. In some areas in the Philippines it is abundant. Some confusion has existed concerning its close relatives and various forms. Its nomenclatorial history in relation to the synonym, lamarcki Sowerby, 1842, is discussed be- low. We consider bulla (Roding) to be a separate species, but aratrum (Roding) from northern Aus- tralia to be a subspecies. Below is a synopsis of the characters of these three taxa: S. aurisdianae aurisdianae— exterior rough and with a dull finish; interior of aperture usually pink- ish and with 8 to 12 fine lirae on the upper part of the outer lip; spire glazed over on the ventral side of 3 or 4 whorls above; without brown splotches in the glazed portions; siphonal canal reflected about 90 degrees. S. aurisdianae aratrum— exterior rough and with a dull finish; interior of aperture brownish orange to salmon and with 5 to 9 fine lirae on the upper part of the outer lip; sjfire glazed over only on the ventral side of 1 or 2 whorls above; with brown splotches in the glazed portions; siphonal canal reflected about 75 degrees. S. bulla— exterior smoothish ( except for single row Plate 101. Figs. 1 and 2, Strombus aurisdianae Linne, Zanzibar. 3, immature S. bulla (Roding) from the Luzon Island. 4, “wingless” abnormality of aurisdianae from Cebu Island, Philippines. 5, S. bulla (Roding), brown-spotted form from Cebu Island, Philippines. All natural size. [09-991] 126 Euprotomus R. T. Abbott Strombidae of knobs) and with a glossy finish; interior of aper- ture reddish orange and with no or rarely 1 or 2 fine lirae on the upper part of the outer lip; spire usually glazed over the entire spire in adults; glaze white in color, sometimes with violet on the spire; siphonal canal reflected 85 to 95 degrees. Typical ciurisdianae is characterized by its rough exterior, smoothish brownish to reddish orange aperture, the single wing-like extension at the top of the outer lip, the strongly recurved siphonal canal, and by the 5 to 9 shoulder knobs on the dorsal side of the body whorls. In the Western Pacific it is fairly constant in characters, although Ryukyu Island specimens (form chrysostomus ) may take on the cream-orange apertural coloration of the Australian subspecies, aratrum. However, certain Philippine and Indian Ocean specimens may also exhibit this probable dietary form. Certain Zanzi- bar and Seychelles specimens show a tendency to- wards stronger rugosity and a uniform orangish ex- terior. They intergrade with normal Indian Ocean aurisdianae, and, at best, might be considered an “incipient subspecies.” We collected an adult malformed specimen in 1958 on Olango Island, Cebu, Philippines, in which the upper part of the outer lip is rounded over and entirely lacking the posterior projection. (PI. 101, fig. 4.) Habitat— Occurs in moderate numbers in shallow water from the low tide mark to a depth of about 12 feet, either on grassy sand flats, on dead coral reefs, or on a coral sand and grass bottom. Description— Shell 46 to 76 mm. (2 to 3 inches) in length, solid, rugose, with a posterior projection on the outer lip, with a strongly recurved siphonal canal and with a smoothish, glossy, cream and orange aperture. Nuclear whorls 3, glossy, trans- lucent, rounded, and either whitish, tan or brown in color. First 3 postnuclear whorls well-rounded, neatly reticulate. Following whorls with numerous axial, knobbed ribs crossed by a dozen uneven spiral threads. There is a large, wavy cord just be- low the suture. Shoulder of last whorl with 5 to 9 small, short knobs ( other knobs are glazed over by the parietal wall). Below are two spiral rows of very obscure nodules. Base of shell with numerous rather smooth spiral, crowded, rounded cords. Parietal glaze cream to whitish and extending up on to 3 to 5 of the whorls in the spire. Columella straight, glossy, tan or light-cream, and smooth ex- cept for indistinct lirae and wrinkles at the very top. Inner lip smooth, brownish orange to pinkish, rarely yellowish orange, bordered with a broad band of whitish to tan-cream; aperture smooth, ex- Plate 102. Strombus aurisdianae Linne. Figs. 1-5 from Dutch New Guinea. 1, central radular tooth. 2, lateral. 3, inner marginal. 4, outer marginal. 5, verge, showing cross-section,. 6 and 7, verge and operculum from Zanzibar specimen. cept for a dozen lirae below the deep stromboid notch and 8 to 12 fine, irregular, white lirae on the floor of the posterior canal region. Outer lip moder- ately sharp; rounded, glazed, smooth posteriorly where there are 5 to 7 grayish brown spiral bars. Posterior projection of outer lip extending half-way to all the way back as far as the apex of the spire. It may extend straight back or curve slightly to- wards the apex. Siphonal canal recurved about 90 degrees and slightly twisted to the right. Periostra- cum very thin, translucent yellowish, flaking off when dry, and usually worn away from the shell in life. Operculum stromboid, dark-brown, strongly arched, pointed, and with 6 to 7 serrations. Radula ribbon 6 to 8 mm. in length, with about 40 to 50 rows of teeth. Formula for New Guinea: 2- 1-2; 1-3 (without peg); 4; 5. Zanzibar: 2-1-2 (also 3- 1-3); 1-3 (without peg); 4 or 5; 5 or 6. Verge simple, 16 to 18 mm. in length, with a laminated pad. Posterior mantle filament 4 mm. in length. M easurements (mm .)— length width no. whorls 76.5 42.9 9 + ( large; Ryukyu Islands ) 61.0 34.0 9 ( average; Cebu Id. ) 46.5 25.7 10 ( small; Negros Id. ) [09 - 992] November 23, 1960 INDO-PACIFIC MOLLUSCA, vo). 1, no. 2 Strombus 127 Plate 103. Geographical distribution of Strombus auris- dianae Linne. Synonymy— 1758 Strombus auris-dianae Linne, Systema Naturae, ed. 10, p. 743, no. 429 (In O. Asiae); 1767, ed. 12, p. 1209, no. 497; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 256-258. 1798 Lambis stiva lidding, Museum Boltenianum, Ham- burg, pt. 2, p. 64, no. 815. Substitute name for aurisdianae L. 1798 Lambis buris Roding, ibid., p. 64, no. 819. 1842 Strombus lamarckii Gray, Sowerby, Thesaurus Con- ehyl., vol. 1, p. 35, pi. 9, figs. 98, 99. Non Gray, 1826; 1938, Adam and Leloup, Result. Sci. Voy. Indes Orient. Neerl., vol. 2, fasc. 19, p. 117. Non Gray, 1826. 1942 Strombus chnjsostomus Kuroda, Venus, vol. 12, p. 7, figs. 1 and 2 on p. 4 (Okinawa Ids.). 1880 Strombus striatogranosus “Moreh,” von Martens in Mobius, Beitr. Meeresfauna I. Mauritius Seychellen, Berlin, p. 277. Types— Linne’s types are in the Linnaean Society of London and are a mixture of two species. I hereby designate tire first figure reference in Linne’s 10th edition synonymy as being the “type figure” or the most representative of the species— Rum- phius, pi. 37, fig. R. The original type locality was “O. Asiae”. I further restrict it to Amboina, Indo- nesia. N otnenclature— The exact identity of Linne’s aurisdianae has been a subject of contention at various times, commencing with Hanley ( 1855, pp. 268-269 ) and continued by Dodge ( 1956 ) who gives a lengthy summary of the matter, but still confuses the problem. It is a fact that Linne had a mixture of specimens and his cited figures rep- resented two species. The choice is between the smooth species (which we call bulla Roding) and the rough species referred to by authors as lamarcki Sowerby. I prefer to reserve the name of aurisdi- anae for the very common species which has its “dorso muricato” and is figured in Sowerby ’s The- saurus, pi. 9, figs. 98 and 99. The name lamarcki was first instituted by Gray (not Sowerby, 1842) in 1826 and applies to the smooth species ( bulla Roding). On this basis, we disagree with Oostingh ( 1925, p. 54 ) that Sowerby 1842 was the first re- visor. Selected records (see accompanying map, pi. 103); solid dots: specimens examined; open circles: literature records) — MOZAMBIQUE: Port Amelia (MCZ). TANGANYIKA: Mboa Magi (R. T. Abbott, USNM). MADAGASCAR: Ma- junga; Tufear; Sarodrano (all Dautzenberg, 1929, p. 467). Nossi-be (A. Chavane, ANSP). Gloriosa and Providence Id. (USNM). SEYCHELLES: Cerf Id. (Yale Peabody Mus.). MALDIVES: Hulule Id. (E. A. Smith, 1903, p. 612, no. 202). CEYLON: Pearl Bank, Gulf of Manaar (G. and M. Kline, NSF, 1956). MALAYA: off Pulau Sudong, Singapore (R. D. Purchon, ANSP). SUMATRA: near Poelo Raja, and Tjalang, west Atjeh (Oostingh, 1929, no. 39, p. 2). RYUKYU ISLANDS: Shioya, Shanawan Bay, Okinawa Id. (USNM). PHILIPPINES: common on the following islands: Luzon; Samar; Cebu; Bohol; Mindoro; Negros; Catanduanes; Bala- bac; Mindanao; Sanga Sanga, Sulu Archipelago (all ANSP). Lubang; Marinduque, Ramblon; Panay; Camiguin ( all USNM). AUSTRALIA: Queensland: Hope Id. (USNM); Danley Ich, Torres Straits (MCZ); Low Isles (Tony Marsh, ANSP). SOLOMONS: Bougainville Id. (MCZ). Fossil records— Altena (1942, p. 58) reports one specimen ( as lamarckii Sowerby ) from the Pliocene Upper Kalibeng layers, Java, Indonesia, but I have not seen this specimen. Records for the Ryukyu Islands are fossil specimens of S. vomer hawaiensis Pilsbry. KENY A: Pliocene, Crag: South Mombasa Id., just west of Mbaraki Creek ( J. Weir, Mono- graph 5, 1938, p. 68, pi. 5, fig. 2). HAWAIIAN CHAIN: Pleistocene: Oahu Island: entire fossil from Honolulu Har- bor (J. M. Ostergaard, 1928, p. 27; this may be a vomer hawaiensis Pilsbry). Strombus aurisdianae subspecies aratrum (Roding, 1798) (PI. 14, figs. 1, 2) Range— Northeast Queensland, Australia. Remarks and Description— This subspecies occurs nearer to the mainland and in muddier waters than the typical subspecies which is a coral-water form. S. aurisdianae aratrum is somewhat more elongate, with a less recurved siphonal canal, usually with [09 - 993] 128 Euprotomus R. T. Abbott Strombidae smaller and more numerous knobs on the shoulder of the last half of the body whorl, and with a darker more orange-brown aperture. The nuclear whorls are whitish and very similar to those of aurisdianae. The upper portion of the parietal wall is poorly glazed and well decorated with numerous, dark blackish brown zigzag stripes. Black-brown staining is generally present around the glossy underside of the shell. An ecologic and life history study would clarify the relationship of these two subspecies. Measurements (mm.)— length width no. whorls 90.6 42.3 10 (large; Cairns) 73.8 41.2 9 + ( average; Pilot Point ) 58.8 30.6 10 (small; Bowen) Habitat— Moderately common on sand in inter- tidal areas. Synonymy— 1798 Lambis aratrum Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 64, no. 820. Refers to Conchy!. -Cab., vol. 10, figs. 1487-88. 1816 Strombus aurisdianae Linne, Lamarck, Le Liste, p. 4. Encycl. Meth., pi. 409, figs. 3a, b. 1817 S trombus aurisdianae var. adusta “Chemnitz”, Dillwyn, Descr. Cat. Recent Shells, London, vol. 2, p. 664. 1822 Strombus melastomus Swainson, Appendix to Cat. Shells of Mrs. Bligh, London, p. 8 (Pacific Ocean). 1823 Strombus aurisasini Dillwyn, Index Hist. Conchyl. Lister, London, p. 39 (no locality). Refers to Lister pi. 872, fig. 27. 1825 Strombus melanostomus Sowerby, Cat. Shells Earl of Tankerville, p. 68. Emendation of melastomus Swain- son. 1826 Strombus adusta Gray, in King Narrative Survey— Australia, London, vol. 2, appendix, p. 490. Refers to Conchyl. -Cab., vol. 10, figs. 1487-88. 1835 Strombus melanastomus Swainson, Exotic Conchology, 2nd issue, pt. 6, pi. 47. Types— Boding’s species is based upon figures 1487-88, vol. 10, Conchylien-Cabinet. Chemnitz’s locality of East Indies was evidently erroneous. Some old collections bear labels of Java, Mauritius and Amboina, but we believe these are manufac- tured locality records. We hereby designate Bowen, Queensland, Australia, as the type locality. Nomenclature— As indicated by our synonymy, the name aratrum Boding must take precedence over melastomus Swainson and melanostomus Sowerby. The name aratrum Martyn 1784 (applied to the species vomer Boding) is non-binomial and invalid because the figures are entirely hand pro- duced and not printed. Therefore, it does not pre- occupy Boding’s 1798 name. This species is the “adustus Chemnitz” of early authors. Records— AUSTRALIA: Queensland: Thursday Island ( USNM ) ; Cape York ( MCZ ) ; Green Island, Cairns ( ANSP ) : Pilot Point. Brampton Reef and Sinclair Bay, all near Bowen (ANSP); Dunk Id. (USNM); Bedford Beach (USNM); Port Douglas, 50 mi. north of Cairns (MCZ and ANSP); Turtle Bay, Cape Grafton, near Cairns ( MCZ ) ; Alexandra Reef (MCZ); Bustard Head (ANSP); Whitsunday Id. (USNM). Strombus bulla (Boding, 1798) (PI. 14, figs. 5, 6; pi. 101, fig- 5) Range— Byukyu Islands to Melanesia and Samoa. Remarks— We consider S. bulla to be a species separate from aurisdianae. Their ranges overlap without signs of interbreeding in the western Pa- cific. To the east of the Solomons one finds only bulla and to the west of the East Indies one evidently only finds aurisdianae. S. bulla fives in deeper water. The two are closely related. One could hazard a guess that bulla originally evolved as a Central Pacific subspecies, and later, as a full spe- cies, re-invaded the territory of the parent aurisdi- anae, although the reverse could also have been possible. The key to identifying features of this glossy smooth, lavender-tipped species are listed under the remarks of aurisdianae. Habitat— S. bulla occurs sparingly from just below the lowest tide line to a depth of 10 fathoms on clean sand bottoms. Description— Shell 49 to 72 mm. (2 to 3 inches) in length, solid, with a smoothish, glistening sur- face, with a posterior projection on the outer lip, with a strongly recurved siphonal canal and with a smooth, glossy, white and rose-orange aperture. Nuclear whorls 3, glossy, translucent, rounded, and Plate 104. Living animal of female Strombus bulla (Reel- ing). (from Quoy and Gaimard, 1833, pi. 51, fig- 1). [09 - 994] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 129 Plate 105. Radula of Strombus bulla (Roding), Dutch New Guinea. Fig. 1, central. 2, lateral. 3, inner marginal. 4, outer marginal. either whitish, tan or purplish in color. First 3 postnuclear whorls well-rounded, and neatly reticu- late. Following whorls with numerous axial, knobbed ribs crossed by about a dozen uneven spiral threads. The subsutural cord is very weak and is weakly crinkled. Shoulder of last whorl with 5 to 9 small, short, smoothish knobs (other knobs are glazed over by the parietal wall). There are no spiral rows of smaller knobs below. Base of shell with crowded, weak, spiral cords. Parietal glaze enamel-white, usually quite thick, and extending up nearly to or over the apical whorls. In some instances, the glaze may cover the entire spire. Columella straight, glossy, whitish and entirely smooth. Outer lip enamel- white, except for a lav- ender stain at the base. Deep interior of aperture reddish to rose-orange and smooth. Posterior pro- jection of the outer lip extending % way back as far as the spire or rarely slightly beyond. In some speci- mens it leans towards tbe apex. Siphonal canal re- curved from 85 to 95 degrees and slightly twisted to the right. Body whorl with large mottlings of light brown, and small specklings of white; rarely with mauve undertones. Some Philippine specimens may have an irregular dark chestnut-brown patch on the dorsum of the last whorl. Periostracum very thin, translucent, and usually worn away even in live specimens. Operculum stromboid. Animal tan with large white spots. Verge with a “heel on the distal blade. Radula 6 mm., and with 42 rows of teeth. Formula for New Guinea: 2-1-2; 1-3 (with- out peg); 4; 5. Measurements ( mm.)— length width no. whorls 73.0 37.4 11 ( large; Cebu Id. ) 61.5 33.5 9 + ( average; Okinawa Id 49.2 28.0 10 ( small; Luzon Id. ) Plate 106. Geographical distribution of Strombus bulla ( Roding ) . Synonymy— 1798 Lambis bulla Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 64, no. 814 (no locality). Refers to Conchyl.-Cab., vol. 3, fig. 840. 1811 Strombus laevis Perry, Conchology, London, pi. 13, fig. 4 (African Seas). 1826 Strombus lamarckii Gray, in King, Narrative Survey— Australia, London, vol. 2, appendix, p. 490. Refers to Conchyl.-Cab., vol. 3, fig. 840 and Seba, pi. 61, figs. 1, 2. Non Sowerby, 1842. 1840 Strombus lamarckii Swainson, in Lardner’s Cabinet Cyclopaedia, Treatise Malacology, London, p. 139. Non Sowerby, 1842. 1842 Strombus aurisdianae Linne, Reeve, Conchologia Sys- tematica, vol. 2, p. 206, pi. 251, fig. 4. 1843 S trombus guttatus “Martini” Kiener, Coquilles Vivantes, vol. 4, Strombus, p. 24, pi. 15, fig. 1 ( Mer des Indes); 1851, Reeve, Conch. Icon., vol. 6, Strom- bus, pi. 14, fig. 33. 1925 Strombus (Strombus) aurisdianae Linne, Oostingh, Mededeel. Landbouw. Wageningen, vol. 29, pt. 1, p. 55. 1938 S trombus (Euprotomus) aurisdianae Linne, Adam and Leloup, Result. Sci. Voy. Indes Orient. Neerl., vol. 2, fase. 19, p. 117. Types— Rodin g’s bulla is based upon figure 840, vol. 3, Conchylien-Cabinet. The latter gave no lo- cality. We hereby designate Cebu City, Cebu Is- land, Philippines, as the type locality. Nomenclature— This is the species referred to in the old literature as Strombus guttatus Martini, a non-binomial name. This was not validated until 1843 by Kiener, but had two earlier names, bulla Roding and laevis Perry. Many authors between 1847 and 1938 referred to this species as aurisdi- anae L., but on the basis of Gray’s 1826 revision we [09 - 995] 130 Euprotomus R. T. Abbott Strombidae have restricted the latter name to the commoner, rough species figured by Rumphius, pi. 37, fig. R. S. bulla is S. lamarcki of Gray not Sowerby, Hanley, Oostingh and Adam and Leloup. Records (see map, pi. 106) -RYUKYU ISLANDS: Itoma Jima Id. (MCZ); Okinawa Id. (Mrs. A. A. Scott, ANSP). TAIWAN: Hoko ( Kuroda, 1941, p. 97). PHILIPPINES: Luzon: Lusong Cove, Bataan, 7 fms.; off Corregidor Id., 6-10 fms.; San Miguel Id., Tabaco Bay, Albay Prov. (all duPont-Academy Exped., 1958, ANSP). Mindoro: Mansalay Bay (ANSP); Calapan (MCZ); Lubang (MCZ). Cuyo Id., Palawan (ANSP). Cebu City, Cebu Id. (A. B. Franco, ANSP). Sulu Archipelago: Bongao Channel, Sanga Sanga Id. (J. Root, ANSP); Jolo Id. (MCZ). Samar Id. (MCZ). INDONESIA: Amboina and Bouro Ids. (MCZ); Misol, Timor, Savu, Flores, Java and Banka Islands ( Oostingh, 1925, p. 56). DUTCH NEW GUINEA: Soepiori Id., Schouten Ids. and Aoerori Id., east Padaido Ids. (both NSF, 1956, ANSP). PAPUA: Finschhafen Bay (MCZ). SOLO- MONS: Ugi Id. (ANSP); Kieta, Bougainville Id. (W. J. Eyerdam, ANSP). NEW CALEDONIA: (MCZ). SAMOA: Tutuila Id. (ANSP). Fossil records— KENYA: Pleistocene: reef-limestone, east- ern shore of Mombasa Harbour (L. R. Cox, 1930, Mono- graph 4, p. 137, not figured). Also an unverified report (see Cox, above) from the Pleistocene of Dar-es-Salaam, Tan- ganyika, by Koert and Tornau, 1910). Strombus vomer subspecies vomer (Roding, 1798) (PI. 14, figs. 7, 8) Range— Ryukyu Islands and New Caledonia. Remarks— To date, this very attractive species has been recorded in fair numbers from only the Ryukyu Islands and New Caledonia. Despite its ap- parent absence over a stretch of 3000 miles, there is no discernible difference in specimens from these two distant places. It would not be entirely sur- prising if it were to turn up in the Philippines and Indonesia, although we believe that this species, which was once widespread in the Pliocene, is now shrinking in range and becoming extinct. This shrinkage has left two peripheral subspecies, ire- dalei in northern Australia and hawaiensis in the Hawaiian Chain. The two isolated colonies of vomer vomer may, in future geological times, pro- duce a further differentiation. This entire vomer complex presents an example of the various stages of development of allopatric species, with iredalei being the most advanced, and the vomer vomer colonies representing potential subspeciation. Strombus vomer is a polytypic species contain- ing three subspecies vomer (Roding) from the Central Pacific Arc, iredalei Abbott from northern Australia and hawaiensis Pilsbry from the Hawai- ian Chain. Iredale’s donnellyi is quite likely a slightly malformed or immature specimen of vomer Plate 107. The discontinuous geographical distribution of Strombus vomer vomer (Roding) in the Western Pacific Arc, its Australian subspecies, iredalei Abbott, and its Ha- waiian subspecies, hawaiensis Pilsbry. vomer. It was obtained dead near Sydney, Aus- tralia, from dredged piles of sand from the ship, “Triton”, which had been working in New Cale- donia under the command of Captian Comtesse. Other New Caledonian, as well as Australian, spe- cies occurred in dead condition in these sand piles, and the authenticity of their origins is very ques- tionable. Typical vomer is distinguished from the other members of the subgenus Euprotomus by the in- terior of its orange to lemon-yellow aperture which bears 45 to 50 smooth, rather even-sized, white spiral lirae, by the brown patch on the upper part of the parietal wall, and by the very weak or obso- lete rugae on the base of the columella. S trombus vomer iredalei and vomer hawaiensis have a whit- ish to light yellowish aperture with numerous, crowded, uneven-sized lirae, and the base of the columella in each usually has moderately devel- oped rugae. The former subspecies, from Australia, has an extra one or two spiral rows of weak, small knobs on the lower part of the body whorl and may or may not have black-brown stains on the parie- tal wall. Two of the lirae inside the upper part of the aperture are usually larger than the others. The smoothish, glossy exterior of vomer vomer and the rough, non-glossy exterior of iredalei are similar to the difference between the smooth Strom- bus bulla (Roding) and the rough S. aurisdianae. Habitat— Found by George and Mary Kline in New Caledonia in 4 to 10 feet of water on coral sand and coral rubble on the barrier reef in Touho [09 - 996] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 131 Plate 108. Radula of Strombus vomer vomer ( Roding), New Caledonia. Fig. 1, central. 2, lateral. 3, inner marginal. 4, outer marginal. Bay. Mrs. Anita A. Scott says (in lift.) it is un- common in sand at dead low tide on Okinawa Is- land, Ryukyus. Description— Shell 55 to 88 mm. (2 to 3M inches) in length, solid, glistening, knobbed, with a high spire and projecting upper and outer lip, and with an orange inner aperture over which are white, spiral lirae. Whorls 11. Nuclear whorls 2, smooth, translucent lavender and convex. First 4 postnu- clear whorls angulate, lavender, and with 10 to 12 spiral threads crossed by fine, axial threads which give the first 2 or 3 postnuclear whorls a micro- scopically cancellate appearance. At the angular periphery of the whorls, beads appear which be- come larger and fewer (7 to 9 in the body whorl; 9 to 10 in the penultimate; 11 to 16 in the whorl above this). Except for these knobs and for the 20 to 24 weak spiral cords at the base of the shell, the outer shell is smoothish, glossy, with a whitish back- ground over which are maculations, speckles, tiny bars and arrows and axial streaks of mauve-brown. Outer lip moderately flaring, thickened and with a single, whitish, long or short projection at the top. Upper parietal wall with a glaze of blackish brown (like scorched paper); lower half swollen and cream. Inner columella wall brownish orange and weakly lirate above, smooth in the center, and with 5 or 6 small teeth near the white base. Sipho- nal canal long and bent back about 90 degrees. To the left of the strong, deep stromboid notch there is a large, cream, glossy, twisted flange. Inside of entire outer lip, including the dorsal projection, smooth and enamel-white. Interior of aperture orange to lemon-orange over which are 45 to 50 distinct, raised, smooth, white spiral lirae. Perios- tracum thin, smoothish, translucent. Operculum stromboid, dark- to light-brown, slightly arching, and with 7 to 8 very small serrations. Mantle ridged, its edge smooth. Radula ribbon (shell 83 mm.) 8 mm. long, with 44 rows of teeth. Formula: 2-1-2; 1-4 or 1-5 (without peg); 5; 5 or 6. Verge not observed. Measurements ( mm.)— length width no. whorls 88.0 50.0 1 1 ( large; locality unknown ) 72.5 40.5 10 (average; New Caledonia) 55.1 36.0 10 ( small; Ryukyu Ids. ) Synonymy— 1784 Aratrum , T. Martyn, Universal Conchologist, London, vol. 1, pi. 1 (Friendly Isles), non-binomial. 1798 Lambis vomer Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 64, no. 821 (no locality). Refers to Conchyl.-Cab., vol. 10, figs. 1485-6. 1811 Strombus acutus Perry, Conchology, London, pi. 12 fig. 2 (Pacific Ocean). 1826 Strombus zelandiae Gray, in King, Narrative Survey— Australia, London, vol. 2, appendix, p. 490. Refers to Conchyl.-Cab., vol. 10, figs. 1485-86. 1821 Strombus pacificus Swainson, Exotic Conchology, Lon- don, pt. 3, pi. 17 (Friendly Isles); 1841, ibid., 2nd ed., p. 10, pi. 17. 1840 Strombus chemnitzii Pfeiffer, Kritisches Register Mar- tini Chemnitz Konch.-Kab., Kassel, p. viii, no. 7. Refers also to Conchyl.-Cab., vol. 10, figs. 1485- 1486. 1842 Strombus novae zelandiae Chemnitz, Reeve, Con- chologia Systematica, vol. 2, p. 206, pi. 250, fig. 2; 1844, Duclos, in Chenu, Illust. Conchyl., vol. 4, p. 11, pi. 7, figs. 5, 6; 1851, Reeve, Conch. Icon., vol. 6, sp. and fig. 35. 1869 Strombus nova-seelandia , Ch., Paetel, Molluscorum Systema et Catalogus, Dresden, second p. 46. ?1931 Euprotomus donnelUji Iredale, Records Australian Mu- seum, vol. 18, no. 4, p. 212, pi. 23, fig. 19 ( Sydney Harbour, N. S. W.). 1942 Strombus hirasei Kuroda, Venus, vol. 12, p. 8, figs. 3 and 4 on p. 6 (Okinawa Ids.). 1950 Euprotomus atratum Allan, Australian Shells, Mel- bourne, p. 99, pi. 17, fig. 3 [error for aratrum Martyn], Types— Roding’s species is based upon figures 1485 and 1486, vol. 10, Conchylien-Cabinet. Chem- nitz believed bis specimen was from New Zealand, but this is evidently erroneous. Two other localities are mentioned in the early literature, Friendly Isles [Fiji] by Thomas Martyn (1784) and Pulo Condore [120 miles S.E. of Cambodia, China Sea] in the Portland Catalogue (1786, p. 29 and p. 64). The latter two records are unconfirmed. We hereby designate Noumea, New Caledonia, as the type locality. The type of S. hirasei Kuroda is presum- ably in the Kyoto University collection and is from Okinawa. From the description and figures it ap- pears to fit within the limits of vomer vomer. The type of Euprotomus clonnellyi Iredale is presum- ably in the Australian Museum. Nomenclature— Thomas Martyn’s name Aratrum appears without a generic name and is evidently non-binomial. This is generally and rightfully re- jected because all of the illustrations and ruled [09 - 997] 132 Euprotomus R. T. Abbott S trombidae borders were hand painted and not printed. None of the four paintings 1 have examined of this spe- cies are the same. Records (see map, pi. 107) -RYUKYU ISLANDS: Yae- yama and Oshima, Amami-Osima Ids. (Y. Hirase, ANSP); Satsuma (MCZ); Okinawa (Mrs. A. A. Scott and A. R. Calm, ANSP). NEW CALEDONIA: Touho Ray (G. and M. Kline, NSF, 1959); Bourail (Mme. Reverce, ANSP). Brampton Reef, 19° 51' S.; 158° 20' E. (J. Brazier, 187f, p. 585). Kuroda, 1942, p. 8, questions a Wakayama-Ken, Kii, Honshu Id., Japan, record, since it was probably from an Okinawan fisherman. The Pulo Condore Id., South China Sea, record in the Portland Catalogue (1786, p. 29 and 64) is unconfirmed, but possible.] Fossil records— None reported. Strombus vomer subspecies hawaiensis Pilsbry, 1917 (PI. 14, figs. 9, 10; pi. 109) Range— Hawaiian Chain from Midway to Maui Island. Remarks and description— Shell 64 to 98 mm. (2% to 4 inches in length), very similar to vomer vomer, but differing in having a white to yellowish tinted aperture; in having finer, more numerous, crowded, uneven-sized spiral lirae (55 to 70 instead of 45 to 50 ) on the inside of the body whorl which may ex- tend to the edge of the outer lip; in having 10 to 15 weak teeth or spiral lirae at the base of the columella (instead of 5 to 6); in lacking the brown stain on the parietal wall; and in having stronger spiral cords on the lower half of the whorls, of which 2 to 4 show in the bottom half of the whorls in the spire. The length of the projection at the top of the outer lip is variable in adults, and usually has a longitudinal furrow down the middle inner side. Periostracum not seen. Operculum yellowish brown, thick, longitudinally furrowed and with a saw-toothed edge. Soft parts not examined. I am greatly endebted to several Hawaiian col- lectors for the gift and loan of specimens of this rare subspecies; Clifton S. Weaver, Dr. Tom H. Richert, Mr. Bobby Lee, Mr. John Duarte, and Mr. Crawford N. Cate. S. hawaiiensis, not hawaiensis, is a misspelling. Habitat— Lives on sand and coral rubble bottom from 3 to 21 fathoms. A rare subspecies, and rarely cast on shore. M easurements (m m .)— length width no. whorls 75.0 41.0 10 ( holotype ) 98.3 52.0 10 ( large; Maui Id. ) 83.5 41.5 11 ( average; Kauai Id 72.0 41.0 11 ( small; Oahu Id. ) 64.0 32.5 8 + ( small; Oahu Id. ) |§ Plate 109. Figs. 1 and 2, holotype of Strombus vomer hawaiensis Pilsbry, Hawaii. Natural size. Synonymy— 1917 Strombus hawaiensis Pilsbry, Proc. Acad. Nat. Sci. Philadelphia, vol. 69, p. 329, pi. 22, figs. 1, 2 (Pearl and Hermes Reef, Hawaiian Chain ) . 1952 Strombus hawaiiensis Pilsbry, Tinker, Pacific Sea Shells, Honolulu, p. 52. Types— The holotype is in the B. P. Bishop mu- seum, Honolulu. It was collected by Lt. W. H. Munter. One beaehworn paratype in ANSP no. 46724. The type locality is Pearl and Hermes Reef, Hawaiian Chain. Records (see map, pi. 107 )-HAWAIIAN CHAIN: Mid- way Id. (ANSP and MCZ). Pearl and Hermes Reef (BPBM). KAUAI: Haena (W. A. Bryan, ANSP); Milolii (W. A. Bryan); Wailua Bay ( John Duarte). MOLOKAI: Moomumi (W. A. Bryan). OAHU: Paumalu (Cliff Weaver); Waianae (W. A. Bryan); Ewa (T. H. Richert and ANSP). MAUI: southwest end ( Bobby Lee ) . Fossil records— RYU KY U ISLANDS: Gabusoga and Na- kosi, Okinawa Id. (lower part of Pliocene) (Nomura and Zimbo, 1936, Science Reports Tohoku Imp. Univ., Sendai, 2nd ser., vol. 18, no. 3, p. 259, pi. 11, figs. 26a, 26b, as S. aurisdianae L.). A specimen closely resembling hawai- ensis was collected by H. S. Ladd (station F 238, in USNM) of probable Miocene age near Nasongo, Viti Levu Id., Fiji. [09 - 998] November 23, I960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 133 Strombus vomer subspecies iredalei Abbott (new name) (PI. 14, figs. 11, 12) Range— Western Australia to the Gulf of Car- penteria, Australia. Retnarks and description— This subspecies ap- pears to be limited to about 2000 miles of coast along western and northern Australia. It is closer in morphological characters to the geographically distant hawaiensis than to the central, typical vomer vomer. S. vomer iredalei has an exterior which is less shiny and more rugose, with the body whorl bearing 2 to 4 fairly strong spiral rows of coarse beads or poorly developed knobs below the row of small, somewhat even-sized pointed knobs at the shoulder. The spiral cord just below the suture bears numerous, elongate beads. Interior of aperture whitish with numerous spiral, white lirae of unequal size and with 2 or 3 of these at the upper end being much larger than the others. Lower half of parietal wall and columella strongly swollen, cream, tan or rarely brownish orange, and usually with small rugae on the inner, lower end. Parietal wall not glazed over at the center, but may be bordered with brownish markings. The outer lip may be somewhat flaring or turned in- ward. The nuclear and early whorls are whitish to pinkish but otherwise like those of hawaiensis. Radula similar to that of vomer vomer, its form- ula 2-1-2; 1-4 (also 1-3); 5; 5. Verge not observed. Habitat— Little is known about its habitat or eco- logy. A live specimen was collected in shallow water at Broome by Dr. Hubert Lyman Clark of Harvard. Measurements (mm.)— length width no. whorls 76.5 42.0 11 (large; west Australia) 64.8 35.2 10 (average; Australia) 55.5 33.5 11 ( small; Broome, Australia 37.0 22.5 6 + ( small; Australia ) Synonymy— 1826 Strombus australis Gray, in King, Narrative Survey— Australia, London, vol. 2, appendix, p. 489 (Aus- tralia). Non Schroter, 1805. 1842 Strombus australis Sowerby, Thesaurus Conchyliorum, vol. 1, Strombus, p. 36, no. 53, pi. 9, figs. 96-97 (Australia); 1843, Kiener, Coq. Vivantes, vol. 4, Strombus, pi. 14, fig. 1; 1851, Reeve, Conch. Icon., vol. 6, pi. 14, fig. 34. Non Schroter, 1805. 1854 Strombus (Monodactylus) australis Sowerby, H. and A. Adams, Genera of Recent Mollusca, London, vol. 1 p. 259. 1885 Strombus australis Gray, Rrazier, Proc. Linn. Soc. New South Wales, vol. 10, pt. 1, p. 88. Types— The type of Strombus australis Gray is presumably in the British Museum of Natural His- tory in London. Australia is the type locality which we do not restrict until more is known about this subspecies. Nomenclature— Gray gave a fairly diagnostic Latin and English description, but somewhat con- fused the picture with badly punctuated remarks which contained some typographical errors (for “Martini, vii”, read “Martini, hi"). Some have felt that Gray referred his new species to Martini, Conchy!. -Cab., vol. 3, figs. 338, 339 and to Seba’s pi. 61, fig. 5, 6, but it may be noted under the Latin description that Gray put “Icon. ?” This meant he could not find an illustration of it in any of the iconographs available to him at that time. In view of the description, and subsequent illustra- tions of this species by Sowerby and Kiener, I am accepting australis Gray as described and illus- trated in our present monograph. Strombus australis Gray, 1826, is preoccupied by Strombus australis Schroter, 1805 (which is the high-spired, “ turritus ” form of Strombus vittatus). I hereby rename australis Gray, 1826, and australis Sowerby, 1842 as iredalei after Tom Iredale of Sydney, Australia. Records (see map, pi. 107 )-WESTERN AUSTRALIA: Rowley Shoals (J. Rrazier, 1885, p. 88); Nicol Ray (J. Brazier, 1885, p. 88). Geraldton (MCZ); mouth of False Cape Creek, La Grange Bay, Broome; Ridel! s Beach, near Broome; James Price Point, 35 mi. north of Broome (all V. Orr, 1958, ANSP ) . NORTHERN TERRITORY: off Darwin (A. R. Cahn, ANSP). Yirrkola, Arnhem Land (USNM). [09 - 999] 134 Euprotomus R. T. Abbott S trombidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [10-000] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 135 Subgenus Conomurex P. Fischer, 1884 Type: Strombus luhuanus Linne, 1758 This subgenus is limited to the Indo-Pacific area, and contains two species— the Pacific Ocean luhti- anus Linne, and the Indian Ocean decorus Roding. The latter has a subspecies, persicus Swainson, in the northwest section of the Indian Ocean. The subgenus may be a rather recent development, since it has not been recorded in the fossil record earlier than the Upper Pliocene of Indonesia. The shells are characterized by their somewhat conic shape, depressed spire, and rose to red aper- ture. The penis is scarlet to rose and has a prong arising from one corner of the distal blade. The operculum has 4 to 6 well-developed serrations. The lateral tooth of the radula has a very large cusp on the inner side and a small, pointed peg at the base. Synonymy— 1884 Conomurex “Bayle” P. Fischer, Manuel cfe Conchvli- ologie, fasc. 7, p. 670. Type by monotypy: S trombus luhuanus Linne, 1758. Strombus luhuanus Linne, 1758 (PI. 14, fig. 15; pi. 110) Range— Southeast Japan to Indonesia, Australia and east to Palmyra and Fiji. Remarks— This common, shallow-water, western Pacific species is readily recognized by its black- brown, smooth columella, its blood-red to bright orange-red aperture, and its rather heavy shell. Young or badly beachworn specimens may lack color on the columella and aperture. Specimens at the extreme range of distribution are inclined to be small and lacking the rich watermelon-red of the aperture. Such dwarf, whitish-mouthed eco- logic forms have been found in Vaucluse Bay, Sydney, Australia and on Palmyra Island, south of the Hawaiian Chain. I suspect the dwarfed Pal- myra Island luhuanus is an ecologic form rather than a very localized subspecies. Not all Palmyra specimens have a white aperture and not all have a high, extended spire. The latter character ap- pears to be correlated with the unusually heavy growth of calcareous algae on the spire during the animal’s growth, thus forcing an unnaturally rapid descent of the whorls. I call this the “Palmyra” form. Records and specimens of this species from the Mauritius and Madagascar area are most likely due to misidentifications or mixed labels. The In- dian Ocean counterpart to luhuanus is S. decorus (Roding). The latter has no black on the columella and its aperture is rose to orange with a white border. There are reports (von Martens, 1887, p. 189) of luhuanus in the Bay of Bengal which 1 have not verified. Habitat— This species is usually abundant wher- ever it occurs. Large colonies live in shallow water from the low tide mark to a depth of 30 feet, usually where the bottom consists of coral sand, coral rubble and patches of algae. It is found in Micronesian lagoons and in the bays of large is- lands, providing there are no muddy conditions. The animals give off great quantities of mucus when disturbed. Many shells from certain locali- ties have five Sabia conica (Schumacher) (Hip- ponicidae) attached to the outer shell. They are more commonly on the spire, but may be found on the body whorl. Their attachment causes a round, deep, irregular scar. Description— Shell 32 to 70 mm. ( 1 to 2 :K inches ) in length, solid, well-shouldered, conic, with a black columella and orange-red aperture, and with a thick, rough, brown periostracum. Color of outer shell (when brown periostracum is removed) white with 7 to f2 irregularly-sized spiral bands of light-brown which may contain axial, flame-like bars of brown. Aperture orange- to watermelon- red. Columella darkly suffused with black-brown. Whorls 8 or 9. Nuclear whorls 3, bulimoid, glossy, smooth, translucent-tan or translucent-rose. First Plate 110. Living animal of female S trombus luhuanus Linne. (from Quoy and Gaimard, 1833, pi. 51, fig. 3). [10-005] 136 Conomurex R. T. Abbott S trombidae Plate 111. Geographical distribution of Strombus decorus ( Roding ) and its Arabian subspecies, persicus Swainson, ^ and of the Pacific Ocean Strombus luhuanus Linne. synonymy postnuclear whorl with 15 to 18 microscopic spiral threads which are axially crossed by much smaller scratches. In later whorls, numerous fine, axial rib- lets develop. Spire with 8 to 10 small, whitish, swollen, former varices. After the fourth postnu- clear whorls, the rounded shoulder has about 20 short, even-sized, well-rounded, ribs per whorl. These disappear on the last whorl. Base of shell with about 2 dozen almost obsolete spiral threads. Interior of body whorl with about 100 very fine spiral lirae which do not reach the edge of the sharp outer lip. Stromboid notch usually well-de- veloped. A similar notch occurs at the shoulder, a little below the well-indented, almost channeled, 1758 Strombus luhuanus Linne Systema Naturae, ed. 10, p. 744, no. 432 (In O. Asiae); 1767, ed. 12, p. 1209, no. 500; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 264-265. 1788 Strombus luguanus Herbst, Natur. Abbild. der merkw. W firmer, vol. 9, p. 203, pi. 48, fig. 2 (no locality). S. luthuanus on plate caption. 1798 Lambis luhuana Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 61 (refer to Conchyl.-Cab., vol. 3, figs. 789, 790). No locality. ?1839 Strombus pusillus Anton, Verzerchniss der Conchylien, Halle, p. 86, no. 2812 (Young of luhuanus?) . 1884 Strombus (Conomurex) luhuanus L., Fischer, Manuel de Conchyliologie, Paris, fasc. 7, p. 670; 1929, Thiele, Handb. Syst. Weiehtierkunde, Tena, vol. I, p. 254. 1931 Conomurex luhuanus Linne, Iredale, Records Aus- tralian Mus., vol. 18, no. 4, p. 212; 1959, Kira, Coloured Ulus. Shells Japan, Osaka, p. 35, pi. 15, fig. 8. suture. Columella smooth, glossy, black-brown, but poorly developed. Periostracum thick, axially fim- briated, especially below the suture, tan to brown in color. Operculum stromboid, blackish brown, slightly arching, with 3 to 5 large serrations, and slightly less than one third the length of the shell. Radular ribbon 5 to 8 mm. long, wine-red to brown, with 42 to 46 rows of teeth; main cusps very large. Formula 2-1-2 or 3-1-3; 1-4 or 1-3; 4 to 6; 7 or 8. Eye peduncles usually equal in length. Head and body heavily speckled and mottled with crimson-red (preserved). Eye with a single red ring. Verge with a long, thin, thumb-like append- age. Verge scarlet-red with the laminated, distal pad yellowish. Measurements (mm.)— length width no. whorls 69.5 38.0 9 (large; Helen Reef, Carolines) 56.5 30.5 9 ( average; Samar Id., Phil. Ids. ) 32.2 18.0 6 + (small; Palmyra Id.) Types— According to Dodge, 1956, p. 265, the Linnaean collection in the Linnaean Society of London, contains two cotypes marked by Linnaeus himself. The original locality was given merely as “in O. Asiae”. We are restricting the type locality to Luhu Island, near Amboina and Ceram, Indo- nesia. This was the locality given by Rumphius on his pi. 37, fig. S and upon which Linnaeus named the species, luhuanus. Selected records (see map on this page for others; solid dots; specimens examined; open circles: literature records) — HONG KONG: Port Shelter (A. J. Staple, ANSP). THAI- LAND: all Gulf of Siam: Koh Samit; Koh Tao; Koh Samui; Koh Samet; Maprao Id. (all USNM). SINGAPORE: (Oostingh, 1923, p. 85). JAPAN: Hachijo Id., south of Tokyo; Shirahama, Wakayama Pref., Honshu Id.; Tomioka, Amakusa, Kyushu Id. (all T. Habe, ANSP). RYUKYU IS- LANDS: Shioya, Shanawan Bay, Okinawa (USNM). TAI- WAN: Taihoku-syu; Kiirun; Suo; Hoku; Tusyo (all Kuroda, 1941, p. 97). PHILIPPINES: common throughout the is- lands of Luzon; Lubang; Mindoro; Samar; Leyte; Catandu- anes; Marinduque; Basilan; Bilan; Santa Cruz; Palawan; Cebu; Panay; Mindanao; Sanga Sanga; Tawi Tawi; Siasi (all ANSP, USNM and MCZ). INDONESIA: Bouro Id., Moluccas; Roti Id., Timor; Wodo Id., Halmahera; Poeloe Boeton, Celebes (all MCZ). AUSTRALIA: Vaucluse Bay, [10-006] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 137 Sydney, N.S.W.; Green Id., Palm Id., Brook Id., Queens- land ('all ANSP). MARIANAS: Maug Id. (USNM); Saipan Id. (USNM); Apra Harbor, Guam Id. (A. J. Ostheimer, ANSP). CAROLINES: Yap Id. (C. O. Kile, ANSP); Ponape Id. (V. Wertley, ANSP); lagoon, Ifaluk Atoll; Lukunor Atoll; Ulithi (all USNM). NEW CALEDONIA: Baie de 1’Orphelinat, 8 ft. (G. and M. Kline, NSF, 1959). FIJI: Makongai Id. ( R. T. Abbott, MCZ). ELLICE ISLANDS: Funafuti (Hedley, 1899, p. 429). LINE ISLANDS: Pal- myra Id. ( George Vanderbilt, ANSP). [Records from Tahiti, Samoa, Seychelles, Mauritius, Reunion and Amirante Ids. are in our opinion based upon misidentifications or errone- ous locality data.] Fossil records— INDONESIA: Pleistocene of Timor ( Oos- tingh, 1923, p. 85), and Celebes ( K. Martin, 1890, p. 278); Pleistocene or Upper Pliocene (Tesch, p. 49, pi. 130, fig. 167). NEW HEBRIDES: Efate Id., Upper Pliocene (Ab- rard, 1946, p. 64). MARSHALL ISLANDS: Pleistocene at Bikini, drill hole 2, core 11, core piece 1, 180 to 185 feet ( H. S. Ladd, USNM). MARIANAS: Pleistocene (?), Guam and Tanapag, Saipan (USNM). TAIWAN: Byoritu Beds (Siko), Pliocene (Nomura, 1935, p. 179, pi. 9, fig. 23). Strombus decorus subspecies decorus (Roding, 1798) (PI. 14, fig. 13; pi. 113) Range— Durban, South Africa, to the Gulf of Ben- gal. Remarks— This species (formerly known as mau- ritianus Lamarck) is limited to the Indian Ocean and is evidently closely related to the Pacific Ocean luhuanus. Despite their resemblance to each other, I have considered them full allopatric species, Plate 112. Strombus luhuanus Linne, Sanga Sanga Island, Philippines. Fig. 1, central radular tooth (three views). 2, lateral. 3, inner marginal. 4, outer marginal. 5, verge. 6, operculum, showing scar from muscle attachment. since the differences are many and I have seen no intergrades from the Malayan region. S. decorus is more variable in size, shape and color pattern, especially in the same colony, than is luhuanus, and along its northern distribution in the Arabian Sea it has developed into a rather distinct, smooth- ish, angular-shouldered form which we identify as the subspecies, persicus Swainson (formerly belut- schiensis Melvill). Strombus decorus decorus is distinguished from luhuanus by the absence of black on the columella, by the delicate, light-orange to rose interior of the aperture, by the white border of the inside of the outer lip which produces a white interior to the siphonal canal, and by the usual presence of axial knobs on the shoulder on the last whorl. These smaller, more knobbed specimens, occuring spor- adically in Zanzibar colonies, are form coniformis Sowerby, 1842. Specimens often have circular, sunken scars caused by the attachment of the cap shell, S abia conica (Schumacher). Habitat— Live in colonies on coral sand, sponge and weed bottom from low water to 18 fathoms. Usually common wherever it occurs. Description— Shell 34 to 74 mm. ( lh to 3 inches ) in length, solid, heavy, somewhat conic in shape, with a white and brown-flecked columella, and a rose-tinted aperture. Color of outer shell variable, with a white to cream background over which are sparse mottlings, or nettings, or zigzag streaks, or Plate 113. Strombus decorus decorus (Roding), Portuguese East Africa. Fig. 1, central radular tooth. 2, lateral. 3, inner marginal. 4, outer marginal. 5, verge. 6, operculum. [10-007] 138 Cononmrex R. T. Abbott Strombidae indistinct spiral bands of light- or dark-brown or yellowish orange. Whorls 9 to 10. Spire usually moderately raised, rarely high, rarely low. Nuclear whorls 3, glossy, smooth, rounded. First three post- nuclear whorls with 7 to 9 microscopic, incised spiral lines crossing numerous, small axial riblets (about 24 per whorl). Top of spire with 6 to 8 small, whitish, rounded, swollen former varices. Whorls rounded in the spire and, in the penulti- mate whorl, bearing 14 to 16 small, well-rounded axial ribs or knobs. The last whorl may be smooth- ish at the rounded shoulder or bear 4 to 6 promin- ent knobs. Base of shell smoothish. Columella callus very thin, white or flecked with brown. In- terior of aperture light-orange to rose, with a broad white border. Inner wall of body whorl with ex- ceedingly fine, numerous spiral threads. Stromboid notch moderately deep. Posterior siphonal notch moderately developed and bordered above by a white tongue-like flap which is adherent to the pre- vious whorl. Periostracum thin, smoothish, trans- lucent-tan. Operculum stromboid, light-brown, arching, one third the length of the shell and with 6 or 7 sharp serrations. Radnla formula of Mozambique specimens: 2-1-2 (rarely 3-1-3); 1-3 to 1-5 (plus peg); 4 or 5; 5 or 6. Verge with a rose base and a yellowish distal blade which has a "heel" or slightly developed tlmmb- like process. Measurements (mm.)— length width no. whorls 74.3 40.5 9+ (large; Mozambique) 54.0 27.5 10 (average; Zanzibar) 33.5 18.0 9 (small; Zanzibar) Synonymy— 1798 Lambis decora Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 62, no. 777 (no locality). Refers to Conchyl.-Cab., vol. 10, figs. 1499 and 1500. 1807 Lambis miniata Link, Beschr. Natur.-Samml., Rostok, p. 108. Refers to Conchyl.-Cab., vol. 10, figs. 1499- 1500. 1807 Lambis flammea Link, ibid, p. 108. Refers to Conchyl- Cab., vol. 3, fig. 799 (a young specimen). 1821 S trombus cylindricus Swainson, Zoological Illustra- tions, series 1, vol. 1, pi. 53 (no locality); 1855, Berge, Conchylienbuch, Stuttgart, p. 231, ph 38, fig. 5. 1822 Strombus mauritianus Lamarck, Anim. sans Vert., vol. 7, p. 206 (lie de France [Mauritius]). Refers to Knorr, pt. 6, pi. 15, fig. 3; Lister, pi. 849, fig. 4a; and others. 1823 Strombus lutruanus Dillwyn, An Index to Hist. Con- chy! Lister, London, p. 38. Refers to Lister, pi. 849, fig. 4a. 1828 Strombus laevilabris Menke, Synopsis Method. Mol- luscorum, Pyrmonte, p. 41 (no locality). Refers to Conchyl.-Cab., vol. 10, figs. 1499-1500. 1842 Strombus coniformis Sowerby, Thesaurus Conchyl, London, vol. 1, p. 29, pi. 7, figs. 55 and 61 [the knobbed form]. Types— Roding’s species is based upon figs. 1499 and 1500 in vol. 10 of the Conchylien Cabinet. Chemnitz says that specimen came from Mauritius, which we now designate as the type locality. This is also the type locality for Lamarck’s mauritianus, the type of which is probably in the Museum d’- Histoire naturelle de Geneve. The whereabouts of Swainson’s type of cylindricus is unknown to us. Nomenclature— The more familiar name of mauri- tianus Lamarck, 1822, is unfortunately antedated by three other names, the earliest being decorus Roding. Lambris flammea Link, 1807, is based upon Conchyl.-Cab., vol. 3, fig. 799, and, together with the description, there is little doubt that it is a young specimen of decorus. Morch (1852, p. 63) was the first erroneously to associate Link’s name with floridus Lamarck (i.e. mutabilis Swainson). This error has been continued by many workers. Records (see map, pi. lll)-UNION SOUTH AFRICA: Durban, Natal (USNM). MOZAMBIQUE: Port Amelia (USNM, MCZ; Bazaruto Id. (MCZ); Inhaca Id., Delagoa Bay (W. Macnea, ANSP); Mozambique City ( K. Grosch, ANSP). ZANZIBAR: Chango Id.; Paje; Cliumbe Id.; Bawi Id.; Mnemba Id.; Pwakuu Id. (all NSF, 1957). KENYA: Malindi (USNM). SAUDI ARABIA: Muscat (ANSP). MADAGASCAR: Nossi-be (A. Chavane, ANSP); Nossi Fanihi; Tulear; Tamatave (all Dautzenberg, 1929, p. 471). INDIAN OCEAN ISLANDS: Diego Garcia lagoon, Chagos Ids. (Melvill, “Sealark”, 1909, p. 93). Marie-Louise Id., Amirante Isles (E. A. Smith, “Alert”, 1884, p. 502). Male, Maldives ( R. Jonklass, ANSP). Mauritius (ANSP, MCZ, USNM). CEYLON: (USNM). THAILAND: Koh Pipidon, Puket (Bay of Bengal) (USNM). INDONESIA: Poelau Berhala, Sumatra, 3 live specimens on coral reef, fide Tom- lin ( Oostingh, 1929, no. 39, p. 3); off Deli, Sumatra (Oostingh, 1930, no. 49, p. 4). BURMA: Mergui Archip. (von Martens, 1887, p. 189). Fossil records— According to L. R. Cox ( 1930, Monograph 4, p. 137): KENYA: Pleistocene, reef-limestone, eastern shore of Mombasa Harbour. TANGANYIKA: Pleistocene of Dar-es-Salaam. Strombus decorus subspecies persicus Swainson, 1821 (PI. 14, fig. 14) Range— Arabian Sea and Persian Gulf. Remarks— In the northern range of decorus, most specimens take on a characteristic flat-sided, conic shape, a reduction of the size of the axial riblets, and a reduction in the rose tint within the aperture. We are accepting this as a subspecies, since inter- grades exist. Young specimens closely resemble the genus Conus, but are recognized as Strombus by the minute varices in the apex. Habitat— Lives in sandy mud and coral sand from low water line to a depth of 10 fathoms. [10-008] Noveml>er 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 139 Description— Shell 40 to 51 mm. ( 132 to 2 inches ) in length, solid, conic, smoothish, with angular shoulders and lightly colored with light-brown net- ting and weak yellow-brown spiral bands. Whorls 9. Spire flat-sided, variable in height, and with about 6 small, swollen, former varices. Nuclear whorls unknown but probably like those in decorus. First two or three postnuclear whorls rose or whit- ish, with about 10 microscopic, spiral threads cross- ing larger axial riblets (about 22 per whorl). These riblets disappear three whorls from the last. Top of whorls flatfish, the shoulder angular. Sides of last whorl flat. Columella glossy white. Interior of aperture usually white, rarely tinted with rose. Stromboid notch weakly to moderately developed. Periostracum moderately to very thin, dull, tan to brownish. Operculum typical, with 6 to 8 serrations. Measurements ( mm.)— length width no. whorls 37.0 22.1 8 + (small; Liujab, Persian Gulf) 42.0 24.1 8+ (average; Jask, Persian Gulf) 51.1 27.5 9 (large; Dhahran, Persian Gulf) Synonymy— 1821 Strombus persicus Swainson, Zoological Illustrations, series 1, vol. 1, pi. 53 (Persian Gulf). June. 1844 Strombus ismarius Duclos, in Chenu’s Illustr. Conchyl., vol. 4, Strombus, p. 5, pi. 7, figs. 1 and 2 ( Nouvelle- Guinee). (dwarf form). 1898 Strombus (Conomurex) belutschiensis Melvill, Mem. and Proc. Manchester Lit. and Philos. Soc., vol. 42, no. 4, p. 37, text fig. ( Charbar, Mekran coast of Be- luchistan, 7 fms.). 1901 Strombus (Conomurex) beluchiensis Melvill, in Mel- vill and Standen, Proc. Zool. Soc. London for 1901, p. 380, pi. 21, figs. 13, 15 (in color). Emendation for belutschiensis Melvill. Types— The type of belutschiensis Melvill is pre- sumably in the British Museum of Natural History in London. Its type locality is Charbar, Mekran Coast of Beluchistan [now Iran or Persia], The type locality for persicus is “Persian Gulf”, but we do not know the whereabouts of Swainson’s type, unless, by chance, it is at Cambridge University, England. Duclos’ type of ismarius is probably in the Museum d'Histore naturelle de Geneve. Nomenclature— This subspecies appears to have received several names and one spelling emenda- tion, as is seen in our synonymy. The earliest valid name, persicus Swainson, 1821, is well illustrated and well described. We should like to point out that some argument might arise on whether or not the name persicus is a homonym of Humphrey, 1786 (A Catalogue of the Portland Museum, Lon- don, p. 3, item no. 15). The line reads: “15 Strom- bus Fusus, L. Persicus, or Persian Spindle, a pair fine. Lister. 854. 12." This is a Tibia, and if the name Persicus is to be interpreted as a species name, it would make Persicus Swainson a homonym and unavailable. However, in looking through the rest of the Portland Catalogue, we note that no other trivial or generic name is italicized, and that all geographical names are italicized. We feel cer- tain that Humphrey was merely giving the latin name for “Persian” and not intending a species name. S. ismarius Duclos is undoubtedly this spec- ies, and the locality of “Nouvelle-Guinee” is pro- bably erroneous. Strombus cailliaudi Jay, Tryon (Manual of Con- chology, vol. 7, p. 122) is probably not a Strombus. I concur with Tomlin ( 1937, Proc Mai. Soc. Lon- don, vol. 22, p. 224) that Jay’s species is the long- spired form of Conus mediterraneus Hwass. Jay’s type of Conus cailliaudi appears to have been lost or destroyed by fire. Records (see map, pi. Ill)— PERSIAN GULF: Zaal Id., Tarut Bay; Kuwait (both LTSNM); Ras Tanura (Mrs. B. J. Grantier, ANSP); Liujab, 3/2 fms. (Thaanum Coll’n. ). SALiDI ARABIA: Aden (Melvill and Standen, 1901, p. 381); Abu Musa Id. and Dubae, Trucian Oman Coast, 7 fms. (Melvill, 1898, p. 37). Fossil records— None reported. [10 - 009] 140 Gibberulus R. T. Abbott Strombidae Plate 114. Figs. 1-4, Strombus gibberulus gibbosus ( Rod- ing), Schouten Ids., Dutch New Guinea. 1 and 3, adults with shells of the snail, “Capulus,” attached. 2 and 4, imma- ture specimens. Figs. 5-7, Strombus gibberulus gibberulus Linne, Zanzibar. 5 and 7, immature. 6, adult. All natural size. [10-010] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 141 Subgenus Gibberulus Jousseaume, 1888 Type: Strombus gibberulus Linne, 1758 This subgenus is limited to the Indo-Pacific area, and contains only one species which, however, has broken into three subspecies— true gibberulus Linne from the Indian Ocean, albus Morch of the Red Sea area, and gibbosus Roding of the Pacific Ocean. The shells are characterized by a peculiar, abnormal coiling which gives them a distorted ap- pearance The apical varices are extremely broad. The penis has a “heel or accessory prong on the distal blade. The marginal radular teeth are rather long, delicate and with 5 to 10 dentitions. There is no basal peg on the marginal. The operculum is proportionately broad and with 6 to 9 large serra- tions. The fossil records of this group are not numer- ous, except in the Pleistocene of the Indo-Pacific. Abrard’s prae gibberulus from the Pliocene of the New Hebrides is possibly gibberulus gibbosus. I have serious reservations as to whether or not Plate 115. Strombus gibberulus gibbosus (Roding), New Caledonia. Fig. I, posterior corner of mantle margin, show- ing 4 mm. -long, lleshy appendage. 2, operculum, showing scar from muscle attachment. 3, under view of animal, showing operculum, foot, mucus slit at the anterior edge, egg-laying groove or notch on the right side of the anterior end, and the eye peduncles and proboscis. 4, verge. 6-8, radulae. 5, verge of S. gibberulus gibberulus Linne, Zanzibar. Oostrombus Sacco, 1893 (I Molluschi dei Terreni Terziarii del Piemonte e della Liguria, pt. 14, p.13; type by original designation: Strombus proble- maticus Michelotti, 1861 ) is related to the recent snbgenus Gibberulus. Cossmann, 1904, p. 16 notes the absence of Oostrombus in the Miocene. In any event, the name Gibberulus Jousseaume has prior- ity. Cossmann (1904, p. 14) and Wenz (1943, part 6, p. 1255) place the genus Thersitea Coquand, 1862, of the Eocene in the Fasciolariidae. I agree that it probably does not belong in Strombidae. Synonymy— 1888 Gibberulus Jousseaume, Memoires Soc. Zool. de France, vol. 1, p. 174. Type by original designation and monotypy: Gibberulus gibberulus Gmelin = gibberu- lus Linne, 1758. Strombus gibberulus subspecies gibberulus Linne, 1758 (PI. 14, fig. 28; pi. 114, figs. 5-7) Range— ( Entire species: East Africa to the Tua- motu Islands). Typical gibberulus: Indian Ocean, exclusive of South Africa, Red Sea and Australia. Remarks— This is one of the most abundant and widely distributed, shallow-water Strombus of the Indo-Pacific region. It is divided into three well- defined geographical subspecies: gibberulus from the Indian Ocean, albus from the Red Sea and gibbosus from the Pacific Ocean. All are character- ized by the peculiarly distorted, swollen penulti- mate whorl and the large size of the swollen, whit- ish, former varices in the upper whorls. Typical gibberulus is the largest (usually about 55 mm. in length, but ranging from 30 to 70 mm.); the outer shell is a drab yellowish to grayish tan with numerous fine spiral lines of white; when the columella has coloration, it is always limited to a faint purple well within the aperture; the spiral raised threads are usually pronounced on the varix of the last whorl. One out of several hundred speci- mens seen from East Africa had a rose-tinted mouth, others being purple-tinted. The subspecies gibbosus, confined to the Pacific- Ocean, is smaller (usually about 40 mm. in length, but ranging from 28 to 55 mm.); the outer shell is commonly banded or flecked with bright-yellows and browns; when the columella has coloration, the purple-brown, elongate splotch is always in full view; the spiral threads over the varix of the last whorl are usually weak or absent. The subspecies albus from the Red Sea is pro- 110 -015] 142 Gibbendus R. T. Abbott Strombidae Ocean, albus Morch of the Red Sea, and gibbosus (Roding) of the tropical western Pacific. portionately wider, lias an appearance of being more distorted, is usually milk-white on the out- side and the interior of the outer lip is a strong rose. I have examined several dozen animals from boti the Indian and Pacific Oceans and find consider- able variation in the radula and the number of dentitions on the opercula, so that no differences are meaningful. However, the verges appear to be separable, although not to a degree that would justify treating the Pacific Ocean subspecies as a full species. In most, if not all, of the Pacific speci- mens the right eye peduncle and the tentacle are as long or even slightly longer than the left peduncle and the left tentacle, while in the Indian Ocean specimens (and in all other species of S trombus we have examined ) the left is the longer. Abbott ( 1949, pp. 59-61 ) made a study of the shell length and frequency of the color phases in various colonies of the subspecies gibbosus, and concluded that the shells of males are only slightly smaller than those of the females. In three colonies examined, 40% were males, the remainder females. There was no indication of any sex-linking in the dark- or light-phase. Although shell size varies greatly within the subspecies, there is a fair amount of uniformity within each localized population. Some mean shell lengths of large population sam- ples are 33 mm. (north end, Agana Bay, Guam); 30 mm. for males, 31.5 for females ( S. W. side, Igurin Island, Eniwetok Atoll); 32 mm. (lagoon, Lae Atoll); 41.5 mm. (Puerto Princessa, Palawan Island); 48.0 mm. (Cebu City, Cebu Island); 34 mm. (Helen Reef, S. W. of Palau Ids.). We have noticed in specimens examined that, generally, smaller shells come from smaller atolls or areas with evidently less nitrogenous matter in the water. Similar ecologic size differences exist in the Indian Ocean gibbendus gibbendus. Habitat— S. gibbendus and its subspecies are co- lonial, shallow-water species living from the inter- tidal area to a depth of 10 fathoms. While they are more abundant on clear sand and weed bottoms just below the low water mark, they also occur in sandy patches on barrier reef Hats, on sandy lagoon bottoms, and also in muddy sand bottoms. In some areas the shells are festooned with algal growths, and in deeper, clearer water they are small and brightly colored. It is quite probable that the colo- nies migrate in the breeding season from deep to shallow water. This species lives in or near coral waters, and does not exist, as does S. canarium, in muddy, continental-like areas. [10-016] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 S trombus 143 Description— (of the Indian Ocean gibberidus gibberulus). Shell 30 to 70 mm. ( 1M to 2/1 inches) in length, solid, somewhat fusiform, and with dis- torted whorls. Color of outer shell light-tan to whit- ish with numerous, crowded spiral bands of darker tan. Columella smooth, white or with a faint flush of violet on the portion well within the aperture. Inner edge of outer lip white; within this is a broad violet or dark-purple, axial band crossed by numer- ous, irregular, whitish, spiral lirae. Deep within the aperture the shell is lightly flushed with violet. Posterior canal, short, deep, narrow, pointing di- rectly upward; hounded on the body wall side by a swollen, white callus. Stromboid notch deep. Spire angle 70°. Whorls 10; nuclear whorls 3, the first very small and slightly elevated, the remainder rounded, glossy, smooth and transparent. The first and second postnuclear whorls have only 5 to 6 incised spiral lines on the upper part of the whorl. Apical whorls with 10 to 11 very broad, swollen, whitish former varices which are crossed by 6 to 8 coarse, spiral threads. Penultimate whorl strongly and roundly shouldered, so that the last third of the body whorl descends rapidly, thus giving a distor- tion to the entire shell. Last part and base of body whorl with strong, raised threads. Periostracum translucent-tan to light-brown, smooth and usually glistening. Operculum stromboid, slightly arching, broad at the attachment end, light-brown, and with 7 to 8 sharp, rather long serrations, and slightly less than h the length of the shell. Radular ribbon 2 to 5 mm. in length, with 32 to 46 rows. Formula 4-1-4 or rarely 3-1-3; 1-4 (no peg); 7; 7. Verge up to 10 mm. in length, with a long thumb-like process. Measurements (mm.)— length width no. whorls 70.0 32.5 10 (large; Zanzibar) 55.0 28.5 9 (average; Zanzibar) 30.0 16.0 8 (small; Zanzibar) Synonymy— 1758 Strombus gibberulus Linne, Systema Naturae, ed. 10., p. 744, no. 443 (In. O. Asiae); 1767, Linne, ed. 12, p. 1210, no. 501; 1842, Sowerby, Thesaurus Con- chyl., vol. 1, pi. 6, fig. 19 (not 24-26); 1843, Kiener, Coquilles Vivantes, vol. 4, Strombus, pi. 28, fig. 1 ( not la). 1811 Strombus labiatus Perry, Conchology, London, pi. 12, fig. 3 (no locality). Non Roding, 1798. Types— Linnaeus’ type is in the Linnaean Society of London. Hanley implies that Sowerby’s figure 19 is the type, which is the Indian Ocean subspe- cies. We restrict the type locality to Zanzibar, Brit- ish East Africa. Nomenclature— To our knowledge, this is the first time that gibberulus has been separated into three subspecies. The Red Sea allnis has been long recog- nized. It is difficult to say if Linne included the Pacific Ocean subspecies in his gibberulus. The first available name for the Pacific subspecies is gibbosa Roding. Selected records (see accompanying map, pi. 116; solid dots: specimens examined; open circles: from the literature) —MOZAMBIQUE: Inhaca Id., Delagoa Bay (W. MacNae, ANSP). Mozambique City (Kurt Grosch, ANSP). TAN- GANYIKA: Mboa Maji, intertidal ( R. T. Abbott, ANSP). MADAGASCAR: Nossi-be (A. Chavane, ANSP: MCZ). Gloriosa Id. (UNSM). INDIAN OCEAN ISLANDS: Mahe, Seychelles (USNM). lie des Roches, Amirante Isles (E. A. Smith ) , 1884, “Alert”, p. 503 ) . Chagos Archip. ( Melville, 1909, “Sealark”, p. 93. CEYLON: Powder Bay, Trineomalee (G. and M. Kline, NSF ). ANDAMAN IDS: (MCZ). COCOS KEELING: (USNM). HONGKONG: Castle Peak Bay (A. J. Staple, ANSP). Fossil records— Raised Quartenary beaches of the Red Sea region (Nardini, 1934, p. 222). Pleistocene of French So- malia (Nardini, 1933, p. 171). Cox (1930, p. 138) reports gibberulus from the Quartenary of Mombasa, Kenja. Perim Island, Quartenary of Tanga, Tanganyika ( Koert and Tor- nan, 1910, p. 10). Strombus gibberulus subspecies gibbosus (Roding, 1798) (PI. 14, figs. 26; pi. 114, figs. 1-4) Range— From the Ryukyu Islands to Indonesia and Australia, and eastward to the Tuamotu Is- lands (hut not the Hawaiian Chain). Remarks and description— This is the very abun- dant and widely distributed Pacific subspecies whose differentiating characters are listed in the remarks under the typical gibberulus. The first few postnuclear whorls bear 1 to 12 microscopic spiral threads in gibbosus, in contrast to the 5 to 6 in- cised lines only on the upper two-thirds of the whorl in gibberulus. The aperture of gibbosus may be all-white or tinted with violet, or yellow or brown or orange, while that of gibberulus is almost always tinted with purple to violet. Operculum stromboid, slightly arching, about h to U the length of the shell and with 6 to 7 serrations. Rad- ular ribbon 4 to 8 mm. in length, with 28 to 48 rows. Formula variable: usually 4-1-4 (rarely 3-1-3 or 5-1-5); 1-4 (rarely 1-3) and no peg; usually 6 (rarely 3, 4, 5 or 7 denticles on the inner marginal); usually 7, but rarely 5 to 10 on the outer marginal. Verge with a slightly developed “heel" on the dis- tal blade. The intestines of some contained coral sand, for- aminifera and small, whole gastropod shells. How- ever, the species is undoubtedly herbivorous or omnivorous. [10-017] 144 Gibberulus R. T. Abbott S trombidae Measurements ( mm.)— length width no. whorls 58.0 25.0 10 ( large; Cebu Id. ) 40.0 18.2 9 (average; Suva, Fiji) 30.0 15.2 9 (small; New Caledon: Synonymy— 1798 Lambis gibbosa Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 62, no. 786. (no locality). Refers to Conchyl.-Cab., vol. 3, fig. 794. 1842 Strombus gibberulus Linne, Sowerby, Thesaurus Con- chyl., vol. 1, pi. 6, figs. 24-26; 1843, Kiener, Co- quilles Vivantes, vol. 4, pi. 28, fig. la (not figs. 1); 1949, Abbott, Nautilus, vol. 63, no. 2, pp. 58-61 (sexual dimorphism). 1957, Demond, Pacific Sci- ence, Honolulu, vol. 11, no. 3, p. 295, fig. 14. Not gibberulus gibberulus Linne. ?1946 Strombus (Canarium) praegibberulus Abrard, Ann. de Paleontologie, vol. 32, p. 63, pi. 4, figs. 30-31 (Plio- cene, Malelcula, New Hebrides). Types— To our knowledge no type exists. The species was based upon figure 794 in vol. 3 of Martini and Chemnitz, Conchyl.-Cab., Nurnberg, 1777 which had no locality. We hereby designate Cebu City, Cebu Island, Philippines, as the type locality. Nomenclature— Heretofore, this subspecies has been referred to as gibberulus Linne, but now prop- erly is gibberulus gibbosus Roding. Selected records (see accompanying map, pi. 116, for other records; solid dots: specimens examined; open circles: liter- ature records )— RUYKYU ISLANDS: Okinawa (A. A. Scott, ANSP). TAIWAN: Kurun (Kuroda, 1941. p. 97). PHILIP- PINES: common on most islands, Luzon, Cebu, Catandu- anes, Samar, Balabac, Palawan (all ANSP), Busuanga, Cuyo, Marinduque, Panay, Sanga Sanga (USNM). INDO- NESIA: Amboina; Halmahera Id., Batjan and Bouro Id., Molucca; Marissa, Celebes Id. (all MCZ). AUSTRALIA: Low Islands, Queensland (Tony Marsh, ANSP). Helen Reef, N. W. of New Guinea (V. Orr, NSF). CAROLINES: Truk Id. (Mrs. R. T. Gallemore); Ponape (Mrs. V. Wertley, ANSP). PHOENIX ISLANDS: Canton Id. (Frank Witts, ANSP). SAMOA: Toloa Pt., Upolu Id. (NSF). LINE IS- LANDS: Palmyra (USNM). SOCIETY ISLANDS: Moorea, Tahiti and Borabora ( H. A. Rehder, USNM). TUAMOTU ISLANDS: Raroia, Takume and Tikahau (J. P. E. Morrison, USNM); Marutea (Dautzenberg and Bouge 1933, p. 297). Plate 117. Strombus praegibberulus Abrard. Holotype, Plio- cene of New Hebrides, (from Abrard, 1946, pi. 4, figs. 30, 31). Natural size. Fossil records— Pleistocene, Lanai Id., Hawaii ( USNM ) . Pleistocene, Honolulu Harbor, Oahu Id. ( J. M. Ostergaard, 1928, p. 27). Pleistocene (?) of Aranit Id., Eniwetok Atoll, Marshall Id., drill hole A-l, 136.5 to 138.0 ft., II. S. Ladd (USNM). INDONESIA: Altena (1942, p. 47) reports gib- berulus from the Pliocene of Java, Obi and Timor Islands; also the Quarternary of Timor. The only record I have seen illustrated is that of Tesch's from the Pliocene of Timor which may be nearer the subspecies gibbosa ( Tesch, 1920, p. 49, pi. 130, figs. 166a,b). Tesch also records it from the Quarternary of Koepang, Timor and Macassar, Celebes. Abrard (1946, p. 63) records it from the Pliocene and Quaternary of the New Hebrides. Strombus gibberulus subspecies albus Morch, 1850 (PI. 14, fig. 27) Range— Limited to the Red Sea and Gulf of Aden. Remarks and description— This rather restricted subspecies differs from the typical Indian Ocean gibberulus in being smaller, much paler and usu- ally white on the outside, and in having the inside of the last whorl flushed with rose to carmine-rose. Rarely, the white columella is tinted with rose. There is an inch-long, narrow bar of purplish brown just below the suture on the dorsal side of the body whorl which is rarely, if ever, present in gibberulus, and rarely in gibbosus. Soft parts not available for study. Measurements — length width no. whorls 56.1 28.5 9 ( large; Eilat, Israeli ) 43.0 25.0 9 (average; Aqaba, Red Sea) 34.0 18.0 8 (small; Port Sudan) Synonymy— 1850 Strom bus albus “Mart.” Morch, Cat. Conchyl. C. P. Kierulf, Hafniae, p. 11. no. 264. Refers to Conchyl.- Cab., vol. 3, figs. 797, 798; 1852, Morch, Yoldi Catalog -, p. 62. (Red Sea). 1869 Strombus gibberulus var. rhodostomus “Morch” von Martens, in Decken’s Reisen in Ost-Afrika, vol. 3, Mollusken, p. 64 (Sansibar). Types— The species is based upon Martini’s fig- ures 797, 798, and no type of Morch’s exists. The type locality is “Red Sea”. Records (see map, pi. 116)— EGYPT: Ras Banas (USNM); Geb Zebara (ANSP); Gulf of Suez (USNM). SAUDI ARA- BIA: 20 mi. north of Jidda ( C. Aslakson, ANSP); SUDAN: Berbera (USNM); Port Sudan (ANSP). ISRAELI: Eilat, Gulf of Aqaba (A. Hadar). Fossil records— EGYPT: Pleistocene: Wadi Gueh, 80 ft. alt. (R. B. Newton, 1900, p. 508, pi. 20). FRENCH SO- MALIA: Pleistocene, Ravin de Baghenda; near d’Obock; d’hacoulta; Ras Bir; Ras Doumeira; Khor Ambada ( Abrard, 1942, vol. 18, p. 64, pi. 6, fig. 39). [10-018] November 23, 1960 INDO-PACIFIC MOLLUSCA, vol. 1, no. 2 Strombus 145 INDEX Looseleaf subscribers should place this index at the beginning of the fam- ily Strombidae. Strombus begins on p. [09-831]. accinctus Linne, 100 acutus Perry, 131 adansoni Defiance, 123 adusta Dillwyn, 128 adusta Gray, 128 adustus Reeve, 121 afrobellatus Abbott, 88 alata, Schumacher, 58 alatus Gmelin, 47 albus Morch, 144 aldrichi Dali, 117 Aliger Thiele, 53 altispirus King, 45 anatellus Duclos, 65 aratrum Martyn, 131 aratrum Roding, 127 athenius Duclos, 104 atratum Allan, 131 auratus Spalowsky, 122 aurisasini Dillwyn, 128 aurisdianae Linne, 125 australis Gray, 133 australis Schroter, 113 australis Sowerby, 133 belucbiensis Mel., 139 belutschiensis Mel., 139 bifrons Sowerby, 47, 89 bivaricosus Nomura, 46 bravardi Borehert, 53 bubo Lamarck, 123 bubonius Lamarck, 123 bulbulus Sowerby, 86 bulla Roding, 128 bulla tus Dodge, 86 buris Roding, 127 cailliaudi Jay, 139 campbelli Gray, 114 campbelli G. and P., 114 canarium Dickerson, 50 canarium Linne, 48 Canarium Schumacher, 63 cancellatus Pease, 84 carinata Roding, 99 carnaria Roding, 123 carnea Roding, 123 chemnitzii Pfr., 131 chipolanus Dali, 53 chrysostomus Kuroda, 127 columba Lamarck, 90 coniformis Sowerby, 138 Conomurex Fischer, 135 Conorium Jouss., 63 TO STROMBUS NAMES IN VOL. coronatus Defrance, 122 corrugatus A. and R., 81 costatus Gmelin, 53 crassilabrum Anton, 65 cristatus Lamarck, 61 cylindricus Swains., 138 decorus Roding, 137 deformis G. and P., 90 deformis Gray, 90 deformis Kiener, 92 deningeri Fischer, 114 dentata Roding, 85 dentatum Kira, 68 dentatus Linne, 84 dentatus Reeve, 68 denti Cox, 109 depauperata D. and B., 76, 77 deperditus Sowerby, 92 dilatatus Lamarck, 94, 123 dilatatus Swainson, 93 Dilatilabrum Coss., 62 Dolomena Iredale, 89 dominator P. and J., 53 donnellyi Iredale, 131 Doxander Iredale, 111 dubius Sowerby, 86 dubius Swainson, 86 elatus Anton, 45 elegans Sowerby, 80 elegantissima Rod., 121 epidromis Linne, 107 epimellus Duclos, 73 erythrinus Dillwyn, 79 Euprotomus Gill, 125 Eustrombus Wenz, 53 exnodosus Sacco, 93 expansa Tryon, 108 exustus Swainson, 121 fasciatus Born, 121 fasciatus Gmelin, 122 fennemai Martin, 95 llammea Link, 138 Hammeus authors, 73 flavigula Tryon, 121 Horidus Lamarck, 73 flosculosus Morch, 73 fortisi Brongn., 62 fragilis Roding, 86 fusiformis Sowerby, 78 galeatus Swainson, 53 galliformis P. and J., 53 Gallinula Herrm., 107 Gall inula Morch, 107 , NO. 2 gallus Linne, 53 gendinganesis Abrard, 65 gendinganensis Mart., 69 Gibberulus Jouss., 141 gibberulus Linne, 141 gibbosus Roding, 143 gibbus I. and C., 49 gigas Linne, 53 glaber Martin, 51 glabratus Sowerby, 45 goliath Schroter, 53 gracilior Sowerby, 47 granulatus Swainson, 117 guttatus Kiener, 129 haemastoma Sowerby, 82 haitensis Sowerby, 53 hawaiensis Pils., 132 hawaiiensis Tinker, 132 hebraeus Linne, 45 belli Kiener, 83 hellii Rousseau, 84 herklotsi Martin, 62 hirasei Kuroda, 131 incisus Wood, 64, 65 inflatus Martin, 62 iredalei Abbott, 133 isabella Lamarck, 49 ismarius Duclos, 139 japonicus Reeve, 113 javanus Martin, 102 jugosus Wood, 81 junghuhni Martin, 61 karikalensis Coss., 51 kemedjingensis Mart., 109 kieneri I. and C., 92 klineorum Abbott, 70 labiatus Perry, 143 labiatus Roding, 67 Labiostrombus Oost., 107 labiosus Gray, 96 labiosus Wood, 95 labrosus Menke, 45 laciniatus Dillwyn, 60 lacteus Rondelet, 54 laevilabris Menke, 138 laevis Dodge, 65, 66 laevis Perry, 129 Laevistrombus Kira, 47 lamarcki Sowerby, 127 lamarckii Gray, 127, 129 lamarckii Swainson, 129 latissimus Linne, 57 [09-651] 146 Index to Strombus R. T. Abbott Strombidae latus Gmelin, 122 leidyi Heilprin, 53 lentiginosus Linne, 117 Lentigo Jouss., 117 lineatus Lamarck, 121 lineolatus Wood, 121 liocyclus Dali, 63 listen T. Gray, 115 lituratus Menke, 103 lobata Roding, 60 lobatus Swains., 53 Lobatus Iredale, 53 luguanus Herbst, 136 luhuanus Linne, 135 lutruanus Dill., 138 madiunensis Mart., 114 maculatus Nuttall, 77 maculates Sowerby, 75 malekulensis Abrard, 92 marginatus Linne, 97 martapurensis Mart., 46 martini Oostingh, 50 mauritianus Lam., 138 maximus Martin, 56 melanastomus Swains., 128 melanostomus Sowerby, 128 melastomus Swains., 128 mekranicus Vied., 61 microurceum Kuroda, MS, 72 microurceus Kira, 71 mimasakensis Yok., 45 miniata Link, 138 minimus Linne, 105 minor Abrard, 92 mirabilis Sowerby, 115 moisei Cuvillier, 45 Monodactylus Herrm., 125 muricatus Martini, 65 mutabilis Swainson, 72 nodosus Borson, 123 nodosus Sowerby, 93 nova-seelandia Paetel, 131 novae zelandiae Rve., 131 ochroglottis Abbott, 74 olydius Duclos, 69 Oncoma Mayer-Eymar, 62 Oostrombus Sacco, 141 orientalis Duclos, 54 orosminus Duclos, 93, 94 orrae Abbott, 66 ostergaardi Pils., 75 otiolum Ieixlale, 68 overbecki Cox, 51 pacificus Swainson, 131 palabuanensis Mart., 92 palmata G. Fischer, 60 papilio Dillwyn, 120 parvulus Krumbeck, 45 persicus Swainson, 138 pertinax Duclos, 54 peruvianus Swainson. 53 picta lidding, 58 pipus Roding, 119 plicatus Bolten, 90 plicatus Lamarck, 68 plicatus lidding, 89 polyfasciatus Dill., 121 ponderosus Philippi, 56 praegibberulus Abrard, 144 preoccupatus Finlay, 123 problematicus Mich., 141 proximus Sowerby, 47 pugilis Linne, 47 pugiloides Guppy, 47 pulchellus Reeve, 92 pusillus Anton, 136 radians Duclos, 80 rana Roding, 118 rana Oken, 118 raninus Gmelin, 53 rembangensis Martin, 95 reticulata Link, 65, 68 rhodostomus von Mart., 144 robustus Sowerby, 100 rubicunda Perry, 80 rugosus Sowerby, 81 ruppelli Reeve, 80 rutteni Altena, 97 samar Dillwyn, 85 samar ensis Reeve, 85 scalariformis Duclos, 83 sedanensis Martin, 102 semperi Smith, 102 Septimus Duclos, 101 sibbaldi Kiener, 90 sibbaldi Sowerby, 91 sinuatus Humphrey, 60 sonde Dickerson, 70 sondeianus Martin, 46 sowerbyi Coss., 93 spinosus Martin, 123 spolongensis Mart., 77 stiva Roding, 127 striatogranosus von Mart., 127 Strombella Schliiter, 47 Strombidea Swainson, 63 Strombus S.S., 47 subalata Herbst, 121 subulatus Duclos, 121 succinctus Linne, 99 sulcata Watson, 114 sulcatus Anton, 45 sulcatus Holten, 112 sumatranus Woodward, 102 swainsoni Reeve, 94 taeniatus Q. and G., 49 taiwanicus Nomura, 95 tankervillii Swainson, 91 taurus Reeve, 58 terebellatus Sowerby, 87 teschi Cox, 97 textile Linne, 45 thersites Gray, 56 thersites Martin, 51 thersites Swainson, 55 thersites Wood, 56 Thersitea Coq., 141 tjilonganensis Mart., 62 togopiensis Cox, 102 triangulatus Martin, 115 tricornis G. Fischer, 54 tricornis Humphrey, 53 tricornis Lamarck, 54 Tricornis Jouss., 53 tridentatus Gmelin, 85 trigonus Grateloup, 53 troglodites Kiener, 106 troglodytes Lamarck, 106 tubercularis Anton, 45 tuberosus Martin, 62 turritus Lamarck, 113 turturella Roding, 49 uccinctus Linne, 100 unifasciatus Mart., 77 urceus Linne, 63 ustulatum Sebum., 64, 65 valdetuberculatus Inn., 53 vanicorensis Tryon, 49 vanikorensis Q. and G., 49 variabilis Swainson, 103 varinginensis Mart., 50 vittatus Linne, 111 vomer Roding, 130 yerbnryi Smith, 92 zebriolata A. and L., 72, 73 zelandiae Gray, 131 [09 - 652] & -A. INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 147 September 28, 1961 ' 6)L zst THE GENUS LAMBIS IN TIIE INDO-PACIFIC by R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia The nine known species of scorpion or spider conchs of the genus Lambis are limited to the tropical Indo-Pacific. Their general anatomy, egg masses and radulae are extremely close to those of Strombus, and, from a biological standpoint, some workers might wish to consider them a subgenus of Strombus. The main distinguishing characters are the great development of long spines or digitations on the apertural borders of the shell, and the ab- sence or great reduction of the posterior mantle filament. Like members of S trombus, the Lambis are shallow water, active snails, usually associated with algal-rich coral reefs. C. M. Yonge (1932) gives an excellent account of the herbivorous feed- ing methods and the nature of the crystalline style. Their diet is evidently largely restricted to the deli- cate red algae (see under Lambis lambis Linne)- The Strombus subgenus, Euprotomus , has more characters in common with Lambis than do any other subgenera. The verge is almost identical and the lateral radular tooth lacks a basal “peg” in both groups. However, the early co-existence of fossil Lambis crocata (Link) and Strombus (Euprotomus) aurisdianae Linne in the Pliocene of Zanzibar sug- gests that these two groups had already arisen, perhaps from a common stock, during the early Pliocene or late Miocene. Fossil records are not numerous, except in the Pleistocene where they are not uncommon from the Red Sea to the Hawaiian Islands. Today, they are extinct in Hawaii. H. S. Ladd obtained a Lambis lambis- like specimen in the Miocene-Pliocene, tuffaceous limestone of La- kemba, Lau Group, Fiji. Fossil records of Pterocera in the nineteenth century European literature refer to members of the family Aporrhaidae. There are no known Lambis species restricted to the Tertiary or earlier periods. Of the nine living species four are polytypic with subspecies of varying degrees of morphological separation, ranging from the distinctly different chiragra chiragra and chiragra arthritica to the weakly separated scorpius scorpius and scorpius indomaris. Most abundant and most ubiquitous is Lambis lambis which is the largest and dominant gastropod in many shallow-water areas of the Indo- Pacific. Equally widespread from East Africa to Polynesia are truncata (moderately common), digi- tata (rare) and crocata (uncommon). Three spe- cies have rather limited ranges: violacea (rare in Mauritius), robusta (rare in southeast Polynesia), and millepeda (abundant in the central part of the Western Pacific arc). Lambis , particularly lambis and truncata, are -A': Plate 118. Living animal of male Lambis lambis (Linne). Between the eyestalks and tentacles is the darkly colored proboscis. Below is the long muscular foot at the end of which is the corneous, sickle-shaped operculum. Arising from the back of the animal is the long, external verge or penis, (natural size; from Quoy and Gaimard, 1833, pi. 50, fig. 11). [10-051] 148 Lambis R. T. Abbott Strombidae used as food, both cooked and raw, in most areas of the Indo-Pacific. The common lambis is seen on sale alive in most country fish markets in the Philip- pines. Lambis millepeda is generally not eaten be- cause of its bitter taste. In Ceylon, according to Deraniyagala (1933, Ceylon Journal of Sciences, sect. C, Fisheries, vol. 5, p. 63), lambis and rugosa [? = chiragra ] are roasted in the shell and eaten by Tamil fishermen. The flesh of the Mullu Shanku or Lambis acts as a mild narcotic. Both bitterness and narcoticness may be associated with the algal diet of the animal. Accounts of the gross anatomy of Lambis, which do not differ materially from those of Strombus, have been published by M. F. Woodward (1894), R. Bergh (1895), Jean Risbec (1925) and J. H. Prince ( 1955 ) . The latter treats with the details of the eyestalks of Lambis lambis. Geographical Distribution of Sexual Dimorphism It has been pointed out by many workers that in Strombus, as well as in other dioecious proso- branchs, the shells of the males are smaller than those of the females. Abbott (1949, p. 59) brought attention to tire fact that the sexual difference in shell size varied in S trombus from colony to colony, and suggested that there might be a geographical distribution of the degree of difference. Our soft anatomy examinations of Lambis were limited to thirty specimens, but, together with measurements of over a hundred shells, suggest the existence of sexual dimorphism in the sculpturing and nature of the spines, as well as the expected difference in shell length. The most striking example occurs in Lambis lam- bis in the central part of its range in the area of the Philippines, the Palaus, Carolines, New Guinea, Indonesia and northeastern Australia. The males are from 30 to 45 percent smaller than the females, and are further characterized by two small knobs on the shoulder of the last whorl (rather than a welding of these two into a longer and higher knob ) and by proportionately smaller spines which hook in the plane of the outer lip in tire posterior direction of the apex (rather than long, upwardly curling spines). The differences are much less de- veloped both to the west toward East Africa and to the east into New Caledonia, Fiji and Tonga. When we examine chiragra as a species extend- ing from East Africa to Polynesia, we find that the Indian Ocean subspecies, arthritica, shows little sexual dimorphism (males slightly smaller), but Plate 119. Radulae of Lambis; left to right: two views of the central tooth; two views of the lateral; the inner mar- ginal; and the outer marginal. The formula in the text refers to the cusps on each tooth, e.g. for fig. 1: 2-1-2; 1-3; 4; 6. Fig. 1, Lambis lambis (Seychelles). 2, L. crocata (Zanzi- bar). 3, L. chiragra chiragra (Palau Islands). 4, L. truncata sebae (New Caledonia). All y 80. Plate 120. Verges from male Lambis. Fig. 1, Lambis lambis, showing enlargement of the distal pad and its cross-section (Palau Islands). 2, L. crocata (Zanzibar). 3, L. chiragra arthritica. All x 2. [10-052] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 149 Plate 121 Fig. 1 Lambis (Millepes) violacea (Swainson, 1821). Mau- ritius. Rare. 2 Lambis (Millepes) robusta (Swainson, 1821). Soci- ety Islands. Rare. 3 Lambis (Millepes) digitate (Perry, 1811). Upolu Island, Samoa. Uncommon. 4 Lambis (Lambis) lambis (Linne, 1758). A female from Cebu Island, Philippines. Abundant. 5 Lambis (Millepes) scorpius scorpius (Linne, 1758). Okinawa Island, Ryukyu Islands. Moderately com- mon from the East Indies to Samoa. Note small flange on left side of base of topmost digitation which is absent in the Indian Ocean subspecies, indomaris. 6 Lambis (Millepes) millepede (Linne, 1758). Cebu Island, Philippines. Common in the S. W. Pacific. 7 Lambis (Harpago) chiragra arthritica Roding, 1798. Mozambique. Uncommon in the Indian Ocean. 8 Lambis (Lambis) crocata (Link, 1807). Zanzibar Island. Moderately common in the Indo-Pacific. 9 Lambis (Lambis) truncate sebae (Kiener, 1843). Saipan Island, Marianas. Moderately common in the Red Sea and Western Pacific. 10 Lambis (Harpago) chiragra chiragra (Linne, 1758). The uncommon rugosa form found in certain males. 11 the same, but a male from the Palau Islands show- ing very weak columellar sculpturing. 12 Lambis ( Harpago ) chiragra chiragra (Linne, 1758). A female from Luzon Island, Philippines. (all 2/5 natural size; center shell, fig. 9, is 12 inches in length) [10-053] 150 Lambis R. T. Abbott Strombidae that the subspecies chiragra shows in most cases a great size difference, as well as a columellar differ- ence, in the Western Pacific. The male very com- monly takes on the so-called rugosci-like columella, while the large females have a smooth, rose colu- mella. Curiously, further to the east in the Line, Tuamotu and Society Islands, only the rugosa type has so far been found. I have not seen many speci- mens from there and have had no opportunity to examine soft parts, but I suspect that both males Plate 122. Young shells of Lambis. Fig. 1, L. truncata sebae (Tahiti). 2, L. truncata truncata (Mauritius). 3, L. lambis (Saipan Island). 4, L. chiragra chiragra ( Sanga Sanga Is- land, Sulu Sea). All natural size. and females have a rogosa-like columella. I have not been able to find any significant differences, other than size, in the shells of the males and fe- males of the other species of Lambis. List of Recognized Taxa Genus Lambis Roding, 1798 Subgenus Lambis s.s. Roding, 1798 lambis (Linne, 1758). Type truncata (Humphrey, 1786) subsp. sebae (Kiener, 1843) crocata ( Link, 1807 ) subsp. pilsbnji Abbott, 1961 Subgenus Millipes Morch, 1852 millepeda (Linne, 1758). Type digitata (Perry, 1811) scorpius (Linne, 1758) subsp. indomaris Abbott, 1961 robusta ( Swainson, 1821 ) violacea ( Swainson, 1821 ) Subgenus Harpago Morch, 1852 chiragra (Linne, 1758) subsp. arthritica Roding, 1798 Bibliography Alcasid, G. L. 1947. A Review of Philippine Strombidae. Philippine Jour. Science, vol. 77, no. 2, pp. 179-203. Butot, L. J. M. 1955. Duivelsklauwen, boksbeugels, schor- pioenen of enterhaken. Penggemar Alam, vol. 35, pp. 71- 84, 3 pis. Gabb, William M. 1868. An Attempt at a Revision of the two Families Strombidae and Aporrhaidae. American Journal of Conchology, vol. 4, pt. 3, pp. 137-149. Gill. Theodore. 1870. On the Pterocerae of Lamarck, and their Mutual Relations. American Journal of Conchology, vol. 5, pt. 3, pp. 120-139. Prince, J. H. 1955. The Molluscan Eyestalk, Using as an Example Pterocera lambis. Texas Reports on Biology and Medicine, vol. 13, no. 2, pp. 323-339, 10 figs, llisbec, Jean. 1925. De 1’Anatomie de Trois Strombides et du Modulus candidus Petit. [Pterocera lambis L.]. An- nales du Musee d’Hist. Nat. de Marseilles, vol. 20, pp. 186-201, pi. 29. Woodward, M. F. 1894. On the Anatomy of Pterocera, with some Notes on the Crystalline Style. Proc. Malacological Soc. London, vol. 1, pp. 143-150, pi. ix. Yonge, C. M. 1932. Notes on Feeding and Digestion in Pterocera and Vermetus, with a Discussion on the Occur- rence of the Crystalline Style in the Gastropoda. Great Barrier Reef Exped., vol. 1, no. 10, pp. 259-281. [10-054] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 151 Subgenus Lambis Roding, 1798 Type: Lambis lambis (Linne, 1758) The typical subgenus of Lambis contains three species each of which is characterized by a smooth columella and an inner body wall which lacks spiral lirae. There are 7 digitations, 6 of which arise from the edge of the outer lip, and the seventh being an extended anterior siphonal canal. All three species, lambis, truncata and crocata are moderately com- mon to abundant over a large part of the Indo- Pacific. The latter two are polytypic, one having a subspecies in the Indian Ocean, the other a sub- species in the Marquesas Islands, Polynesia. The earliest fossil records of Lambis belong to this sub- genus, one of these being Lambis crocata (Link) from the Pliocene of Zanzibar. Synonymy — 1798 Lambis Roding, Museum Boltenianum, Hamburg, pt. 2, p. 61 (Type by absolute tautonomy: Lambis lambis Gmelin = Linne; p. 66). 1799 Pterocera Lamarck, Memoires Soc. d’Hist. Naturelle Paris, (Vol. 1), p. 72. Type by monotypy: S trombus lambis Linne, (and by Children, 1823, p. 127); 1929, Thiele, Handbuch Syst. Weicht., vol. 1, P- 255; 1940, Wenz, Handbuch der Palaont., vol. 6, pt. 4, p. 946. 1807 Pteroceras Link, Beschr. Naturalien-Samml., Rostock, part 3, p. 109. (type hereby selected: Pt. lambis Linn. ). 1810 Pteroceres Montfort, Conchy liologie Systematique, Paris, vol. 2, p. 607 ( substitute name for Pterocera Lamarck, 1799). Invalid type by original designa- tion: P. scorpius Linne. 1823 Digit ata O. Fabricius, Fortegnelse over Fabriciusses efterladte Naturalien, Copenhagen, p. 86. (nude name ) . 1829 Pterocerus Brongniart, Tableau Terrains, p. 410. 1852 Heptadactylus “Klein”, Morch, Catalogus Conchyl. . . . Yoldi, Hafniae, p. 60 (type hereby designated: lambis L. ); 1859, Chenu, Manuel de Conchyl. et Paleont. Conchyl., Paris, vol. 1, p. 259 (type by monotypy: Pterocera lambis Linne). 1878 Heptadactylus Kobelt, Illustrirtes Conchylienbueh, vol. 1, p. 106 (type hereby designated: Pterocera lambis Linne ) . Nomenclature — The genus name Pteroceres Montfort, 1810, must evidently be interpreted as a substitute name for Pterocera Lamarck. Montfort customarily changed the gender of Lamarck’s femi- nine genera to masculine. He refers to Lamarck’s genus. A similar case was similarly interpreted in Opinion 120, I.C.Z.N., Smithsonian Misc. Coll., vol. 73, no. 7, p. 29 (The Status of Achatinus Montfort, 1810). Lamarck’s once well known name, Pterocera, is a junior synonym of Lambis and has not been in general use for over twenty years, despite Thiele’s and Wenz’s usage in 1929 and 1940, respectively. Lambis lambis (Linne, 1758) (PI. 121, fig. 4; pi. 118; pi. 123, figs. 1-3) Range — East Africa to Micronesia and eastern Melanesia. Remarks — This is the commonest and one of the most widely distributed species of Lambis. It is one of the few marine gastropods to exhibit pronounced sexual dimorphism, other than size, in its shell Plate 123. Male and female shells of Lambis lambis (Yap view of females. % natural size. Island, Carolines). Fig. 1, male. 2 and 3, dorsal and side [10-057J 152 Lambis R. T. Abbott Strombidae characters. The size of the shell of the male is usu- ally about 60 percent that of the shell of the female. The degree of sexual dimorphism varies geographi- cally. It is most pronounced in the Caroline Islands and in most, but not all, areas of the Philippines, Indonesia, Queensland and the Solomons. The dif- ferences are less obvious in the Indian Ocean and at the extreme eastern end of the range in New Caledonia, Fiji and Tonga. In extreme examples in the male, the lowest three labial digitations are short and hooked posteriorly, while in the female, they are two or three times as long and are strongly curved upward. The female usually has a very large, pinched, somewhat-bilobed knob on the dor- sal part of the shoulder. In the male this is reduced to two small, equal-sized nodules. The two most posterior digitations are usually more widely sepa- rated in females than in males. It is possible that the upturning of the digitations in the females facilitates mating. When Lambis is in the position of coition, the right anterior end, or region of the “stromboid notch,” of each shell is approximately in juxtaposition. A closer contact would be possible because of the shortness of the digitations of the male and because the female’s digitations are curved upward out of the way. There is considerable geographical variation in the shape and size of the dorsal knobs in females; within a colony there is a uniformity. At Yap and Ponape Islands in the Carolines, the dorsal knob is very large, axially pinched, and bordered below by a smaller, sharp ridge. In Zanzibar, the dorsal knob is not as high as and is more elongate than those in most East Indian and Western Pacific specimens. In general, there are three color forms which occur throughout the range of the species in either sex — light-cream with sparse paintings of light- brown; a cream background with heavy mottlings of dark purplish brown; and an over all light- brown with flecks of white. Intergrades are not un- common within any colony. In the young of some, but not all, shells there is a large burnt-brown splotch on the upper section of the inner parietal wall. Couturier’s (1907, p. 154) reference to “Ptero- cera lambis ” from Mangareva and Tahiti undoubt- edly are specimens of truncata sebae Kiener. Al- though lambis has been reported from the Red Sea on several occasions, I have not seen authenticated specimens from there. C. M. Yonge (1932, p. 260) discussed in detail the herbivorous habits of this species, and included a detailed account of the crystalline style. The ani- mal feeds only on the fine red algae and not on the gross brown algae or eelgrass. A powerful cellulase digestive enzyme is present in the stomach. Yonge identified his species as “ Pterocera crocata,” but unquestionably he had Lambis lambis (Linne), since he figures the latter species in his 1930 book on “A Year on the Great Barrier Reef” (p. 83, pi. XXV, fig. C). He states (in 1932, p. 260) that his species is abundant and the most characteristic member of the fauna of the reef flat and mangrove areas of Low Isles on the Great Barrier Reef. The length of the shell ranged from 180 to 200 mm. and the aperture was red. All of these features are char- acteristic of Lambis lambis in Queensland and do not apply to the uncommon, smaller, outer reef crocata ( Link ) . I have been unable to verify the literature rec- ords for this species in the Red Sea. R. Sturany (1903, p. 46) reported the less common L. truncata from ten Red Sea localities, but evidently failed to find L. lambis. Miss Virginia Orr made the following observa- tions on the egg mass being laid by a female lambis at Poum, New Caledonia, on January 4, 1961: “the tangled clump was laid under an old slab of coral in 2 feet of water. The clear jelly tube was several feet in length and about 1 mm. in diameter. The orange eggs were about 0.3 mm. in diameter and there were about 30 of them per 10 mm. of tube.” Risbec also briefly described the egg mass of lambis which he says is most commonly laid in February in New Caledonia (Risbec [in Vayssiere], 1927, Ann. Mus. d’Hist. Nat. Marseilles, vol. 21, p. 185). Habitat — L. lambis occurs in colonies on reef flats and on sand or coral-rubble bottom, usually in association with algae, at depths ranging from the zone of low tide to several fathoms. Demond ( 1957, p. 297) says it “lives in sand on both seaward and lagoon reef flats, and on sandy lagoon floors, rang- ing from a few feet below low tide line to depths of 10 feet” in the Marshall Island atolls. M. Couturier ( 1907, p. 154 ) records it at depths of 3 to 5 fathoms in the Tuamotu Islands, but I believe that he was observing L. truncata sebae. Melvill (1909, p. 94) reported a specimen at 30 fathoms at Cargados Carajos, Indian Ocean, but this may have been a dead shell. Description — Shell, including the digitations, 90 to 200 mm. (about 3% to 8 inches) in length, with 6 slender, labial digitations which may be either short and bent posteriorly or long and bent up- wards towards the dorsum. Siphonal canal moder- ately long and slightly twisted to the left. Whorls 10 to 11. Nuclear whorls smooth. Postnuclear whorls [10-058] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 153 slightly concave above the carinate shoulder. The carina may bear numerous small beads or nod- ules. Spiral sculpture between the indented suture consists of about a dozen, irregular, microscopic threads. Last whorl with strong, blunt nodules on the shoulder. Mid-area of last whorl with a spiral row of 3 or 4 small nodules or rarely a raised, elon- gate nodule. Aperture and columella smooth, ex- cept for 4 to 7 weak lirae at the posterior end of the aperture, just behind the deep, smooth pit. Outer lip crenulated in young specimens, but smooth in old specimens. Color of aperture uniform tan, cream, or rarely orange tan or rose. A black- brown blotch may or may not be present on the upper part of the columellar wall of young speci- mens. Color of outer shell variable: cream with light- or dark-brown flecks and maculations; or mottled and spotted with purple-tan or blue-black. Periostracum thin, smoothish, translucent-tan. Oper- culum corneous, brown, elongate, arching, and without serrations. Radula formula: 2-1-2; 1-4 or 1-3; 5 or 6; 5 or 6. Verge as long as the aperture ( see pis. 119 and 120). Measurements (mm.) — (including digitations) length width no. whorls 200 111 10 + (large; north Ceylon 180 100 10 + (average; Luzon Id.) 92 43 10 (small; Palau Ids.) Synonymy — 1758 Strombus lambis Linne, Systema naturae, ed. 10, p. 743, no. 425; 1767, ed.' 12, p. 1208, no. 493 (O. Asiae); 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 250-251. 1798 Lambis lambis Gmel., Roding, Museum Boltenianum, Hamburg, pt. 2, p. 66, no. 844. Refers to Conchyl. - Cab., vol. 3, fig. 855, 888. 1798 Lambis lobata Roding, loc. cit., p. 68, no. 872. Refers to Conebyl. -Cab., vol. 3, fig. 902. Non lobata Rod- ing loc. cit., p. 65, no. 826, which is Strombus sinuatus Humphrey. 1798 Lambis lamboides Roding, loc. cit., p. 66, no. 845. Substitute name for lambis Gmelin, 1791. 1798 Lambis cerea Roding, loc. cit., p. 66, no. 846. Substi- tute name for lambis Gmelin, 1791. 1798 Lambis hermaphrodita Roding, loc. cit., p. 68, no. 875. Substitute name for lambis Gmelin, 1791. 1798 Lambis laciniata Roding, loc. cit., p. 68, no. 877. Sub- stitute name for lambis Gmelin, 1791. 1798 Lambis maculate Roding, loc. cit., p. 67, no. 854. Re- fers to Conchyl-Cab., vol. 3, figs. 858, 859. 1817 S trombus lambis Linne, Dillwyn, Descriptive Cata- logue Recent Shells, London, vol. 2, p. 658. 1826 S trombus camelus “Chemn.” Gray in King, Narrative Survey— Coasts of Australia, London, vol. 2, ap- pendix, p. 490. ( refers to Conchyl. -Cabinet, vol. 10, fig. 1478); 1830, Menke, Synopsis Method. Mol- luscorum, ed. 2, Pyrmonte, p. 72. 1842 Pteroceras lambis Lin., Sowerby, Thesaurus Conchyl., vol. 1, Pteroceras, p. 41, pi. 11, figs. 5 and 7 not 6. 1932 Pterocera crocata, Yonge, Great Barrier Reef Exped., Scientific Reports, vol. 1, no. 10, pp. 260-267. [feed- ing and digestion]. Types — Hanley ( 1855, p. 267 ) states that a speci- men like that figured in Sowerby’s Thesaurus Conch., vol. 1, pi. 1, fig. 5 is marked as lambis in the Linnaean collection in London. From that fig- ure and from Linne’s description it appears that this species was founded upon a male specimen. We restrict the type locality to Amboina, Indonesia. Nomenclature — Most of the synonyms of lambis Linne refer to the male shell. This is true of all of Roding’s names listed in our synonymy. Among the early figures of the male are: Schroter, 1782, Musei Gottwaldiani, pi. 21, figs. 142, 142; Rumphius, pis. 36A and 35E; Lister, pi. 866, fig. 21; Argenville, pi. 14, fig. E; Regenfuss, pi. 4, fig. 45 [excellent]; Knorr, pt. 1, pi. 28, fig. 1 [excellent]; and the Con- chyl.-Cab., vol. 3, figs. 858, 859, 902, 903. It is also figured in Rutot, 1955, pi. 2, fig. 2; Rlainville, 1827, pi. 25, figs. 4, 4a [excellent]. The female shell, with its long recurved, lower digitations, has received only one other name: ca- melus Chemnitz, 1788 [non-binomial] which was validated in 1826 by Gray. It has been figured in: Conchyl. -Cab., vol. 10, fig. 1478; also vol. 3, fig. 855; Schroter, 1782, pi. 21, fig. 139a; Rumphius, pi. 35F; Plate 124. Geographical distribution of Lambis lambis be based upon specimens of L. truncata sebae (Kiener). ( Linne ) . The Red Sea literature records ( open circles ) may [10-059] 154 Lambis R. T. Abbott Strombidae Reeve, 1851, pi. 5, fig. 8; Abbott, 1949, p. 324 ( Sci- entific Monthly, vol. 69). It may be pointed out that the use of the terms “male” and “female” among native collectors, even today, and among early writers, such as Rumphius, refers not to the sex of the animal but to the im- mature or light-weight (“female”) and the older, heavier (“male”) specimens. Selected records (see accompanying map for others; solid dots; specimens examined; open circles: literature records) — SAUDI ARABIA; Muscat, Gulf of Oman ( Melvill and Standen, 1901, p. 381). MOZAMBIQUE (von Martens, 1880, p. 279). TANGANYIKA: Inner Sinda Id. ( R. T. Ab- bott, USNM). ZANZIBAR: common around the island; Ki- wengwa; Chumbe Id.; Mnazi Moja; Paje; ( NSF, 1957). RED SEA: Suakin; Aqaba; Massana (all Issel and T.- Canefri, 1876, p. 357). Persian Gulf (MCZ). MADA- GASCAR: Nossi-be (A. Chavane, ANSP). Cargados Carajos (Melvill, 1909, p. 94). Mauritius (N. Pike, MCZ). MAL- DIVES: Gan Id., Addu Atoll; North Male Atoll (both Yale Peabody Mus.). INDIA: Bombay ( Homell, 1951, p. 87). CEYLON: Fort Frederick, Dutch Bay (Yale Peabody Mus.); Weligama Bay; Hikkaduwa; Eluvativa Id. (all G. and M. Kline, NSF, 1957). ANDAMANS: Port Blair; Long Id. (both Brit. Mus.). THAILAND: Phuket, Bay of Bengal (F. N. Crider, ANSP). SINGAPORE: Raffles Light ( R. D. Purchon, ANSP). RYUKYU IDS.: Shioya, Shanawan Bay (USNM). TAIWAN: Hoko ( Kuroda, 1941, p. 98). PHILIP- PINES: common throughout the islands of Luzon, Samar, Cebu, Catanduanes, Mindoro, Bohol, Masbate, Burias, Panay, Basilan, Lubang, Marinduque, Polillo, Palawan, Mindanao, Jolo, Sanga Sanga (all ANSP, USNM and MCZ). INDONESIA: Pulu Pandjang and Biliton Id., west Sumatra; Madura Id. and Bay of Batavia, Java; Obi Id., Moluccas; Lintido, Celebes (all ex Rijksmus. Nat. Hist., Leiden, ANSP) Banka, Kangeang (east of Madura), Flores, Timor, Amboina, Nusa-Laut, Halmahera, Waigen, Kei Islands (all Oostingh, 1923, p. 78). COCOS KEELING ATOLL (A. R. Whitworth, ANSP). AUSTRALIA: Western Australia: Cape Leveque (A. R. Whitworth). Northern Territory: Darwin (A. R. Whitworth. ANSP). Queensland: Rubber Reef, Port Douglas (Tony Marsh, ANSP); Hayman Id., Brook Id., Orpheus Id., Palm Ids. ( all H. A. Pilsbry, ANSP); Mossman (ANSP); Black Reef and Lupton Reef, Whitsunday Group (MCZ); Green Id., near Cairns (MCZ). DUTCH NEW GUINEA: Mios YVoendi Atoll, Padaido Ids.; Sowek, Soepiori, Schouten Ids.; Aoeri Id. (all NSF, 1956). Port Moresby, Papua ( P. Coleman, ANSP). ADMIRALTY IDS.: Koruniat (ANSP). SOLOMONS: Choiseul, Malita, Bougainville, Shortland Ids. (all W. J. Eyerdam, ANSP). NEW HEBRIDES: Mallicolo Id. (G. Massoulard, ANSP). NEW CALEDONIA: Touho Bay; Yate; Bourail (all G. and M. Kline, NSF, 1959). BONIN IDS.: Chichi Jima (Y. Kondo, USNM). MARIANAS: Saipan (ANSP); Apra Har- bor, Guam Id. (A. J. Ostheimer, NSF, 1953). PALAUS: common throughout the islands of Koror, Babelthuap, Kayangel, etc. (NSF, 1955); Yap Id. (C. O. Kile, ANSP); CAROLINES: Ulithi (ANSP); Oneop Id., Lukunor Atoll (MCZ). MARSHALLS: Bikini, Eniwetok, Rongelap and Rongerik Atolls (all USNM). FIJI: Mbau Id. and Suva ( H. S. Ladd, USNM); Levuka, Ovalau Id. (R. T. Abbott, MCZ); Ongea Levu, Lau Group (H. S. Ladd, USNM). TONGA: Vlitoa ( BPBM no. 68528); Vava’u (ANSP). Fossil records — KENYA: Pleistocene; reef limestone, Mom- basa (L. R. Cox, 1930, p. 139). TANGANYIKA: Pleisto- cene of Dar-es-Salaam (A. Ortmann, 1892, p. 642). SU- DAN : Port Sudan, coral reef. Pleistocene ( Hall and Stan- den, 1907, p. 67). INDONESIA: Post-Tertiary Pleistocene, Billiton [Belitong Id.], Borneo. ( K. Martin, 1881, vol. 3, p. 17). Young Quaternary at Goenoeng Medong, east Borneo Id. (Beets, 1948, p. 8). FIJI ISLANDS: Lau Id., Futuna limestone, Pliocene (?) , Station L 493 ( specimen resembling L. lambis seen in USNM). NEW HEBRIDES: Espirito Santo Island, Pliocene (?), H. T. Stearns Station ES— 14, alt. 240 ft. (specimen of L. lambis seen in USNM). Lambis tnincata (Humphrey, 1786) This is the largest and one of the most widely distributed members of the genus in the Indo-Pa- cifie. Adults reach a length of 10 to 15 inches. It is distinguished from the smaller L. lambis by the large knobs on the spire and by the absence of the strong, raised, axially-pinched knob which is pres- ent on the dorsum of female lambis. The dark pur- ple-brown maculations found in some lambis are absent in truncate. This species may be readily separated into two subspecies: truncata tnincata (Humphrey, 1786) — with a flat apex, thus giving the spire a truncate appear- ance. Limited to the Indian Ocean. truncata sebae (Kiener, 1843) — with a pointed apex, thus giving the spire an acute angle. With a curious, discontinuous distribution in the Red Sea and the Pacific Ocean as far east as the Tuamotu Islands. We cannot on the basis of the few authentic specimens we have seen distinguish Red Sea and Western Pacific specimens of sebae. The dorsal hump or shoulder knob in Red Sea specimens is much more pronounced than that of Pacific speci- mens. This is a not uncommon feature of some of the truncata truncata from Zanzibar. L. t. sebae occurs also in Pleistocene deposits in the Red Sea area. It is possible that the subspecies truncata has more recently developed in the Indian Ocean and has now isolated the extreme ends of the parent subsjoecies, sebae. Some, but not all, specimens from eastern Poly- nesia take on an elongate shape with the spire angle being less than that of East Indian and Mela- nesian specimens. The interior of the aperture, at the upper end, may have a few axial folds or creases. This is what Morch named sowerbyi which we are synonymizing with sebae. The chocolate coloration of the aperture mentioned by Morch occurs in only a few specimens, and may be only due to environmental conditions. At best, it is a very weak subspecies. Lambis truncata subspecies truncata (Humphrey, 1786) (PI. 122, fig. 2) Range — Indian Ocean from central East Africa to the Ray of Bengal and Cocos Keeling Atoll. Remarks — This subspecies is characterized by its truncate, flat apex which is brought about by the first 5 whorls revolving in the same plane. The sixth and subsequent whorls descend at a normal [10-060] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 155 rate, so that the adult shell takes on a normal Lam- bis shape. The young shell is so strikingly different from the adult that Grateloup described it as a new species of “Pyrula” in 1840. There is little differ- ence in the adult shells of sebae and truncata, ex- cept that the latter generally has a rather large, indistinct hump on the dorsum, grows to a slightly larger size and usually has one or two fewer knobs per whorl in the spire. This species is moderately common throughout its Indian Ocean range. Recently matured specimens have six long, slightly arching labial digitations. Rarely, a small, aberrant seventh may occur between the first and second digitations. Old specimens, usually found in shal- low water on reefs and in lagoons, are very heavy and encrusted with calcareous algae, vermetid snails and rarely coral. The digitations and siphonal canal are greatly reduced, the aperture constricted and the parietal wall and outer lip covered with a creamy brown, granular glaze. Habitat — Occurs near reefs in 12 to 15 feet of water (Issel and T.-Canefri, 1876, p. 356). I have seen natives dive for this species in 20 feet of water off the reefs of Kenya. It is rarely seen on the reefs, mainly, I believe, because the natives remove most of the larger shells for food purposes at each low tide. Description — Shell massive and, including the digitations, 9 to 15 inches in length, with 6 slightly arching labial digitations. Siphonal canal propor- tionately short and almost straight. Whorls 9 to 10. Apex with about 4 to 5 flat whorls. Shoulder of whorls in spire with rather large, rounded nodules, usually 10 to 12 per whorl, but becoming obsolete in the last whorl. Dorsum of body whorl with 2 or 3 knob-like swellings, one of which may be large and mound-like. Spiral sculpture of numerous, weak, irregularly-sized threads or cords. Aperture and parietal area with an enamel gloss. The glaze at the upper end of the columella extends over the nodules on the spire and sometimes over the apex. Color of aperture white within and becoming tan or purplish brown near the edges. Outer lip wavy in recently matured specimens, but very thick and smooth in old specimens. Columella straight and smooth. “Stromboid notch” bounded anteriorly by a small, pointed spur. Color of outer shell whitish cream with sparse, light-brown specklings or some- times with chestnut-brown spiral bands and macu- lations. Periostracum moderately developed, trans- lucent-tan and flaking off when dry. Operculum chitinous, elongate, very slightly curved, brown, and with smoothish edges. The radulae are propor- tionately massive and with about 55 transverse rows. The center cusp of the central tooth is broadly rounded. Formula: 2-1-2; 1-3 or 1-4; 5; 6 to 8. Measurements (mm.) — (including digitations) length width no. whorls 375 212 10 + ( large; Zanzibar ) 387 225 8 + ( large; Seychelles 250 150 8 + ( small; Zanzibar ) Synonymy — 1786 Strombus truncatus Humphrey, Portland Catalogue, p. 133, no. 2967 (East Indies); refers to Davila, vol. 1, pi. 12, fig. 14; ibid., p. 150, no. 3307 (young); ibid., p. 169, no. 3655 (China); 1817, Dillwyn, Descr. Cat., vol. 2, p. 659. 1791 Strombus bnjonia Gmelin, Systema Naturae, 13th ed., p. 3520, No. 33 (no locality). Refers to Lister, pi. 882, fig. 4 and others; 1825, Wood, Index Testace- ologicus, pi. 24, fig. 8. 1798 Lambis davilae Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 66. Refers to Davila, vol. 1, pi. 14, pi. 13. 1798 Lambis radix Roding, loc. cit., p. 67, Refers to Con- chy].-Cab., vol. 10, fig. 1514. 1840 Pyrula bengalina Grateloup, Actes Soc. Linn. Bor- deaux, vol. 11, pp. 170 and 448, pi. 4, fig. 5 (Ben- gal ) - 1852 Pterocera (Heptadactylus) radix-bryoniae Ch., Morch, Catalogus Conchyl. . . . Yoldi, Hafniae, p. 60. [10-061] 156 Lambis R. T. Abbott Strombidae 1854 Harpago (Heptadactylus) radix-bryoniae Gmel., H. and A. Adams, Genera of Recent Mollusca, London, vol. 1, p. 261. 1822 Pterocera tmncata Lamarck, Anim. sans Vert., vol. 7, p. 195; 1834, Kiener, Coquilles Vivantes, vol. 4, pi. 1, pi. 10, fig. 5. 1872 Pterocera (Heptadactylus) tmncata Humphrey, Morch, Jour, de Conchyl., vol. 20, p. 132. 1955 Lambis (Lambis) tmncata (Humphrey), Butot, Peng- gemar Alam, vol. 35, p. 76 (in part). Types — The whereabouts of Humphrey’s type is unknown to me. It was probably sold at auction in 1786 in London. N omenclature — The eighteenth century figures of tmncata were a mixture between the flat-topped Indian Ocean and acute-spired Red Sea and Pacific Ocean subspecies. Humphrey’s 1786 name of trun- cata refers to Davila, pi. 12, fig. 14, a fiat-topped specimen probably from East Africa. The earliest valid name for the acute-spired subspecies appears to be that of sebae (Kiener, 1843). Grateloup’s Pyrula bengalina is a very young specimen of tmncata tmncata, a fact which was pointed out by Kiener in 1843. Although Butot (1955, p. 76) seemed to have understood the characters of L. tmncata which has a truncate spire, he confused the issue by referring his own acute-spired specimen ( his plate 2 ) from Kangean Island, Indonesia, and that of Rumphius (“Cornuta decumana” Rumphius 1705, pi. 35, fig. H) to tmncata, rather than to sebae (Kiener). Records (also see accompanying map) — KENYA: Diani Beach, south of Mombasa ( R. T. Abbott, USNM, MCZ). MADAGASCAR: Nossi Iranja, south side of Nossi-be ( Acad- emy-Madagascar Exped., I960); Nossi Fanihi; Tulear; Am- bodifotatra; Tamatave (all Dautzenberg, 1929, p. 472). SEYCHELLES: Southeast Id., off Mahe Id. (Yale Peabody Mus.). MAURITIUS: Port Louis (MCZ). CEYLON: ( Issel and T.-Canefri, 1876, p. 356). ANDAMANS: Port Blair (Issel and T.-Canefri, 1876, p. 356). BENGAL: ( Grate- loup, 1840, pi. 4). COCOS KEELING ATOLL: shallow water in lagoon (USNM). Fossil Records — None recorded. Lambis truncata subspecies sebae (Kiener, 1843) (PI. 121, fig. 9; pi. 122, fig. 1) Range — Red Sea and the tropical Pacific Ocean from the East Indies to eastern Polynesia. Remarks — This subspecies differs from the typi- cal tmncata tmncata of the Indian Ocean in having a pointed, rather than a truncate, apex. Adults rarely exceed 13 inches in length (including spines), while tmncata may reach 16 inches. The curious distribution of this subspecies in two distantly sepa- rated areas, one in the Red Sea, one in the Pacific, can be explained by at least two possibilities. Either, two morphologically similar forms have evolved in- dependently of one another, or the once continuous distribution has recently been broken by the inva- sion of a geographically intervening race of flat- topped Indian Ocean specimens. The fossil record is too poorly known to settle this matter. The flesh of this Lambis is commonly eaten by natives. Hedley (1899, p. 429) reports that the Ellice Islanders eat the snail raw or roasted. Habitat — Lives in colonies on sandy, algal and coral rubble bottoms in the vicinity of coral reefs. “Commonly found at depths of 15 to 30 feet off the edge of seaward reefs . . . and on sandy lagoon shelves among seaweed or on minor reef promi- nences in 2 to 10 feet of water. Invariably occurs below low tide line [Marshall Islands]” (Demond, 1957, p. 297). Old and worn, but living, specimens occur on gravel flats in the lagoon at Funafuti in waist deep water (Hedley, 1897, p. 429, fig. 18). “It is rare in the central Philippine Islands” (E. Zambo, in litt., 1958). Description — Shell massive, 9 to 13 inches in length, and differing from the Indian Ocean tmn- cata tmncata in having the apex of the shell with a spire angle of 80 to 100 degrees, and in rarely exceeding a total length of 12 inches. Young speci- mens show a predominance of fine axial streaks of light-brown. The radulae are illustrated on plate 119, fig. 4. Measurements (mm.) — (including digitations) length width no. whorls 300 200 9 + ( large; Samoa ) 275 170 8 + ( average; Saipan Id. ) 225 137 8 + (small; Mindanao Id.) Synonymy — 1843 Pterocera sebae “Valenciennes” Kiener, Coquilles Vi- vantes, Paris, vol. 4, Pterocere, p. 4, pi. 2, pi. 4, fig. 2 ( la mer Rouge et l’ocean Indien ) . 1872 Pterocera (Heptadactijlus) sebae Valenciennes, Morch, Jour, de Conchyl., vol. 20, p. 131. 1872 Pterocera (Heptadactylus) sowerbyi Morch, Jour, de Conchyl., vol. 20, p. 131 (Ins. Taiti). 1935 Lambis bryonia Gmelin, Y. Hirase and Pilsbry, 1000 shells in Color (Kai Sen Shu), vol. 4, pi. 73, fig. 368. 1938 Lambis truncata (Solander), S. Hirase, A Collection of Japanese Shells, 6th ed., pi. 87, fig. 2. 1955 Lambis sebae Valenciennes (Kiener), Butot, Peng- gemar Alam, vol. 35, p. 76, pi. 1, fig. 2. 1955 Lambis truncata (Humphrey, 1786), Butot, loc. cit., pi. 1, fig. 1, 3, pi. 2, fig. 7. 1958 Lambis (Lambis) truncata (Humphrey), Oyama, The Molluscan Shells, Tokyo, vol. 2, Lambis (1), figs. 7, 8 ( Okinawa). Types — We hereby restrict the type locality of sebae Kiener to the Red Sea. The type is presuma- bly in the Musee d’Histoire Naturelle de Geneve. [10-062] September 28, 1961 INDO-PACIFIC MOLLUSCA, vo). 1, no. 3 Lambis 157 RED SEA records: Gase Arm, Israel (ANSP); Aqaba, Israel ( Issel and T.-Canelri, 1870, p. 356). Sherm Sheikh; Shadwan Id.; Ras Abu Sorner; Hassani Id., and Jidda ( all R. Sturany, 1903, p. 28). Port Berenice, Egypt (ANSP). 10 mi. oft Port Sudan ( R. C. Spencer, coll n.). Berbera, Gulf of Aden ( USNM ) . PACIFIC OCEAN records (also see accompanying map) -RYUKYU ISLANDS: (USNM). PHILIPPINES: Iba, Zambales, Luzon Id. (P. de Mesa, ANSP). Lubang Id., Min- doro Prov. ( P. de Mesa, ANSP). Cuyo Id., Palawan Prov.; Zamboanga, Mindanao Id. (both du Pont-Academy Exped., 1958) . Sibuyan Id., Romblon Prov. (Alcasid, 1947, p. 180). Sarangani Bay, S. Mindanao Id.; Tara Id. (both USNM). INDONESIA: Amboina ( Rumphius, 1705, pi. 35, fig. H). Borneo and Ceram Ids.; Kangean Id., N. E. of Java (all Butot, 1955, p. 76). Waigeu Id., west of Dutch New Guinea (Butot, 1955, p. 76). AUSTRALIA: Torres Straits (ANSP); Keeper Reef, 40 mi. from Townsville, Queensland (Tony Marsh, in litt.). NEW HEBRIDES: Erromanga ( Hedley, 1899, p. 429). NEW CALEDONIA: barrier reef, Touho Bay; Plage de Poe, Bourail (both G. and M. Kline, NSF, 1959) . FIJI ISLANDS: Namuka Id., Kaloka Levu, Viti Levu Id. ( R. T. Abbott, MCZ). Suva, Viti Levu Id. ( H. S. Ladd, USNM). MARIANAS: barrier reef, Tanapag, Saipan Id. ( R. Sutcliffe, ANSP). Tinian Id. (J. L. Chamberlin, MCZ). Guam Id. (USNM). Maug Id. (USNM). CARO- LINES: 4 fms., Main Pass, Ifaluk Atoll (USNM). MAR- SHALL ISLANDS: Bikini, Eniwetok, Rongelap Atolls (J. P. E. Morrison, USNM). GILBERT ISLANDS: Onotoa Atoll (P. E. Cloud, USNM); Apiang (A. Garrett, MCZ). HA- WAIIAN CHAIN: [Evidently absent. Spicer, 1941, P- 2, re- ports that Guam specimens were taken to and sold on Mid- way]. LINE ISLANDS: Christmas Id. (Fred Barnett, ANSP). ELLICE ISLANDS: Funafuti (Hedley, 1899, p. 429). SAMOA: Satalo Id., Upolu Id. (NSF, 1955). Fagaitua Bay, Tutuila Id. (MCZ). SOCIETY ISLANDS: District of Anau, Bora Bora; Aua, District of Paea, and Ative, District of Punaauia, Tahiti (all R. Robertson, 1952, ANSP). Nauarei, Moorea Id. ( H. A. Rehder. USNM). TUAMOTU ISLANDS: Ship Pass, Raroia (IT Rufon, ANSP). Hao and Ohura [= possibly Takapoto] (Couturier. 1907, p. 154). Maran- ganai Id., Toau [Elizabeth Atoll] ( H. A. Pilsbry, 1929, ANSP). Fossil records — Pleistocene of Khor Ambado, French So- malia ( Abrard, 1942, p. 62, pi. 6, fig. 34, as Pterocera cf. millepeda Linne). HAWAIIAN CHAIN: “Pleistocene” raised beach, 15 feet altitude, Makua, south coast of Oahu Id. ( Children’s Museum, Honolulu ) . Lambis crocata subspecies crocata (Link, 1807) (PI. 121, fig. 8, pi. 126) Range — East Africa to Samoa and the Ryukyu Islands to northern Australia. Remarks — This moderately common species is readily recognized by its solid-orange, smooth aper- ture. The outer lip bears 6 slender digitations, and the siphonal digitation is long and gracefully curved. We have divided the species into two races— the typical crocata of the Indian and western Pacific Oceans which becomes progressively rare towards the east where it aj^pears to have reached its limits in Samoa— and the giant subspecies pilsbryi which appears to be limited to the Marquesas Islands in eastern Polynesia. The exterior of the shell is usually mottled in creamy-white and soft brownish orange, although a pure-orange color form rarely occurs in colonies Plate 126. Immature shells of Lambis crocata ( Kayangel Island, Palau Islands). Fig. 1, with digitations beginning to be formed. 2, with digitations almost completed. Both nat- ural size. from the Palau Island and the Philippines. A speci- men from Chumbe Island, Zanzibar, is malformed and has 9 labial digitations. This species has not been recorded from the Red Sea or the Hawaiian Chain. It was formerly called aurantia Lamarck. Habitat — This species lives on seaward reefs from the low tide mark to a dej)th of about 10 feet. Melvill (1909, p. 94) reported it at depths of 31 and 34 fathoms in the Seychelles, but did not indi- cate whether or not these were live specimens. Alan J. Kohn recovered live specimens on the reefs in the Seychelles (Yale Peabody Exped., station 41, 1957). Description — Shell 100 to 150 mm. ( 4 to 5M inches) in length (including the digitations), with 6 labial digitations, and a long, slender, curved siphonal digitation. Aperture a solid, whitish orange and smooth. Nuclear whorls 3, smooth, glossy and light-brown in color. 7 to 8 postnuclear whorls with numerous spiral threads. The shoulder of the whorls in the sjhre bears a small beaded carina which lies just above the finely indented suture. Last whorl with 3 spiral, rows of knobs, the upper row having 2 to 4 widely-separated rather large, rounded knobs and the lower two rows having 4 to 6 smaller knobs. Color of outer shell solid orange-tan or whitish with orange-tan maculations. Lower 3 labial digi- tations hooked posteriorly at an 80 to 90 degree angle. Aperture elongate, solid orange, smooth, constricted within by a long axial ridge on the in- side of the body whorl. UjDper end of aperture with [10 - 063] 158 Lambis R. T. Abbott Strombidae Plate 127. Geographical distribution of Lambis crocata cro- cata ( Link ) and its Marquesan subspecies pilsbryi Abbott. a deep well which is bounded by a small spiral ridge on the upper end of the columella. Columel- lar callus enameled, swollen, and with a sinuate line of demarkation on the left side. “Stromboid notch” large and fluted. Periostracum heavy, brown and slightly rough, but usually worn off the body whorl. Operculum slightly curved, brown and its edges smoothish. The verge is slender, with a small distal pad, maculated with cream spots, and about h the length of the aperture. The odontopliore is about 8 mm. in length with 40 transverse rows of teeth. The radular formula is 2-1-2; 1-2; 5; 5. Measurements (mm.) — (including digitations) length width no. whorls 150 73 8 + (large; Tutuila Id., Samoa) 128 55 8 + ( average; Cebu Id., Philippines 103 45 5 + ( small; Gulf of Manaar, India Synonymy — 1798 Lambis scorpius Gmelin, Roding, Museum Boltenianum, Hamburg, pt. 2, p. 67, no. 60. Refers to Conchyl.- Cab., vol. 10, figs. 1508-09. Non Gnrelin, 1791; non Linne, 1758. 1807 Pteroceras crocatus Link, Beschr. Naturalien-Samm- lung, Rostock, pt. 2, pp. 109, 110. Refers to Conchyl.- Cab., vol. 10, figs. 1508-1509. 1811 S trombus aculeatus Perry, Conchology, London, pi. 13, fig. 2 (no locality). Perhaps a nomen dubium. 1822 Pterocera aurantia Lamarck, Anim. sans Vert., vol. 7, p. 198 ( Indes orientales). Refers to Conchyl.-Cab., vol. 10, figs. 1508, 1509 and Knorr, pt. 5, pi. 4, fig. 3. 1825 Strombns lambis L., Wood, Index Testaceologicus, London, p. 116, pi. 24, fig. 7. 1825 Pteroceras aurantiacum Sowerby, Catalogue Shells . . . Tankerville, London, p. 67 (substitute name for aurantia Lam.). 1842 Pteroceras aurantia Lam., Sowerby, Thesaurus Con- chyl., vol. 1, Pteroceras, p. 42, pi. 11, fig- 11. 1941 Lambis crocata (Link), Kuroda, Mem. Faculty Sci. Agriculture, Taihoku Imperial Llniv., vol. 22, no. 4, p. 98; 1955, Butot, Penggemar Alam, vol. 35, p. 77, pi. 2, fig. 3. Types — Link’s species is based upon figures 1508 and 1509 of the Conchylien-Cabinet, vol. 10. That specimen is said to come from the East Indies. We hereby restrict the type locality to Amboina, Indo- nesia. Perry’s type of aculeatus probably fell into private hands, and may still be in existence, but probably without identifying labels. Locality records — KENYA: Shimoni, Mombasa (Yale Pea- body Mus.); Diani Beach ( R. T. Abbott, USNM, MCZ). ZANZIBAR: Chumbe Id.; Pange Id.; outer reef, Kiwengwa; lias Nungwe ( NSF, 1957). MADAGASCAR: Nossi-be (A. Chavane, ANSP). INDIAN OCEAN ISLANDS: Gloriosa Id. (USNM); Providence Id., Mascarenes (E. A. Smith, 1884, "Alert”, p. 503); Seychelles: Menai Id., Cosmoledo Atoll; He Platte (both A. J. Kohn, Yale Peabody Mus.). Chagos Islands: ( Lienard, 1877, p. 95). Mauritius ( N. Pike, MCZ). CEYLON: Pearl Bank, Gulf of Manaar (G. and M. Kline, ANSP). INDIA: Pamban and Tuticorn (Thurston, 1895, p. 125). RYUKYU ISLANDS: Okinawa Id. (Mrs. A. A. Scott, ANSP). Amami Islands (Kira, 1959, p. 37). TAIWAN: Ryukyo-syo (Kuroda, 1941, p. 98). PHILIP- PINES: Olango Id., east Cebu ( du Pont-Academy Exped., 1958); Panglao, Bohol Id. (E. Zambo, ANSP); Surigao, Mindanao Id. (ANSP); Jolo Id., Sulu Arch. (ANSP). IN- DONESIA: Amboina (MCZ); Java Id.; Madura Id.; Ambon Id.; and Ceram Id. (Butot, 1955, p. 77). AUSTRALIA: Undine Reef, Queensland (Tony Marsh, in litt.). DUTCH NEW GUINEA: Mios Woendi Atoll, Padaido Ids. (NSF. 1956). PALAUS: Nguarangel (USNM); Kayangel Id. (NSF, 1955). MARSHALLS: Lomuilal Id., Rongelap Atoll: Taka Atoll (both USNM). FIJI ISLANDS: Namuka Id., off Kaloka Levu, Viti Levu Id. (R. T. Abbott, MCZ). SAMOA: Fagaitua Bay, Tutuila Id. (R. T. Abbott, MCZ). Fossil records — Pliocene, south end of Mombasa Id., Kenya (J. Weir, 1938, p. 69, pi. 5, fig. 3). Lambis crocata subspecies pilsbryi, new subspecies (PI. 128, figs. 1, 2) Range — Known only from the Marquesas Islands, Polynesia. Remarks and Description — This giant, isolated race is presumed to be limited to the Marquesas Islands. I have seen only six specimens. One of these was given to Dr. H. A. Pilsbry by Pere Simeon Delmas of the Marquesas. The shells of this sub- species differ from those of crocata crocata in being twice as large (7 to 9 inches in length, including digitations), in having the 3rd, 4th and 5th digita- [ 10 - 064] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 159 Plate 128. Lambis crocata new subspecies pilsbryi. Mar- quesas Islands. Fig. 1, holotype, ANSP no. 156123. 2, para- type, ANSP 255621. % natural size. tions straight or only slightly curved, and in lacking the fairly prominent edge on the left of tire parietal callus. In pilsbryi, the dorsal shoulder knob is much larger, the apex of the spire is usually buried by the first digitation, and the small, columellar ridge at the posterior end of the aperture is very weak. In other characters, such as the color of the outer shell and aperture and in the sculpturing of the spire, it is similar to crocata. Operculum and soft parts unknown. Two names have been proposed for crocata which might possibly refer to this subspecies, although one of them, Lambis yolclii (Morch, 1852) is inade- quately described, and the other, Lambis aculeata (Perry, 1811 ), is based upon a distorted illustration and too brief a description for certain identifica- tion. We consider them nomena (labia. We name this new subspecies in honor of Dr. Henry A. Pils- bry, former Curator of the Department of Mollusks at the Academy of Natural Sciences of Philadelphia. Measurements (mm.) — (including digitations) length width no. whorls 240 113 7 + ( paratype, MCZ no. 49694 ) 212 + 95 7 + (holotype, ANSP no. 156123) 182 + 90 8 (paratype, ANSP no. 255621 J Synonymy — 1842 Pteroceras lambi.s Lin., Sowerby, (in part). Thesaurus Conchyl., vol. 1, Pteroceras, p. 41, pi. 11, fig. 6 only. Not lambis (Linne). Types and Locality Records — The type locality is Nuku Hiva in the Marquesas Islands. The holo- type is in ANSP no. 156123 and was collected by Pere Simeon Delmas. Most of Delmas’ shells were collected on the coral banks in Tai O Hae Bay. One paratype is in MCZ no. 49694. One paratype in B. P. Bishop Mus. no. 68268 from Uahuka, Mar- quesas (S. Delmas, 1922). Miss Martha E. Hunt of Baltimore kindly loaned me a specimen which T. Gavaldon collected in the Marquesas in 1959. [10-065] 160 La mb is R. T. Abbott Strombidae [These occasioned blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [10 - 066] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 161 Subgenus Millepes Moreh, 1852 Type: Lambis millepeda Linne The shells in this group of Lambis are character- ized by elongate apertures bearing well-developed spiral lirae, by a siphonal canal which is either straight or curved to the right, and by the presence of 6 to 10 labial digitations. There are five species, three of which are quite limited in distribution, the other two being found in both the Indian and Southwest Pacific Oceans. Only one, Lambis scor- pius Linne, shows any subspsciation. Below is a synopsis of the differentiating charac- ters of the five living species. Lambis scorpius scorpius ( Linne ) — ( Southwest Pacific). Six labial digitations nodulose, the lower three strongly hooked. Siphonal canal long, strongly curved to the right. Deep in the throat is a white bar or ridge just below the concavity caused by the dorsal tubercle. An ear-like lobe present at the left side of the base of the first digitation. Lambis scorpius indomaris Abbott — ( Indian Ocean). Differing from the above in lacking a well-developed lobe at the base of the first digitation, and in having stunted, shorter 4th, 5th and 6th digitations. Lambis robusta (Swainson) — (Southeastern Poly- nesia). Six axial digitations almost smooth; the 3rd and 4th close together; siphonal canal al- most straight. No white ridge deep in throat. Lambis millepeda ( Linne ) — ( Western Pacific Arc). Nine labial digitations; aperture mauve- brown. Lower parietal callus weakly lirate. Spire low. Plate 129. Anertural details of members of the subgenus Millepes Moreh. Fig. 1, Lambis violacea (Swainson). 2, L. digitata (Perry). 3, L. millepeda (Linne). 4, L. scorpius Lambis digitata (Perry) — (Indo-Pacific). 8 to 9 labial digitations, the first usually being bifur- cate. Spire elongate. Parietal wall callus raised and with prominent, wavy, whitish lirae. Lower 6 digitations very short. Edge of outer lip broad, covered with periostracum. Lambis violacea ( Swainson ) — ( Indian Ocean). 9 to 11 labial digitations, the first usually being bifurcate, and all being enamel-white and flat- tish on the underside. Spire relatively short. Aperture whitish with yellowish bars on the outer lip, and the throat violet to lavender. Apertural lirae fine and numerous. There are no species limited to the Tertiary, but millepeda and scorpius have been recorded from the Quaternary of Indonesia. Synonymy — 1852 Millepes “Klein”, Moreh, Cat. Conchyl. . . . Yoldi, Hafniae, p. 60; 1859, Chenu, Manuel de Conchyl. et Paleont. Conchyl., Paris, vol. 1, p. 259; 1940, VVenz, Handbuch der Palaozool., vol. 6, pt. 4, p. 948 (type by subsequent designation: millepeda Linne). 1854 Millipes “Klein”, H. and A. Adams, Genera of Recent Mollusca, London, p. 261; 1868, Gabb, American Jour. Conch., vol. 4, no. 3, p. 140; Tryon, Manual of Conch., Philadelphia, vol. 7, p. 125; 1878, Kobelt, Illustrirtes Conchlienbuch, vol. 1, p. 106 (type by subsequent designation: Scorpio Linne). 1870 Millipes Moreh, Gill, American Jour. Conch., vol. 5, no. 3, p. 128, 134; 1955, Butot, Penggemar Alam, vol. 35, p. 74. Lambis millepeda (Linne, 1758) (PI. 121, fig. 6; pi. 129, fig. 3) Range — Southwest Pacific. Remarks — This species is rather common wher- ever it occurs, but its distribution is limited to the central portion of the Western Pacific Arc from the Philippines to New Guinea. It is recognized by its 9 labial digitations which arise just behind the edge scorpius (Linne). 5, L. robusta (Swainson). All about ¥2 natural size. [10-071] 162 Millepes R. T. Abbott S trombidae of the outer lip and by the brownish mauve aper- ture which bears whitish mauve lirae well into the throat of the aperture. The lower 3 or 4 labial digi- tations are turned or hooked towards the posterior or spire end. Mr. Evaristo Zambo of Cebu, Philippines, in- forms me that the flesh of millepede is bitter and generally not eaten by natives. No authentic rec- ords of this species are known from the Red Sea or Indian Ocean. An account of the gross anatomy was published by R. Bergh ( 1895, pp. 364-368, pi. 23). Habitat — Lives in shallow water down to a depth of 2 fathoms. Description — Shell 90 to 145 mm. ( 3 M to 5b inches) in length including the digitations. Aper- ture mauve-brown with numerous white lirae. Outer lip with 9 short labial digitations of which the lower 4 or 5 are hooked towards the apex. Siphonal canal rather short and twisted. Nuclear whorls 3, smooth, translucent-tan. Postnuclear whorls 8. Early whorls with a beaded carina; later whorls with 9 to 11 nodules. Last whorl with 3 spiral rows of nodules, the top row with 4 large, somewhat elongate knobs, the lower two rows with 3 or 4 small, round nod- ules. Color of outer shell cream with a heavy suf- fusion or network of dark-brown. Edge of outer lip crenate. Aperture mauve-brown with numerous, wavy, weak lirae of a whitish purple color. Throat yellow within and bounded by an axial whitish ridge on the inside of the body wall. Upper end of aperture has a deep, round, yellow depression or well which is bounded above by two facing, tongue- like whitish ridges. Columella swollen below; bears about 40 to 50 irregular, spiral, mauve-white lirae which cross the blackish to chocolate-brown parie- tal wall. “Stromboid notch” with 2 or 3 short, flat projections at its edge. Periostracum thin, glossy and translucent-brown. Operculum long, dark- brown and with smoothish edges. Bergh’s (1895) figures show a radula similar to that of scorpius and with a formula of 2-1-2; 1-4; 6; 6. Measurements (mm.) — (including digitations) length width no. whorls 145 62 8 + ( large; Luzon Ick ) 100 53 11 ( average; Cebu Ick ) 95 48 9 + ( small; Schouten Ids. ) Synonymy — 1758 Strombus millepeda Linne, Systema naturae, ed. 10, p. 743, no. 426 (In O. Asiae); 1764, Mus. Ludovicae Ulricae, p. 618; 1767; 12 ed., p. 1208, no. 494; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill pt. 3, pp. 251-252; 1801, Bose, Hist. Nat. des Co- quilles, Paris, vol. 4, p. 249. 1798 Lambis millepeda Gmelin, Roding, Museum Bolteni- anum, Hamburg, pt. 2, p. 67, no. 855 (refers to Conchyl.-Cab., vol. 3, figs. 861-862). 1807 Pteroceras millepeda Linne, Link, Beschr. Natur.- Samml., Rostock, pt. 2, p. 110 (refers to Conchyk - Cab., vol. 3, figs. 861-862); 1842, Sowerby, The- saurus Conchyk, vol. 1, p. 43, pi. 11, fig- 3 (Philip- pines ) . 1807 Pterocera millepeda Linne, G. Fischer, Museum-Demi- doff, Moscow, vol. 3, p. 191 (refers to Conchyl.- Cab., vol. 3, figs. 861-862); 1839, Anton, Verz. Conchyk, Halle, p. 84; 1843, Kiener, Coquilles Vi- vantes, Paris, vol. 4, p. 10, pk 9, figs. 1, 2 (not pi. 10, fig. 1); 1851, Reeve, Conch. Icon., vol. 6, pk 6, fig. 10. 1870 Pterocera millipeda Lam. ex Linn., Gill, American Jour. Conch., vol. 5, no. 3, p. 135. 1955 Lambis (Millipes) millepeda (Linne, 1758), Butot, Penggemar Alam, vol. 35, p. 77, pk 2, fig. 4 (Ma- dura, Indonesia). Types — “The Linnaean collection in London con- tains a properly marked specimen of millepeda which thus may be accepted as the type of the spe- cies.” (Dodge, 1956.) We hereby designate Olango Island on the east side of Cebu Island, Philippines, as the type locality. Nomenclature — Linnaeus, Gmelin (1791), Dill- wyn (1817) and Lamarck (1822) considered what was later described as violacea ( Swainson ) and digitate (Perry) to be synonyms or varieties of millepeda (Linne). Swainson in 1821 was the first to accept them as distinct species. Sowerby (1842) and, later, Deshayes (1843) also separated them but gave names which are now considered syno- nyms. Curiously, Dodge (1956, pp. 251-252) con- fused the situation by believing that elongata (Swainson) and digit at a (Perry) were different species, and that the latter was the same as multipes of Chemnitz and of Deshayes. Actually, multipes is a synonym of violacea (Swainson, 1821). Alata polydactylus Martini 1777 is a non-binomial syno- nym of millepeda Linne. Records — ( See accompanying map, pk 130 ) PHILIP- PINES: Calapan, Mindoro (P. de Mesa, MCZ). Luzon Id.: Iba, Zambales (P. de Mesa, ANSP); Tabaco, Albav Prov. (du Pont-Academy Exped., 1958, ANSP). Cebu Id.: Olango (ANSP). Bohol Ick: east side of Jagoliao Id. (both du Pont- Academy Exped., 1958). Mindanao Id.: Davao Bay (MCZ). Masbate Id.; Basilan Id.; Marongas Id., Jolo ids. (all USNM). Romblon Prov., Sibuyan ( Alcasid. 1947, p. 184). INDONESIA: Amboina (MCZ); Batjan Id. (MCZ); Ma- dura Id., Java; Ternate; Ceram; Timor (all Butot, 1955, p. 77). DUTCH NEW GUINEA: Soepiori Id., Schouten Ids.; Japen Id. and Biak Ick, and Aoeri Ids. (all NSF, 1956). AUSTRALIA: Cotton (1953, no. 3, second page, fig. 17) reports it from Northern Australia and from Queensland, although J. Allan (1959, p. 102) states this is not so. Fossil records — INDONESIA: Pleistocene-Pliocene: Koe- pang, Timor Id. (Tesch, 1920, p. 52, pk 130, fig. 172). [Abrard’s 1942, vol. 18, p. 62, pk 6, fig. 34 of millepeda from the Pleistocene of the Red Sea looks like a Lambis truncata sebae ]. [R. B. Newton’s 1900, p. 509, Pleistocene record for Gemsah, Egypt needs confirmation, and may well be L. digit at a]. [10- 072] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 163 Lambis digitata (Perry, 1811) (PI. 121, fig. 3; pi. 129, fig. 2) Range — East Africa to Samoa. Remarks — This is not a common species, but it has a rather wide distribution. It was formerly known as elongata (Swainson). It is characterized by a proportionately high spire, by 8 or 9 labial digitations of which the lower 6 or 7 are very small and the first two, at the upper end of the aperture, are longer and widely separated. The first digita- tion is usually bifurcate. The edge of the outer lip is broad and thickly covered with brown, rough periostracum. The interior of the aperture is similar to that of millepeda, but differs in being less con- stricted and in having a short, spiral, whitish ridge just below the concavity (caused by the last dorsal knob on the outside of the shell). The raised, lirate parietal callus in digitata has a sharp, delimiting left edge. I have seen only two specimens from the Indian Ocean. They have one extra digitation on the outer lip, and other differences, which suggest that there may be an Indian Ocean subspecies, but more specimens are needed to prove this point. Reeve’s crocea looks like the Indian Ocean form and his name is available for it. Habitat— Unknown, although it is probably asso- ciated with coral reefs at depths from 1 to 3 fathoms. Description — Shell 98 to 145 mm. (4 to 5h inches) in length, with 8 (rarely 9) labial digita- tions of which the anterior or first one is bifurcate and the lower 5 or 6 are very short. Spire very high and with an angle of about 40 degrees. Nuclear whorls unknown. Postnuclear whorls 10. Shoulder of whorls carinate and bearing numerous, small, sharp nodules. Body whorl with 3 rows of nodules, the top row with the 5 largest, and the two rows below bearing 4 to 6 small nodules. Color of outer shell whitish with yellow-brown speeklings and mottlings. Outer lip thick. Aperture purplish mauve with numerous whitish spiral lirae. Throat yellow- ish white within and with a deep depression at the posterior end which is bounded anteriorly by a strong whitish ridge. Columella with a thickened mauve callus which bears about 25 whitish, bifur- cating, spiral lirae. This parietal or columellar cal- lus is sharply bordered on its left side. Inside of outer lip purplish with numerous purplish white spiral lirae. “Stromboid notch” well-developed and with the thick outer lip slightly overhanging its upper edge. Siphonal canal rather short and twisted. Periostracum brown and thicker on the edge of the outer lip. Operculum and soft parts unknown. Measurements (mm.) — (including digitations) length width no. whorls 140 70 10 + ( large; Pacific Ocean” ) 132 53 9 + ( average ; “Indian Ocean 118 53 10 + ( small; Pacific Ocean” ) Synonymy — 1811 Strombus digitatus Perry, Conchology, London, pi. 13, fig. 1 ( Eastern Ocean ) . 1816 Pterocera millepeda Lamarck, Le Liste, p. 4; pi. 410, fig. la and b of Encycloped. Method. ( no locality ) ; 1839, Anton, Verzeich. Conchyl., Halle, p. 84, no. 2780. Non I dime, 1758. 1821 Pterocera elongata Swainson, Exotic Conchology, Lon- don, pt. 1, sign. B4, 12th page; 1841, appendix, p. 32 (no locality). Also refers to Conchyl.-Cab., vol. 10, figs. 1479-80. 1842 Pterocera crocea Reeve, Conchologia Systematica, vol. 2, p. 204, pi. 248, fig. 2 (no locality); 1842, Sow- erby, Thes. Conch., vol. 1, pt. 2, pi. 11, fig. 4. [Indian Ocean?]. 1843 Pterocera novem-dactylis Deshayes, in Lamarck's ed. 2, Anim. sans Vert., vol. 9, p. 678, no. 9 (no local- ity); 1845, Kiister, Syst. Conchyl.-Cab., second series, vol. 4, pt. 1, p. 86. 1955 Lamhias (Millipes) elongata (Swainson; 1834), Butot, Penggemar Alam, vol. 35, p. 78, pi. 1, fig. 4. [Lambis], 1955 Lambis violacea Swainson, Abbott, Nautilus, vol. 68, no. 4, p. 124. Types — The type of Perry’s digitata has proba- bly been lost. I hereby restrict Perry’s type locality of “Eastern Ocean’ to Upolu Island, Western Sa- moa. Swainson’s type of elongata may be at Cam- [10-073] 1C4 Millepes R. T. Abbott S trombidae bridge University, or possibly lost. Nomenclature — There can be little doubt from Perry’s illustration that his S trombus digitatus is the species which Swainson later described and named elongate. Records — MOZAMBIQUE: Mozambique City (ANSP); Port Amelia (MCZ). MADAGASCAR: Sarodrano (Dautzen- berg, 1929, p. 473). MAURITIUS: (MCZ). PHILIPPINES: Mindoro Id.; Surigao, Mindanao Id. (both Elera, 1896, p. 255). SAMOA: Upolu Id. (Titian R. Peale, 1839, ANSP). Fossil records — None reported. Lambis scorpius (Linne, 1758) This handsome species is rather widely distrib- uted, and, although well known, it is comparatively uncommon. It is characterized by the gnarled or knobbed digitations and the brightly colored aper- ture which is purple and white-striped within and brownish to reddish orange at the border. In East- ern Polynesia, this species is replaced by the rare Lambis robust a (Swainson). The latter has smoother and heavier digitations and has no small, raised, white spiral ridge deep within the upper part of the aperture, a feature characteristic of scorpius. Indian Ocean specimens of scorpius show slight, but fairly constant, differences from the typical Pa- cific race, and for this reason we are recognizing two geographical races. In Pacific specimens, there is a large, flat ear-like lobe on the left side of the base of the first digitation. In the Indian Ocean subspecies indomaris, this lobe is greatly reduced or pushed back over the apex of the spire. Digita- tions 4, 5 and 6 are considerably more stunted and less protruding in indomaris than in the typical scorpius. Lambis scorpius subspecies scorpius (Linne, 1758) (PI. 121, fig. 5; pi. 129, fig. 4) Range — Indonesia and the Ryukyu Islands to Samoa. Remarks — The typical subspecies appears to be limited to the western Pacific. Its distinguishing characters are discussed above. Habitat — This species is found in shallow water at a depth of 1 to 10 feet, usually on coral reef flats where there is some protection from the ocean waves under or among dead coral slabs and boul- ders. Mrs. Anita Scott reports ( in lift. ) that they are more often, although uncommonly, found in shallow water during the warm summer months in the Ryukyu Islands. Description — Shell 100 to 165 mm. ( 4 to 6)2 inches ) in length, with 6 knobbed labial digitations and a long slender, knobbed siphonal digitation which is strongly curved to the right. Whorls 9 to 11. Nuclear whorls 3, smoothish, opaque-tan, and elevated. Sometimes with a weak spiral band of brown just below the suture. Postnuclear whorls flattish, and bearing numerous small knobs on a strong carina which is located just above the suture. Rody whorl with 3 main spiral rows of low knobs, the top one bearing the 4 largest, the middle row bearing 8 to 9 small knobs and the lowest row bear- ing 5 or 6 knobs of intermediate size. Outer lip bearing 6 digitations, all having 2 or 3 broad knobs, thus giving them a gnarled or jointed appearance. First digitation usually pointed upward in line with the axis of the shell, and bears a large, flat, ear-like lobe at the base on the left side. Digitations 4, 5 and 6 are bent posteriorly, almost at right angles to the axis of the shell. Siphonal canal long, strongly curved to the right, and bearing 4 or 5 weak knobs. Aperture rather narrow, quadrate, and deep within is solid purple. Inside of outer lip with numerous, crowded, white, slightly raised spiral lirae which end before they reach the broad, smoothish yellow- orange edge of the outer lip. Deep within the upper part of the aperture is a strong, white, bar- like, spiral lira. Columella and parietal wall brown- ish purple and overlaid with about 30 to 40 white to tan-white, raised, spiral lirae which may or may not run parallel with the spiral cords on the body whorl. Columella bounded by a long, narrow, white, axial swelling deep within the aperture. Periostra- cum thin, translucent-tan, and heaviest on the under surface of the digitations. Operculum long, slightly curved, light-brown and with 10 small serrations. Verge half the length of the aperture and with a small distal pad. Radula with 46 rows and a for- mula of 2-1-2; 1-3; 4; 5. Measurements (mm.) — (including digitations) length width no. whorls 168.0 66.2 8 + ( large; Okinawa ) 135.0 55.1 10 ( average; Truk Id. 97.2 42.0 8 + (small; Biak Id.) Synonymy — 1758 Stroinbus scorpius Linne, Systema Naturae, ed. 10, p. 743 (In O. Asiatico); 1767, 12th ed., p. 1208, no. 492; 1956, Dodge, Bull. Amer. Mus. Hist., vol. Ill, art. 3, pp. 249-250. 1771 Stroinbus scorpio L., Murray, Fundamenta Test., Amoenitates Academicae, vol. 8, p. 45, pi. L, fig. 29; 1778, Born, Index Rerum Natur. Mus. Caesarei Vindobon, p. 268. 1798 Lambis chiragra Gmelin, Roding, Museum Bolteni- anum, pt. 2, p. 67, no. 856, (Non Gmelin 1791, non Linne 1758). [10-074] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 165 1805 Pterocera scorpius L., Roissy, Buffon’s Hist. Nat. des Moll., vol. 6, p. 91, pi. 58, fig. 5. 1807 Pterocera scorpius Lin., G. Fisher, Museum Demidoff, Moscow, vol. 3, p. 191. 1810 Pteroceres scorpius L., Montfort, Conchyl. Systemat., Paris, vol. 2, p. 607. 1816 Pterocera nodosa Lamarck, in Bruguiere’s Encyclop. Method., pt. 23, Le Liste, Paris, p. 4, pi. 410, fig. 2; 1821, Swainson, Exotic Conchology, London, ap- pendix, p. 32 (sign, B3, lltli page). 1822 Pterocera scorpio Lin., Lamarck, Anim. sans Vert., vol. 7, p. 197. 1842 Pteroceras scorpio Linn., Sowerby, Thesaurus Con- chyl., vol. 1, pi. 11, fig. 1 (Moluccas). 1843 Pterocera scorpio Lam., Kiener, Coquilles Vivantes, vol. 4, pi. 6, (mer des Indes). 1845 Pterocera scorpio Murray, Kilster, Syst. Conchyl. -Cab., second series, vol. 4, pt. 1, p. 87, pi. 14, fig. 7. 1851 Pterocera scorpius Linne, Reeve, Conchologica Icon., vol. 6, Pterocera, pi. 3, fig. 3 (Philippines). 1955 Lambis (Millipes) scorpius Linne, Butot, Penggemar Alam, vol. 35, p. 77, pi. 2, fig. 5 (Madura, Java). Types — We restrict the type locality to Mactan Island, opposite Cebu City, Cebu Island, Philip- pines. Linnaeus’ type is in the Linnaean collection in the Linnaean Society of London, according to Dodge, 1956, p. 249. N omenclature — Linnaeus originally used the Latin substantive noun “Scorpius” for this species. Later authors (Murray, 1771, Born, 1778, and La- marck, 1822) used the more common form, “scor- pio.” We consider the use of the latter name to be an invalid emendation, and are following Butot (1955) who uses scorpius. All of Linne’s figure ref- erences are to the Pacific Ocean form, as are those * in Knorr, pt. 2, pi. 3, fig. 1 and Martini, Conchylien- Cabinet, vol. 3, fig. 860. Records (see map, pi. 131)— RYUKYU IDS.: Okinawa Id. (A. A. Scott and A. R. Cahn Colin., ANSP). TAIWAN: Kasyo-to ( Kuroda, 1941, p. 98). PHILIPPINES: Calapan and Lubang Id., Mindoro ( P. de Mesa, MCZ); Batang Id., Batanes Group (USNM); Borongan, Samar Id.; Gigmoto, Catanduanes Id.; Marivales, Luzon Id. (all du Pont-Acad- emy Exped., 1958); Jolo Id., Sulu Sea (ANSP); Cebu City, Cebu Id. (A. B. Franco, ANSP). INDONESIA: Sumatra: Atje (Zool. Mus. Amsterdam); Pandjang. Java: Djakarta Bay; Panaitan; Madura; Peutjang. Borneo: Sebuku. Also Roti, Timor, Ambon, Ceram, Ternate, Obi and Waigeu Islands (all Butot, 1955, p. 78). Maumerie, Flores Id. ( Rijksmus. Nat. Hist., ANSP). AUSTRALIA: Queensland (Cotton, 1953, no. 3, second page, fig. 16). DUTCH NEW GUINEA: reef at Biak ( NSF, 1956). SOLOMONS: Guadal- canar (AMNH). NEW CALEDONIA: barrier reef, Touho Bay (G. and M. Kline, 1959). MARIANAS: Apra Harbor, Guam Id. (MCZ and ANSP). CAROLINES: Moen Id., Truk (MCZ). Flfl IDS.: Suva, Viti Levu Id. ( H. S. Ladd, USNM). SAMOA: Tutuila Id. (ANSP and USNM); Upolu Id. ( T. R. Peale, Lb S. Explor. Exped., ANSP). [Tahiti rec- ords in the literature are probably based upon specimens of Lambis robusta (Swainson)]. Fossil records — None reported. Lambis scorpius subspecies indomaris new subspecies Range — Limited to western and central Indian Ocean. Remarks and Description — The differences ex- hibited in the shells of this Indian Ocean race and the typical Pacific Ocean race are slight, but con- stant, and warrant, we believe, subspecific recogni- tion. In scorpius indomaris, the lobe on the left side of the first digitation is either very much reduced or bent back around the apex of the shell. Digita- tions 4, 5 and 6 are considerably more stunted, their basal portions being not as long as the curved, distal portions. The dorsal sides of the terminal halves of the digitations and the siphonal canal are usually, although not always, more darkly pig- mented with purple-brown. There appears to be no or little difference in the apertural sculpture or coloring. Operculum with about 10 serrations and the radula indistinguishable from that of scorpius scorpius. Plate 131. Geographical distribution of Lambis scorpius domaris Abbott in the Indian Ocean, and L. robusta ( Swain- scorpius (Linne) in the central Indo-Pacific, L. scorpius in- son) in eastern Polynesia. [10-075] 166 Millepes R. T. Abbott Strombidae This subspecies was named sintiatus Perry, 1811 (non Solander, 1786) which is a homonym. I pro- pose the new name, indomaris. Habitat — This is an uncommon subspecies which lives in shallow water on coral and rock-strewn outer reefs. Also found on coral gravel shallows among beds of algae. Measurements (mm.) — (including digitations) length width no. who rls 135.0 50.0 7 + (holotype, ANSP no. 242210) 130.0 53.0 9 + (paratype, ANSP no. 212384) 111.0 45.0 8 + (paratype, ANSP no. 189165) 170.0 75.0 7 + (Tanikely Id., N.W. Madagascar) Synonymy — 1811 Strombus sinuatus Perry, Conchology, London, pi. 13, fig. 3 (American Seas, and at Madeira [both errone- ous]). Non Solander (or Humphrey) 1786. 1829 Pterocera pseudo-scorpio Lam., Schubert and Wagner, Conchyiien-Cabinet, Nurnberg, vol. 12, p. 16, pi. 218, figs. 3040-41 (locality unknown). Types — The type locality is Nossi-be, northwest Madagascar. The holotype is in ANSP no. 242210. Paratypes in ANSP are listed below. Records — (see map, pi. 131). MOZAMBIQUE: Mozam- bique City (K. Grosclg ANSP no. 247564). ZANZIBAR: outer reef, Kiwengwa (ANSP no. 212384); Pange Id. (ANSP no. 214263); Mangapwani (ANSP no. 212968) (all NSF, 1957). KENYA: Kikambala, 15 n. of Mombasa (Coryndon Mus.). EGYPT: Geb Zebara (ANSP no. 189165). MADA- GASCAR: Nossi-be (A. Chavane, ANSP); Tanikely Id. ( Academy-Madagascar Exped., 1960). CEYLON: [pre- sumed to be this subspecies] Condatchey, Gulf of Manaar ( le Beck, 1799, Philosophical Magazine, vol. 5, p. 339; and Langdon, 1874. p. 74). Andaman Islands: Port Blair; Long Island (both BM). Lienard (1877, p. 38 and 94) reported scorpius [this subspecies?] from Mauritius and Chagos Ids., Indian Ocean. Von Martens (1880, p. 279) reported it from the Seychelles and Amirantes. Fossil records — None reported. Lambis robusta (Swainson, 1821) (PI. 121, fig. 2; pi. 129, fig. 5) Range — Southeastern Polynesia. Remarks — Until recently, exact locality data was not known for this rare and poorly understood spe- cies. Such erroneous localities as Zanzibar and the Philippines had been listed by Reeve ( 1851 ) and others. A specimen was located in the Museum of Comparative Zoology which was collected at Ta- hiti, Society Islands, by W. H. Pease during the latter part of the Nineteenth Century. Dr. Robert Robertson collected five dead specimens on the same island in 1952. Although this species closely resembles scorpius, it is certainly quite distinct and evidently limited to French Oceania. L. robusta differs in having stouter, smoothish digitations of which the third and fourth usually have a common base, in having an almost straight siphonal canal, in lacking a lobe at the base of the first digitation, in lacking the short white bar deep within the upper part of the aperture, and in having, instead, a strong lobe deep inside on the upper portion of the columella. This last feature is difficult to see because it is set so deeply within the aperture. L. robusta ( Swainson, 1821), was formerly known as pseudoscorpio (La- marck, 1822). This species was characterized by Deshayes (1843) as being larger than scorpius, but many adults of robusta are smaller than some scorpius. Habitat — Unknown, but suspected to be in fairly deep water off the ocean edge of coral reefs. Description — Shell 110 to 150 mm. (4/2 to 6 inches) in length, with 6 moderately stout, smooth- ish labial digitations and a moderately long, slightly curved, siphonal digitation. Whorls about 9. Nu- clear whorls not observed. Postnuclear whorls flat- fish, and bearing numerous small knobs on a strong carina which is located just above the finely im- pressed suture. Top of apical whorls with numer- ous, microscopic, spiral threads. Rody whorl with 3 main spiral rows of low knobs, the top one at the shoulder bearing 5 or 6 large, irregularly-sized knobs, the middle row bearing 7 or 8 and the low- est row 6 or 7 smaller equal-sized, rounded knobs. Outer lip bearing 6 smoothish digitations, the first and uppermost being the stoutest and longest. Digi- tations 3 and 4 have a common base. Digitations 4, 5 and 6 have the terminal third bent posteriorly ( or upwards). Siphonal canal slightly curved to the right, and about the same length as the first digi- tation. Aperture rather narrow and quadrate, and deep within is yellowish cream, and, at the upper end has a small sunken depression which is not bounded below by spiral bar. Inside of outer lip tan with an irregular flush of brownish violet which is overlaid by numerous, irregular, raised spiral lirae sometimes extending to the outer edge of the lip. Parietal wall brownish to brownish purple with about 30 irregular, raised, spiral lirae. Columella bounded by a long, weak, narrow, cream-colored, axial swelling deep within the aperture. Periostra- cum moderately thin and light-brown. Operculum and soft parts unknown. [10-076] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 167 Measurements (mm.) — (including digitations) length width no. whorls 190.0 ( large, fide Tryon, 1885, p. 125 ) 153.0 72.0 7 + (U.S.N.M. specimen) 125.0 80.0 7 + (average, Tahiti) 111.0 58.0 7 + (small, Tahiti) Synonymy — 1821 Pterocera robusta Swainson, Exotic Conchology, sign. B 3, 11th page; 1841, appendix, p. 32 (no locality). 1822 Pterocera pseudo-scorpio Lamarck, Anim. sans Vert., vol. 7, p. 197 (no locality). Refers to Bonanni, pi. 3, fig. 312 and Lister, pi. 867, fig. 22; 1823, Dillwyn, An Index to Hist. Conchyl. Lister, London, p. 39; 1843, Deshayes, Anim. sans Vert., ed. 2, vol. 9, p. 674; 1870, Gill, Amer. Jour. Conch., Philadelphia, vol. 5, p. 134. 1842 Pteroceras pseudoscorpio Lam., Sowerby, Thesaurus Conchyl., vol. 1, P- 43, pi. 11, fig- 2 (no locality); 1851, Reeve, Conch. Icon., vol. 6, Pterocera, pi. 3, fig. 4 (Zanzibar). 1852 Pterocera ( llarpago ) pseudoscorpio Lam., Morch, Cat. Conchyl. Yoldi, p. 60. 1854 Harpago (Millipes) pseudoscorpio Lam., H. and A. Adams, Genera Recent Mollusea, vol. 1, p. 261. 1955 Lambis (Millipes) pseudoscorpio Lamarck, Butot, Peng- gemar Alam, vol. 35, p. 78. Types — We hereby designate Tahiti Island, So- ciety Islands, as the type locality. The whereabouts of Swainson’s type of robusta is unknown to me. The type of pseudoscorpio Lamarck is presumably in the Museum d’Histoire Naturelle de Geneve. The earliest figure is plate 867 in Lister, 1685. Records— (see accompanying map, pi. 131). SOCIETY ISLANDS: Tahiti (W. H. Pease, MCZ); Atiue, near Ma- ruapo River; near Punaauia Point ( all District of Punaauia, Tahiti, R. Robertson, 1952, ANSP). LINE ISLANDS: Flint Island (C. D. Voy, ANSP). [Records for Zanzibar, Philip- pines, Australia are probably erroneous]. Fossil records — None reported. Lambis violacea (Swainson, 1821) (PI. 121, fig. 1; pk 129, fig. 1) Range — Indian Ocean. Remarks — This is one of the rarest and most at- tractive of all the Lambis. I would judge that there are no more than a hundred known specimens, and most of these have come from Mauritius. The other localities mentioned in the literature may not be accurate. I am puzzled by Butot’s ( 1955, p. 79 ) record from Indonesia. Lambis violacea is readily recognized by its yel- lowish-white shell, and by its whitish aperture which is tinted with violet or lavender deep within the throat. The spiral lirae on the inside of the outer lip are fine, rather even, numerous and white. The labial digitations may vary in number from 9 to 11. Habitat — Unknown, although it probably lives on sand and algae bottoms at a depth of 20 to 60 feet. Description — Shell 73 to 114 mm. (about 3 to 4h inches) in length, with 10 to 11 thin, blade-like, short labial digitations and a moderately long, slightly twisted, siphonal digitation. Whorls about 9. Nuclear whorls not observed. Post-nuclear whorls slightly concave, and sharply carinate just above the suture. This shoulder carination bears numer- ous, evenly-sized, nodules which are crossed by 4 to 5 small, but distinct, spiral cords. Above the cari- nation, the whorls are crossed by about a dozen fine spiral threads and by numerous axial threads, thus giving a somewhat reticulated effect. Shoulder of body whorl with a row of 5 or 6 paired knobs, the last one being low and elongate and the next to last one being the highest. Below these, and on the center of the body whorl, are 3 strong beaded cords. Remainder of whorl with numerous, smooth- ish, smaller, spiral cords. Outer lip with 9 to 1 1 digitations whose undersurfaces are smoothish, enamel-white and somewhat concave. First digita- tion obscures the apex of the shell and is bifurcate, the prong to the left being shorter and broader. The other 8 or 9 digitations become progressively smaller towards the anterior end. “Stromboid notch” deep and U-shaped. Below it, the base of the outer lip bears 3 or 4 very small digitations. Siphonal canal is twisted but descends almost straight down. Aperture somewhat quadrate and violet or laven- der deep inside, except for a white depression near the top. Outer wall of aperture with about 80 fine, strongly raised, spiral, white lirae. Outer edge of aperture with large, weak, yellowish spots. Parietal wall weakly and unevenly lirate, its lower section strongly swollen, smooth and tan or purplish cream. Outer shell whitish with a few isolated, small, squarish, light-brown spots. Periostracum unknown. Operculum chitinous, elongate, light-brown and with smoothish edges. Radula unknown. Measurements (mm.) — (including digitations) length width no. whorls 113.5 55.1 74 (Mauritius, ANSP) 73.0 42.0 8 (Mauritius, MCZ) [10-077] 168 Millepes R. T. Abbott S trombidae Synonymy — 1817 Strombus millepeda L., variety C, Dillwyn, Descript. Cat. Recent Shells, London, vol. 2, p. 660 ( refers to Conchyl.-Cab., vol. 10, figs. 1494-95). 1821 Pterocera violacea Swainson, Exotic Conchology, sign. B4, 12th page; 1834, ibid., appendix, p. 33 (Red Sea); 1841, ibid., ed. 2, p. 33. 1823 Strombus purpureus Mawe, Linne’s Syst. Conch., Lon- don, p. 127 (nude name). 1825 Pterocera purpurea Swainson, Dubois, An Epitome of Lamarck’s Arrangement of Testacea, London, p. 248; 1837, Swainson [?], Catalogue of the Foreign Shells Manchester Nat. Hist. Soc., p. 74 (both nude names ) . 1842 Pteroceras multipes “Chemn." Sowerby, Thesaurus Conchyl., vol. 1, p. 43, pi. 11, fig- 8; 1842, Reeve, Conchologiea Systematica, vol. 2, p. 204, pi. 248, fig- 1. 1842 Pteroceras purpurascens “Swainson” Sowerby, loc. cit., p. 43 (in synonymy). 1S43 Pterocera multipes Deshayes, in Lamarck’s ed. 2, Anim. sans Vert., vol. 9, p. 677, no. 8 ( mers de l’lnde). Also refers to Conchyl.-Cab., vol. 10, figs. 1494-95; 1850, Traite Element. Conchyl., pi. 115, fig. 7. 1843 Pterocera millepeda var., Kiener, Coquilles Vivantes, Paris, vol. 4, Pterocera, p. 11, pi. 10, fig. 1. 1955 Lambis (Millipes) violacea (Swainson, 1834), Butot, Penggemar Alam, vol. 35, pp. 78-79, pi. 2, fig. 6 ( Madura, Indonesia ) . Types — The location of Swainson’s type is un- known to me, although it may be at Cambridge University. The type locality is the ‘‘Red Sea.” Nomenclature — The earliest valid name appears to be violacea Swainson, 1821. Dillwyn in 1817 used the name multipes Chemnitz only in synon- ymy under his millepeda L. variety C, and is, there- fore, not validly used. The name purpurea used by Swainson and Mawe is nude. Records — (see pi. 130) Mauritius ( N. Pike, MCZ; ANSP). Cargados Island (J. Robillard, Nat. Mus. Viet.; Lienard, 1877, p. 109). Zanzibar (MCZ). Seychelles ( H. Cuming, Nat. Mus. Viet.). Madagascar ( McCill Redpath Mus.). Red Sea (Swainson, 1841, p. 33). INDONESIA: Ambat, east Madura (Butot, 1955, p. 79). [Philippine records have never been confirmed]. Fossil records — None recorded. [10-078] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 169 Subgenus Harpago Morch, 1852 Type: Strombus chiragra Linne, 1758. The speciation problem in this group is very in- triguing and not entirely solved to our satisfaction. Collecting of live material in Indonesia and Poly- nesia will doubtlessly settle the matter. There are three shell forms which have been variously treated by earlier authors as forms, subspecies or even separate species. One of these, arthritica Roding (see our pi. 121, fig. 7), is a shell of quite constant characters and limited to the Indian Ocean. There are no signifi- cant differences between the shells of the males and females, other than the slightly smaller size of males. We have been unable to ascertain the east- ern limits of this form. Two of Butot’s specimens from Indonesia (his pi. 3, figs. 5 and 6) look like arthritica. I have also seen specimens labelled “Philippines” but this locality may be in error. The second kind of shell is the typical and tra- ditionally accepted chiragra (Linne). This is the large, rose-mouthed form with an almost smooth, lower columella, which is common in the western Pacific ( see our distributional map, pi. 134, and our pi. 121, fig. 12). This is the female shell which is further characterized by the welding together of the last two knobs on the shoulder of the body whorl. The third form we believe is the male shell of chiragra. It has two peculiarities which curiously enough resemble characters found in the Indian Ocean arthritica. These are a tendency to develop white spiral lirae on the purplish brown columella, and the presence of equal-sized knobs on the shoul- der. The shell as a whole is generally much smaller than the female shell. The columellar lirae and purple taint vary from a strong development (see our pi. 121, fig. 10) to a weak development (pi. 121, fig- 11). Intermediate shell forms between males and females are not uncommon ( see pi. 132, figs. 2-4). Below, we are presenting our classification and a synopsis of the distinguishing characters. For con- venience, we have retained the form name rugosa ( Sowerby ) for the male, and have listed its syn- onymy separately. Lambis chiragra chiragra (Linne). Typical or female form: 6 to 10 inches ( 150 to 250 mm. ) in length; aperture whitish rose; lower columella smoothish (aperture of Bay of Bengal specimens with a purplish stain possi- bly due to environment); upper end of aper- ture with an elongate, whitish depressed well or shelf; last two knobs of shoulder larger than the others and welded together; the 5th labial digitation produces a rather high ridge on the dorsal part of the base of the last whorl. Male or rugosa (Sowerby) form: 4 to 7 inches (100 to 175 mm.) in length; aperture rose to reddish with slight or strong purplish mark- ings; lower columella slightly to strongly lirate; upper end of aperture with an elongate, purple and white, depressed well or shelf; last two knobs of shoulder small and not welded to- gether; 5th labial digitation produces a very Plate 132. Fig. 1, Lambis chiragra subspecies arthritica rugosa in L. chiragra chiragra (Linne). All about Vz natural Roding from East Africa. Figs. 2-4, intergrading variations size, in the columellar coloration and spiral lirae in the form [10-083] 170 Harpago R. T. Abbott Strombidae low ridge on the dorsal side of the base of the last whorl. The columella of some males is the same as that in the females. Lambis chiragra subspecies arthritica Roding Male and female form: 5 to 6/i inches ( 127 to 173 mm.) in length; aperture yellowish white; en- tire columella purplish brown with strong, white, spiral lirae which parallel the spiral cords ( in form rugosa, the lirae cross the cords at a slight angle ) ; upper end of aperture with- out an elongate, deep depression; last two knobs the same size as the others on the shoul- der; 5th labial digitation produces a very low ridge on the dorsal side of the base of the last whorl. It is rather curious that the few records we have seen from eastern Polynesia are the rugosa form. This suggests either that only males have been so far collected or that the females also take on the mgosa-type characters. If the later be true, we would have an interesting situation in which major sexual dimorphism occurs in the center of distribu- tion while at both ends of the range there is little or no difference in the shells of the two sexes. Synonymy — 1839 Pterocera Lam., Anton, Verzeichniss Conchylien, Halle, p. 84 (type by subsequent designation: chi- ragra Linne)- 1852 Harpago “Klein ”, Morch, Catalogus Conchyliorum . . . Yoldi, Hafniae, p. 60 (type by indirect tautonomy: Lambis harpago Bolten = chiragra Linne); 1854, H. and A. Adams, Genera of Recent Mollusca, London, vol. 1, p. 261; 1929. Thiele, Handbuch Syst. Weicht., vol. 1, p. 255; 1940, Wenz, Handbuch cler Palaont., vol. 6, pt. 4, p. 946 ( type by subsequent designa- tion: chiragra Linne). 1870 Harpago LI. and A. Adams, Gill, American Journ. Conch., vol. 5, no. 3, p. 136. 1870 Pterocera Lamarck, Kobelt, Illustrirtes Conchylien- buch, vol. 1, p. 105 (type by subsequent designa- tion: Pterocera chiragra Linne). Lambis chiragra subspecies chiragra (Linne, 1758) (PI. 121, figs. 10-12; pi. 132, figs. 2-4) Range — Eastern Indian Ocean to eastern Poly- nesia (but not now living in Hawaii). Remarks — Comparative notes are given above in the subgeneric discussion. The females have large shells with a whitish rose aperture and a whitish, smooth lower columella. The male shells are usu- ally smaller and sometimes with strong, white, spi- ral lirae laid over a darkly-stained, purple colu- mella. These lirae cross the spiral cords of the body whorl at a slight angle, whereas in arthritica they are parallel to the spiral cords. In the Bay of Ben- gal and Western Australia, specimens of chiragra usually lack the rose coloration in the aperture and, instead, may have a dirty brownish stain which is probably due to environmental conditions. Suites of dwarf specimens sometimes found in museum col- lections are probably male shells selected out by collectors in the field, and do not represent a dwarf race. Dautzenberg’s (1929, p. 474) records of chiragra from Madagascar ( which are based upon identifica- tions by Sganzin, von Martens, Thiele and Odhner), undoubtedly are misidentifications of specimens of Lambis arthritica Roding. The latter is moderately common in that area, and we have not seen a speci- men of typical chiragra with reliable locality data from the western part of the Indian Ocean. E. A. Smith (1903, p. 613) reports chiragra from the Maldives and Laccadives southwest of India, but these are probably also arthritica. Thiele’s Hand- buch figure ( 1929, p. 255, fig. 266 ) of “ chiragra ” is certainly arthritica. We have seen less than a dozen specimens of chiragra from eastern Polynesia, and they have all been the male rugosa form. Whether or not the fe- male shell exhibits the characters of large size and whitish rose aperture is not known. It is possible that there is less sexual dimorphism in the east- ern part of the range. I have seen typical rugosa forms from Okinawa, Ryukyu Islands; “Philip- pines”; Touho Bay and Noumea, New Caledonia (ANSP); Ngarumaoa Id., Raroia Atoll, Tuamotu Ids. (USNM); Hikueru, Tuamotu Ids. (San Diego Soc. Nat. Hist.); Saipan Id., Marianas; the Society Islands; and from the Line Islands (B. P. Bishop Mus.). Dautzenberg and Bouge’s (1932, p. 302) records of Pterocera rugosa from the Societies and Tuamotus are probably male chiragra. Lambis chiragra has not been found alive in Hawaii, hut we have examined several late Pleisto- cene specimens from Oahu Island ( through the kindness of Clifton Weaver, H. M. Baker and Karl Greene ) . Habitat — Demond (1957, p. 297) states that this species, in the Marshall Islands, “lives on seaward reef Hats, in sand between rocks and coral heads, and in tide pools. Also found among masses of coral (Heliopora) in channels between seaward reefs in 8 to 10 feet of water. Invariably found below low tide line.” George and Mary Kline collected males and females on the barrier reef in 4 to 10 feet of water at Touho Bay, New Caledonia. In other areas, such as the Philippines and Palau Islands, it is also associated with sand, coral and algal covered reefs where there is surging of oceanic waters. [10-084] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 171 Description of shell of female — Shell large, 150 to 250 mm. (6 to 10 inches) in length, massive, with 5 large labial digitations and with the siphonal canal turned to the left. Large “stromboid” notch located between the 4th and 5th labial digitations. Whorls 10 to 11. Whorls in spire concave, bordered above and below by two raised, spiral cords, be- tween which runs the finely indented suture. Spiral sculpture of numerous, small threads and, on the body whorl, of 4 or 5 large spiral cords. The upper cord at the shoulder bears 6 to 7 large knobs of which the last two are the largest and are welded together. The lowest fourth cord is the weakest, but it extends on to the dorsal side of the 5th labial digitation and there forms a raised ridge. Color of outer shell whitish to cream with crowded, zigzag streaks of light- or purple-brown. Color of deep interior of aperture is white. Throat of aperture somewhat constricted, flushed with rose and bear- ing short, white, raised, spiral lirae. Parietal wall slightly glazed, cream with brown, axial streaks. Lower columella area made into a raised shield which is smoothish, cream and tinted with rose. At the upper or posterior end of the throat of the aperture there is an elongate, concave, greatly de- pressed, whitish lobe attached to the inner parietal wall. Ceylon specimens usually have a brown or purple-brown stain over all of the apertural area. Periostracum moderately thin, brown, axially stri- ate, but usually worn off on the dorsal sides of the digitations and body whorl. Operculum chitinous, dark-brown, fusiform, with about f6 fine serrations on one side (commonly worn away), usually filling the constricted throat of the aperture and with the muscle attachment scar being half the area of the entire operculum. Description of shell of male ( form rugosa ) — Shell similar to that of the female, but about ’2 to h the size; fOO to 175 mm. (4 to 7 inches) in length. Shoulder with 4 to 6 rather evenly-sized small knobs. Throat and columella pinkish or strongly stained with purple-brown over which run weak to strong, white, raised spiral lirae which cross the spiral cords of the parietal wall at an oblique angle. The inside of the outer lip may be reddish in some specimens. Operculum and periostracum like those in the female. Radula ribbon with about 46 transverse rows of teeth. Formula: 2-1-2; 1-3 (sometimes 1-4); 5; 7. Verge moderately long, simple and with a lami- nated pad near the distal end. Measurements (mm.) — (including digitations) length width no. whorls 260.0 190.0 9 + (large, female; Ceylon) 246.0 170.0 8 + (large, female; Philippines) 150.0 85.0 9 + (large, male; New Caledonia) 95.0 65.0 8 + (small, male; Palau Ids.) Synonymy — The names listed below are of chi- ragra (Linne). For convenience and for a clearer understanding of the history of the names applied to the female and male forms, we are listing the synonyms of both forms separately. Synonymy of female — 1555 Strombus , Belon, La Nature et Diversite des Poissons, Paris, p. 423, fig. ( non-binomial ) . 1758 Strombus chiragra Linne, Systema naturae, ed. 10, p. 742, no. 423 (Ad Bandam Asiae); 1767, ed. 12, p. 1207, no. 491; 1956, Dodge, Bull. Amer. Mus. Nat. Hist., vol. Ill, art. 3, pp. 247-249. 1798 Lambis harpago Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 67, no. 860, Substitute name for chi- ragra Gmelin, 1791. 1798 Lambis undulata Roding, loc. cit., p. 68, no. 873 (re- fers to Conchyl.-Cab., vol. 3, fig. 898, a young of chiragra? ) . 1842 Pteroceras chiragra L., Sowerby, Thesaurus Conch., vol. 1, p. 42, pi. 11, fig. 12. Plate 133. A Pleistocene specimen of L. chiragra chiragra ( Linne ) from a raised beach near Sunset Beach, Oahu Is- land, Hawaii, (from the Children’s Museum of Honolulu). Slightly reduced. [10-085] 172 Harpago R. T. Abbott S trombidae 1855 Pterocera kochii Freyer, Sitzungb. Math.-Natur. der Kaiser. Akad. der Wissenschaften Vienna, vol. 15, p. 22 (no locality). Refers to Kiener, 1843, pi. 5. 1938 Lambis (Harpago) chiragra (Linne), Hirase, A Collec- tion of Japanese Shells, Tokyo, 6th ed., pi. 88, fig. 2; 1949, Abbott, Scientific Monthly, vol. 69, p. 325, left fig. 1940 Pterocera (Harpago) chiragra (Linne), Wenz, Hand- buch der Palaozoologie, Berlin, Lief. 6, band 6, p. 948, fig. 2764. 1950 Lambis chiragra Linne, Abbott, Bull. Raffles Mus., Singapore, no. 22, p. 74 (Cocos Keeling Ids.). Synonymy of male or form rugosa — 1823 Strombus chiragra L., Mawe, Linnaean Syst. Conch., London, p. 125, pi. 25, fig. 4. Not Linne, 1758. 1842 Pteroceras rugosum Sowerby, Thesaurus Conchyli- orurn, vol. 1, p. 42, pi. 11, fig. 9 (not 10). 1851 Pterocera rugosa Sowerby, Reeve, Conchologia Iconica, vol. 6, Pterocera, pi. 4, sp. and fig. 6 ( Society Is- lands). Non Sowerby, 1842; 1936, Robert, Kunstge- bilde des Meeeres, Bern, p. 15, pi. 14 (Neu- Kaledonien ) . 1911 Pterocera (Harpago) rugosa Sowerby, Dautzenberg, Bull, l’lnstitut Oceanograph., Monaco, no. 161, p. 3. 1938 Lambis (Harpago) chiragra rugosa (Sowerby), S. Hi rase, A Collection of Japanese Shells, Tokyo, pi. 88, figs. 3. 1949 Lambis rugosus, Platt, National Geographic Magazine, vol. 96, no. 1, p. 68, fig. 1 (in color). 1949 Lambis arthritica Roding, Abbott, Scientific Monthly, vol. 69, no. 5, p. 324, fig. upper center. 1953 Harpago chiragra Linne, Cotton, Malacol. Section, Royal Soc. South Australia, no. 3, Family Strom- bidae, fig. 18. 1959 Lambis chiragra rugosa (Sowerby), Sakurai in Oyama, The Molluscan Shells, Resources Exploitation Insti- tute, Tokyo, vol. 2, Lambis pi. 3, figs. 2, 3. Types — Dodge (1956, p. 247) states that “the specimen marked for chiragra in the Linnaean col- lection in London is a typical and perfect specimen of the chiragra of all authors.” Hanley ( 1855, p. 366) likened the type to the shell illustrated by Sowerby (Thesaurus Conchyl., vol. 1, pi. 11, fig. 12). That shell is probably a large female and the color form from Ceylon. Mr. Peter Dance (1960) was unable to locate for us the type of Sowerby’s Pteroceras rugosum. Sowerby illustrated both arthritica Roding (his fig. 10) and the male form of chiragra (his fig. 9). He speaks of the two varieties. His reference to the “rose aperture” suggests the Pacific male form. I hereby designate his pi. 11, fig. 9 as representing the holotype. Records (see accompanying map, pi. 134) — [We believe literature records for Madagascar and East Africa are er- roneous]. CEYLON; Hikkaduwa (G. and M. Kline, NSF); reef, Foul Point (W. D. Hartman, Yale Peabody Mus.). ANDAMAN IDS.: Port Blair ( W. N. Carpenter, USNM). THAILAND: Phuket, Bay of Bengal (F. N. Crider, ANSP). Siantan Ich, Anambas Ids., east of Malaya (ANSP). SOUTH CHINA SEA: Paracel Islands ( Saurin, 1960, p. 204). COCOS KEELING: (USNM). RYUKYU ISLANDS: Oki- nawa Id. (W. A. McCarty, ANSP); Shiyo, Shanawan Bay, Okinawa Id. (USNM). TAIWAN: Kasyo-to; Hukukaku (Kuroda, 1941, p. 98). PHILIPPINES: Tabaco, Albany Prov., Luzon Id.; Cuyo Id. (both clu Pont-Academy Expech, 1958); Iba, Zambales Prov., Luzon Id. (P. de Mesa, ANSP); San Pedro Bay, Samar Id.; Mindoro Id.; Surigao, Mindanao Id.; Tataan Id., Tawi Tawi Ids.; Camiguin Id.; Dumaguete, Negros Id.; Batan Id., Batanes Group; Leyte Id.; Busuanga Id. (all USNM); Sanga Sanga Id., Sulu Arch. (ANSP); Jolo Id., Sulu Arch. (E. Gutianjo, ANSP). INDONESIA: Sumatra: Sinabang Bay, Simalur Id. (west coast); Biliton Id. (east coast). Java: Leiden Id., Bay of Batavia; Bali Id.; Timor Id.; Great Obi Id., Moluccas; Gulf of Madjene, Cele- bes (all ex Rykmus. Nat. Hist., Leiden, ANSP). Moluccas: Bouro Id.; Saparua Ich; Batjan Id. (all MCZ). AUS- TRALIA: 14 mi. north of Warroora Sheep Station, 150 mi. north of Carnarvon, West Australia ( Arch Whitworth, ANSP). Green Id., Queensland (Tony Marsh, in lift.). DUTCH NEW GUINEA: Mios Woencli Atoll, Padaido Ids.; Maransabadi Ich, Aoeri Ids. (both A. J. Ostheimer, NSF). SOLOMON ISLANDS: Tai Lagoon, Malaita Ich (Calif. Acach Sci. ). Roriana Lagoon, South New Georgia Ich (MCZ): Lutee, Choiseul Ich (W. J. Eyerdam, ANSP). NEW HEBRIDES: Lamap, Mallicola Id. ( G. Massoulard, ANSP). NEW CALEDONIA: barrier reef, Touho Bay ( G. and M. Kline, NSF). MARIANAS: Saipan Id. (MCZ; USNM); Guam Id. (USNM). PALAUS: Kayangel; Babel- thuap; Koror; Eil Malk; Gorokottan (all A. J. Ostheimer, NSF). Helen Ich, Helen Reef (V. Orr, NSF). CARO- LINES: Ulithi; Ponape; Elato; Ifaluk (all USNM). MAR- SHALLS: Bikini; Rongelap; Eniwetok; Wotho; Ujelang (all USNM); Ebon and Jaliut (MCZ). GILBERTS: Onotoa (USNM). SAMOA: Tutuila Ich (ANSP). SOCIETY IDS.: (San Diego Soc. Nat. Hist, form rugosa ). WAKE ID.: (A. E. Tara, AMNH ). PHOENIX GROUP: Baker’s Id. ( Linne) . [10-086] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 3 Lambis 173 (B. P. Bishop Mus.). LINE IDS.: Fanning and Christmas Id. ( B. P. Bishop Mus.). TUAMOTU IDS.: Hikueru (San Diego Soc. Nat. Hist.); Ngarumaoa Id., Raroia Atoll (J. P. E. Morrison, USNM); both form rugosa; Hao and Otepa (M. Couturier, 1907, p. 153), form rugosa. MARQUESAS: (Tony Gavaldon, coll., 1959), form rugosa. Fossil records — HAWAIIAN CHAIN: all Late Pleisto- cene; Midway (USNM); Sunset Beach, Oahu (Childrens Museum); Kapaloma Basin, Oahu ( H. M. Baker, coll’n.); side of hill, 15 feet above high tide, Makua, Oahu (Clifton Weaver, coll’n.); lava-coral formation, Waimea Bay, Oahu ( H. M. Baker, coll’n.); Kauaiu Stream, 250-290 ft. alt., Lanai Id. (Id. Stearns, USNM 496365); Honolulu Harbor and Kupikipikio, Oahu Id. (J. M. Ostergaard, 1928, p. 26). Lambis chiragra subspecies arthritica Roding, 1798 (PI. 121, fig. 7; pi. 132, fig. 1) Range — East Africa to the Central Indian Ocean. Remarks — This distinct subspecies appears to be limited to the western half of the Indian Ocean. Literature reports of it in the East Indies are prob- ably based upon the similar-looking male phase or rugosa form of L. chiragra. L. chiragra arthritica differs from male chiragra chiragra in lacking the deep-set, elongate well or depression at the upper end of the aperture, in having the white spiral lirae on the parietal wall running parallel with the spiral cords (instead of slightly oblique), and in gener- ally having a yellowish rose (rather than a pinkish rose) background color to the aperture. Both Abbott (1950, p. 74) and Butot (1955, p. 79) seem not to have recognized the problem of subspecific and sexual differences in the chiragra- arthritica complex. Although Abbott saw inter- grades in columella characters in the Pacific shells and rightfully suspected a close connection between the two forms in chiragra, he did not realize that he was working with a sexually dimorphic species. He also failed to recognize the morphological and geographical uniqueness of the western Indian Ocean subspecies arthritica. Butot recognized the distinctiveness of arthritica, but erroneously consid- ered the Pacific males of chiragra (the form rugosa) as part of arthritica. Thus his records of arthritica from the Ryukyu Islands, Funafuti, New Caledonia (Ceylon and Indonesia?) are doubtlessly based upon male chiragra. In the shells of male and female arthritica I can find no startling differences, other than the slightly smaller size of males and a very slight indication of larger or more elongate shoulder knobs on the shells of females. Habitat — The Natural Science Foundation expe- dition to Zanzibar in 1957 found this subspecies reasonably common just below the low tide mark on flat, offshore reefs which were largely covered with algae and marine grass. They found them in company with Lambis crocata (Link), Vasum rhi- noceros (Gmelin), Conus, Haliotis and some live coral. A label written by Mr. Kurt Grotsch (ANSP 192618) reports that in Mozambique they are found rarely at all seasons “just above the low water level on weed-covered, stony reefs where a swift current moves. In 2 to 3 fathoms of water they live among corals and gravel and their shells are eroded.” Description — Shell ( including digitations ) 120 to 190 mm. (5 to 7 inches) in length with 5 labial digitations and with its siphonal canal turned to the left. Large “stromboid notch” located between the 4th and 5th labial digitations. Similar to chi- ragra chiragra, but instead of having a deep de- pression at the upper end of the aperture, there is an arching broadly rounded, slightly depressed shelf projecting from the upper part of the parietal wall. The color of the inside of the outer lip is usu- ally yellowish or pinkish yellow with splotches of purplish underlying the white spiral lirae. The columella is brownish purple with strong whitish spiral lirae which tend to parallel the 4 spiral cords on the parietal wall. On the dorsum of the body whorl, the top spiral cord bears 7 to 9 evenly-sized, rounded nodules. Periostracum thin, varnish-like and translucent-yellowish. Measurements (mm.) — (including digitations) length width no. whorls 190 93 7 + (large; Mozambique) 140 75 8 + (average; Zanzibar) 125 70 7 + ( small; Zanzibar ) 120 70 9 + ( small, male, Seychelles ) Synonymy — 1798 Lambis arthritica Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 67, no. 858 (no locality). Refers to Conchyl.-Cab., vol. 3, fig. 857 (which is from Mauritius). 1811 S trombus divergens Perry, Arcana, London, vol. 2, pi. 74 (no locality ) . 1811 Strombus nigricans Perry, Arcana, London, vol. 2, pi. 74 (no locality). Additional name for divergens. 1842 Pteroceras rugosum Sowerby, Thesaurus Conehyliorum, vol. 1, p. 42 (South Sea), pi. 11, figs. 10 (not 9). In part. 1842 Pterocera rugosa Sowerby, Reeve, Conchologia Syste- matica, vol. 2, p. 204, pi. 247, fig. 1. 1855 Pterocera chiragra Linne, Freyer, Sitzungb. der Kaiser Akad. des Wissenschaften, Vienna, vol. 15, p. 22. Refers to Lister, pi. 870. 1859 Pterocera chiragra Linne, Chenu, Manuel de Conchy!., Paris, vol. 1, p. 258, fig. 1612. Non Linne 1758. 1929 Pterocera (Harpago) chiragra (Linne), Thiele, Hand- buch der System. Weich., fena, vol. 1, p. 255, fig. 266. [10-087] 174 Harpago R. T. Abbott Strombidae Types — Roding’s arthritica is based upon a figure of a specimen from Mauritius which we hereby designate as the type locality. Records — ( see map on pi. 134) MOZAMBIQUE: Mo- zambique City (K. Grotsch, ANSP); Port Amelia ( MCZ; USNM); KENYA: Diani Beach, 20 miles south of Mom- basa ( R. T. Abbott, MCZ; USNM); Wasin Icb, off Shimoni (J. K. Howard, MCZ); Malindi (USNM). ZANZIBAR: rock reef, 5 mi. south of Paje; reef off Ras Nungwe; outer reef at Kiwengwa (all NSF, 1957). MADAGASCAR: Nossi- be (A. Chavane, ANSP). SEYCHELLES: Bird Id., Frigate Id.; Beau Vallon, Mahe Id. (all Yale-Peabody Mus., 1957); Anse Boileau, Mahe Id. (Wickworth, Brit. Mus.). MAURI- TIUS: (N. Pike, MCZ). REUNION ID.: (MCZ). MAL- DIVES: Kureduls, Fadiffolu Atoll (Yale-Peabody Mus., 1957). CHAGOS IDS.: ( Melvill, 1909, p. 94, “Investiga- tor”. [the MCZ has records for “Fiji, Singapore and Am- boina”, but I suspect manufactured data, since they came from old private collections]. Fossil records — None recorded. Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [10-088] INDO-PACIFIC MOLLUSCA, vo). 1, no. 4 Pinnidae 175 ~dS September 28, 1961 '•J- - I fx THE FAMILY PINNIDAE IN THE INDO-PACIFIC by Joseph Rosewater Division of Mollusks lTnited States National Museum Washington, D. C. The bivalve family Pinnidae is unique among the anisomyarian clams which have a large posterior adductor and a small or no anterior adductor mus- cle. Commonly called Pen-, Fan-, Wing-Shells or Spanish Oysters, their shells are large, thin, broad and fragile. There are two layers of limy material: an outer prismatic sheath of very large crystals of calcite produced at a right angle to the long axis of the shell (pi. 135, and pi. 136, fig. 2), and a par- tial inner, shingled layer of nacreous material of aragonite produced parallel to the long axis of the shell (pi. 136, fig. 2). The periostracum is usually worn away, and hinge teeth are lacking in the Pin- nidae. The valves are united along their dorso- anterior margins by a dark, nonelastic primary liga- ment; and along their dorso-posterior margins by a fusion of the outer prismatic shell layer called the secondary ligament (Yonge, 1953). The anatomy of these animals is even more specialized than their shells, exhibiting several unique organs, including the protrusible pallial organ, the eye-like Organs of Will found along the margin of the mantle, and a pair of elongate, gutter-shaped waste canals which aid in ejecting debris from the mantle cavity (pi. 137). These morphological modifications aid the sessile pen-shells in surviving in their habitat rooted deeply in sandy mud, where currents and predators are likely to uproot and destroy them. Little is known of the reproduction and develop- ment of the Pinnidae. Species which have been studied were found to be dioecious. Cahn ( 1951 ) reported dates of spawning and growth rates of Atrina japonica in Japan ( = A. pectinata). Yoshida (1956) studied the early life history of pectinata. However, to date, no work has been done on the embryology of any species. Young pinnas begin their existence as minute equilateral bivalves (pi. 139). Growth proceeds in a posterior direction leav- ing the equilateral embryonic valves perched on the umbos of the young adult form. The former are usually worn away before the latter reaches much more than a centimeter in length. At present we know of twenty well-defined spe- cies and subspecies of Pinnidae, but when more adequate material is available for study from col- lections from shallow waters in the tropical and temperate regions of the world this number may be increased to as many as thirty or more. We recog- nize three distinct Recent genera: Pinna, Atrina and S treptopinna. The latter is known only from the Indo-Pacific, while the other two are found in all warm seas. Turner and Rosewater (1958, John- sonia, vol. 3, no. 38) dealt with the Western At- lantic species in considerable detail. In this study, nine Recent and several Tertiary species of Indo- Pacific Pinnidae are considered. Fossil Pinnidae The Pinnidae first appeared in the geologic rec- ord during the Paleozoic Era. Of the Recent genera, Atrina Gray appeared during the Carboniferous and Pinna Linne later, in the Jurassic. The order of appearance of these genera may indicate a tend- ency for development through time of more com- plex shell structure in this family. The genus Streptopinna probably developed during the late Tertiary. Several extinct genera have been described in the Pinnidae. Turner and Rosewater (1958) have discussed these, and Vokes [1951, pp. 40, 116] gave a complete list of the genera. These genera are represented by species apparently having more generalized shells than Recent Pinnidae and show affinities with other closely related bivalve groups such as the Mytilidae, Isognomonidae, Pteriidae and Ostreidae. There has been a relatively large number of fos- sil Pinnidae described from strata from widespread geographic areas in both Europe and the Western Hemisphere. Many of these are from the Paleozoic, but Mesozoic and Quaternary fossils are also plenti- ful. Species described by early workers are almost impossible to identify. The Pinnidae, because of the fragility of their valves, became poor fossils and usually are represented by fragments only. Descrip- 153-501] 176 Economic Importance J. Rosewater Pinnidae tions are based on supposed shapes of valves pro- jected from the fragments and characteristics of visible sculpture which, even in Recent species, may tend to be convergent. In the present study an attempt has been made to include all of the Indo- Pacific fossil species described from the Tertiary to Recent, and fossil records are given for Recent species. Economic Importance The Pinnidae have considerable economic impor- tance in many parts of the world. They produce pearls of moderate value. In the Mediterranean area, material made from the holdfast or byssus of Pinna itobilis Linne has been utilized in the manu- facture of clothing for many centuries: gloves, shawls, stockings and cloaks. Apparel made from this material has an attractive golden hue and these items were greatly valued by the ancients. Today, Pinnidae are eaten in Japan, Polynesia, in several other Indo-Pacific island groups, and on the west coast of Mexico. In Polynesia, the valves of Atrina vexillum are carved to form decorative articles, and entire valves of larger specimens are sometimes used as plates. Turner and Rosewater ( 1958 ) give a more complete discussion of the economic importance of Pinnidae; also see Calm (1951); Salis von Marschlins (1795); Simmonds (1879); Yates (1843); Gilroy (1845); Haas (1955); Feen (1949), and Reyne (1947). Commensalism The Pinnidae serve as hosts to a number of organisms, both internally as commensals and ex- ternally as holdfasts for barnacles, sessile bivalves, tube worms, algae and other forms of marine life. Aristotle and Pliny recorded the classic association of the small crab. Pinnotheres, which lives in the mantle cavity of Pinna. The two were supposed to live together in intimate friendship with the crab warning the bivalve of approaching danger and gaining refuge within the latter’s mantle cavity. According to Christensen and McDermott (1958), it is only the crab which receives any benefit from this association. Plate 135. Shell structure of Pinna carnea Gmelin (West- to show hexagonal shape; from same area of shell as in Fig. ern Atlantic). Fig. 1. Longitudinal section to show the 1 (both x 110; from Johnsonia, vol. 3, no. 38, pi. 151). length of crystals of the prismatic layer. Fig. 2. Cross section [53 - 502] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinnidae 177 Shrimps are also known to live in the mantle cav- ity of the Pinnidae. Turner and Rosewater (1957, p. 297) reported the observation of a shrimp in Pinna cornea from the Western Atlantic. Holtlmis (1952) reported several cases of shrimp-pinnid commensalism in the Indo-Pacific and they are mentioned below in the systematic portion of this study under the appropriate species. Remarks on Anatomy The usual habitat of Pinnidae is in a substrate of soft, sandy mud with the narrow, umbonal tip of the shell downward. They are sessile animals and, after becoming imbedded, there is probably little horizontal movement carried on, excursions being limited to vertical burrowing. As Yonge ( 1953 ) has stated, the Pinnidae are structurally well suited for the life they lead and are further aided in their adjustment by several unique anatomical features (see Plate 137). The gross anatomy of the Pinnidae is similar in the several genera with minor differences in pro- portions. The pallial organ is visible atop ( posterior and dorsal to) the posterior adductor muscle. It differs somewhat in shape among the genera. In Atrina, it has a thickened stalk and a conical to rounded head; in Pinna and Streptopinna, it is usu- ally thin and the head nearly aciculate-conical, de- pending on the degree of contraction. The fragile shells of the Pinnidae are easily broken posteriorly and the pallial organ functions in clearing away pieces of broken shell and debris from the mantle cavity. The organ is made turgid with body fluids and is thus able to move posteriorly. Intrinsic mus- cles cause lateral movements. With obstructions re- moved the extensible mantle may make necessary repairs to the shell. The pallial organ has an interesting nomencla- torial history. Poli (1795) called it a “trachea”; Rogers (1908) an osphradium; Grave (1911) the mantle gland; and Yonge ( 1953 ) the pallial organ. The last appears to be the best name for this struc- ture as it consists of differentiated parts made up of glandular and muscular tissue which are thought to have been derived from the pallium or mantle. Another useful and unique structure is the waste canal located ventral to the gills (plate 137). Be- cause of the vertical habit of the Pinnidae, consid- erable material might enter and accumulate in the mantle cavity were it not for this structure. Its function is the removal of rejected food and debris from the anterior portion of the cavity. The canals consist of two open, ciliated gutters, one on either half of the mantle, which produce a strong current carrying material rapidly to the posterior border where it is caught in the exhalent respiratory-feed- ing discharge and so expelled. Another unusual set of structures whose function is not yet well understood is present in the mantle margin of members of the genera Pinna and Strep- topinna. Will (1844) was probably the first to re- cord their presence, calling them eyes. They are located between the middle and inner mantle lobes and occur in greatest numbers posteriorly, becom- ing widely spaced anteriorly. Rawitz ( 1890, pp. 64- 72, pi. 4, figs. 28, 29) declared they were not eyes but glandular in nature. Sections kindly prepared by C. E. Cutress, Division of Marine Invertebrates, United States National Museum, show what appear to be secretory granules, so that a visual function probably does not exist. The eye-like structures of Will have been noted in the following species of Pinnidae: P. bicolor, P. muricata, P. cornea and Streptopinna saccata. Their comparative appear- ance is commented upon in descriptions of the spe- cies studied in this report. None have been seen in any specimens examined of the genus Atrina. The real purpose and function of these structures has not yet been discovered (see Patten, 1886, pp. 606, 607; Braun, 1954). Winckworth (1929, p. 282) called attention to them in S. saccata, referring to them as pigment spots. In connection with a possi- ble excretory pigment-secreting ability, it is here tentatively suggested that these glands may pro- duce the colored rays which are found in the shells of certain species in this group (see Comfort, 1951 ). The relative size and degree of pigmentation of the glands appear to vary in the several species and may be correlated with the degree of shell pigmen- tation. List of Recognized Taxa The following list contains the names of the genera and subgenera of all living and fossil Pinni- dae. All living species, which we consider valid, together with the Tertiary fossils from the Indo- Pacific are also included. Brackets [ ] indicate other oceans; daggers f are fossil genera and species; E.A. is Eastern Atlantic; W.A. is Western Atlantic; E.P. is Eastern Pacific. Genus f Palaeopinna Hall, 1870 [\flabella Hall, 1884] Type? Devonian Genus fS nlcatipinna Hyatt, 1892 [f flexicostata (McCoy, 1844)] Type. Carbon- iferous [53 - 503] 178 Taxonomic Characters J. Rosewater Pinnidae Genus f Laevipinna Paul, 1941 [t spatula (McCoy, 1853)] Type. Carboniferous Genus f Aviculipinna Meek, 1864 [t prisca (Muenster, 1837)] Type. Permian Genus f Stegoconcha Boehm, 1907 [f granulata (Sowerby, 1822)] Type? Jurassic Genus f Tricliites Deshayes, 1832 [f nodosus (Lycett, 1850)] Type. Jurassic-Creta- ceous Genus f Pinnigena Bronn, 1836 [f ampla (Sowerby, 1812)] Type? Jurassic-Creta- ceous Genus f Oxysma Rafinesque, 1819 [ [bifida Rafinesque, 1819] Type. Tertiary? Genus f Curvulites Rafinesque, 1831 (New name for Curvula Raf., 1819) [f striata Rafinesque, 1831] Type. Tertiary? Genus Pinna Linne, 1758 f punjabensis Eames, 1951 f rembangensis Martin, 1910 f asakuraensis Nagao, 1928 muricata Linne, 1758 f blanfordi Boettger, 1880 bicolor Gmelin, 1791 incurva Gmelin, 1791 [ nobilis Linne, 1758] E.A. [ruclis Linne, 1758] Type. E.A., W.A. [cornea Gmelin, 1791] W.A. [ rugosa Sowerby, 1835] E.P. Genus Atrina Gray, 1842 Subgenus Atrina s.s. Gray, 1842 f pachyostraca (Davies, 1923) vexillum (Born, 1778) Type. squamifera (Sowerby, 1835) South Africa [ chautardi (Nickles, 1953)] West Africa [fragilis (Pennant, 1777)] E.A. [rigida (Solander, 1786)] W.A. [tuberculosa (Sowerby, 1835)] E.P. Subgenus S ervatrina Iredale, 1939 pectinata (Linne, 1767) Type. subsp. zelandica (Gray, 1835) New Zealand f cordata (Pritchard, 1895) f janjukiensis (Crespin, 1950) f tateana (Hedley, 1924) tasmanica (Tenison-Woods, 1875) [seminuda (Lamarck, 1819)] W.A. [serrata (Sowerby, 1825)] W.A. [maura (Sowerby, 1835)] E.P. Genus Streptopinna von Martens, 1880 ?t reticosa (Chapman, 1912) saccata (Linne, 1758) Taxonomic Characters of the Pinnidae Early workers depended wholly on external de- tails of the shells to distinguish species in the Pin- nidae. Although external shell form is very useful, it is so subject to change by the environment that other details of the soft anatomy and the internal shell must be used for positive identification. In most instances adult specimens must be used. Iden- tification to species is based on a combination of characters which include the general shape and texture of the shell, the comparative numbers of ribs, the development of the spines, the contour and extent of the nacreous layer and, when avail- able, the details of certain parts of the soft anatomy. As more anatomical information becomes avail- able, it may well become the main basis for precise specific identification. The present methods of pres- ervation of the animals produce a variety of effects Plate 136. Diagrammatic sketch of the valves of the genera ters. Figs. 1-2. External and Internal surfaces ot the valves Atrina, Pinna and Streptopinna to show diagnostic charac- of Atrina. Figs. 3-4. The same of the valves of Pinna. Figs. [53 - 504] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinnidae 179 such as shrinkage and hardening so that apparent differences in the proportions and shapes of various organs cannot always be considered reliable. Live animals must be relaxed in some effective way, killed rapidly in a substance which fixes the soft parts in the relaxed state, and then preserved in a standard medium. An excellent procedure for some bivalves which produces very pliable specimens easily manipulated for study is the following: relax the living animal in Propylene Phenoxetol; fix in buffered formalin; then preserve in Propylene Phenoxetol (Turner, 1960, p. 7). Another method utilizes 10% Nembutal as a relaxant; A.F.A. ( Lavandowsky’s mixture of alcohol, formalin, and acetic acid) as a fixing me- dium and 70% alcohol for preservation. The latter also produces fairly well relaxed specimens, al- though not so pliable as those preserved in Propyl- ene Phenoxetol. (See Van der Schalie, 1953, for in- structions on the use of Nembutal as a relaxant. ) These methods are relatively simple, easy to learn, and many times increase the taxonomic value of material collected during costly expeditions. In spite of the difficulty of obtaining well-pre- served material, some anatomical work has been done in the Pinnidae which may be of use in tax- onomy. Purchon ( 1957 ) found the stomachs of Atrina vexillum ( Born ) and Pinna atropurpurea [= P. bicolor Gmelin] to differ in some details, the latter being more complicated. As no other work has been done on this organ in other species of Pin- nidae it is not known whether these differences should be considered of generic or specific value. Various other characters, such as the curvature and position of the waste canal, the shape of the pallial organ and coloration of internal organs, have served to discriminate species of Pinnidae ( see Turner and Rosewater, 1958, p. 300). Color of the organs is of little value unless the specimens can be seen alive, since they tend to fade in preservation. Notes taken on a single species, Pinna bicolor, by D. F. Mc- Miehael, Australian Museum, are the only data of this type available in the present study. They con- cern the coloration of the ovary of a female speci- men (see description of bicolor). Nothing is known of the coloration of the male organ. Winckworth (1929) expressed confidence in de- termining species by means of correlating the num- ber of gill filaments per plica with the size of the apical filaments (see plate 138). This character was described by Ridewood ( 1903 ) who gives counts for several species of Pinna and Atrina. Ridewood found that Pinna pectinata [= A. fragilis (Pen.)] possessed 16-17 filaments per plica. This count was verified by Winckworth. The other species exam- ined by Ridewood ( Pinna nobilis ; Pinna nigra [= Atrina vexillum?]-. Pinna zealandica [error for Atrina zelandica] and Pinna virgata [ Pinna muri- cata?] ) all possessed filament counts from 10 to 13. Differences were noted in the sizes of the apical filaments. In fragilis, nobilis and bicolor the apical filament was found to be much larger than the ordinary filaments, but in vexillum it was of about the same size or only slightly larger. In our present study the gills of seven species of Pinnidae have been examined grossly by means of a binocular dis- secting microscope (P. bicolor; P. muricata; P. car- nea; A. vexillum; A. pectinata; A. seminuda ; and S. saccata). It was noted in all specimens examined that the number of ordinary gill filaments was about 12 (6 on either side of a plica). In addition there is one principal and one apical filament ( see plate 138). Differences were noted in the relative sizes of the apical and ordinary filaments and prob- ably in the shapes of the plicae. Unfortunately these specimens were preserved in several different ways, some obviously having undergone violent contraction and shrinkage of the gills. For this rea- son any attempt to compare the gill anatomy of these species will be deferred until properly relaxed and preserved specimens are available. In the final analysis a combination of as many taxonomic characters as possible should be utilized in separating species. Unfortunately, however, in the usual museum mollusk collection only shells are found. For this reason, in the following key to spe- cies of Indo-Pacific Pinnidae, the characteristics of the shells are stressed (see pi. 136 for orientation in shell morphology ) . [53 - 505] 180 Key to Pinnidae J. Rosewater Pinnidae Key to the Recent Indo-Pacific Pinnidae 1. Internal nacreous layer divided by a longitudinal sulcus Pinna (3) Internal nacreous layer not divided (2) 2. Nacreous layer occupying entire interior of anterior portion of valves Atrina (5) Nacreous layer limited to dorsal anterior portion of valves only; shell usually misshapen and contorted Streptopinna saccata 3. Posterior shell-margin squarely truncate; posterior adductor scar often extending onto ven- tral lobe of nacreous layer Pinna muricata Posterior shell-margin not squarely truncate but arcuate or broadly rounded; posterior ad- ductor scar never extending onto ventral lobe (4) 4. Posterior nacreous borders both sharply oblique, forming a deep medial “V”; posterior extension of dorsal and ventral lobes about equal; shell long and narrow, thin and fragile; color light-horn to light reddish brown Pinna incurva Posterior nacreous borders not both sharply oblique, not forming deep “V”; the ventral usu- ally oblique, the dorsal truncate; posterior extension of dorsal and ventral lobes sub- equal, the ventral may be longer or shorter; shell often thickly produced, not long and narrow; light-horn to dark brownish purple, often colorfully patterned . . Pinna bicolor 5. Posterior adductor muscle scar protruding beyond posterior border of nacreous layer or contiguous with it Atrina s.s. (6) Posterior adductor scar not protruding; located well within posterior border of nacreous layer Servatrina (7) 6. Shell reaching large size, thick, heavy, black, broadly ham-shaped; early growth showing prickly major ribs which often have 2-3 rows of minor ribs between them. Atrina (Atrina) vexillum Shell not especially large, thick or heavy, sculptured with large, erect, nearly tubular spines; without minor ribs (South Africa) Atrina (Atrina) squamifera 7. Shell olivaceous to black, or with few color stripes; rather thin; with 14-30 rows of prickly to spiny ribs; posterior margin truncate (8) Shell reddish brown (olivaceous when young); moderately heavy; ribs usually less than 14; posterior margin rounded (southeastern Australia and northern Tasmania). Atrina (Servatrina) tasmania 8. Sculpture reduced to few scattered spines or low uniform imbrications; color olivaceous tan to black ( India to Melanesia but not southeastern Australia, Tasmania or New Zealand ). Atrina (Servatrina) pectinata Sculpture not obsolete but rather uniformly spinose; color olivaceous tan, often with verti- cal bars of brownish purple (New Zealand) Atrina (Servatrina) zelandica Geographic Distribution of Recent Species World-wide Distribution — The geologic history of the family Pinnidae, which began in the late Paleozoic, is largely unknown because of the poor preservation of specimens. Inferences must be made largely from today’s distribution of living forms. The family probably has a history of diverse genera and species which once flourished but have since disappeared without fossil traces. It also seems likely that certain conservative stocks, such as the Carboniferous Atrina, have survived to Recent times without appreciable change, much in the same manner as members of the brachiopod genus. Lingula, and the eurypterid-like horseshoe crab, Limulus. In the main, the family has remained in shallow seas of a tropical and warm temperate nature, rarely having representatives in cooler waters. Because of the pelagic nature of the larval stages and the ease of transportation of young individuals which have attached themselves to floatable objects, the pen shells have had ample time and occasion to reach [53 - 506] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinnidae 181 comb pallial muscles ventral polliol retractor rl or adductor muscle poslerior pedal retractor orsal palliol retractor anterior pedal retractor a'nterior adductor muscle Plate 137. Semidiagrammatie drawing of the anatomy of Atrina. The right half shows only the muscles and character- istic features of the mantle (from Tohnsonia, vol. 3, no. 38, pi. 152). all parts of the world’s oceans. Today’s distribu- tions are a reflection of those of former species, some of which may have had wider, others smaller, ranges. Despite the world-wide distribution of Atrina, Servatrina and Pinna throughout today’s distinct faunal regions, there is a remarkable similarity in the morphological characters among the species of each group. Some of the Recent species probably evolved from forms which had continuous distribu- tions during the Mesozoic or possibly the early Cenozoic. One genus, S treptopinna, is limited to the tropical Indo-Pacific and appears to be a more recent development, probably during the late Ter- tiary. The gastropod genus Lambis (Strombidae) and the pelecypod genus Triclacna, both known only from the Indo-Pacific, have analogous histories. Within the genus Pinna there are two types of species groups, those which are limited to one province, such as the Indo-Pacific, and those which extend across two or more provinces either as sub- species or very closely related allopatric species. To the former group belong P. rnuricata (widespread in the Indo-Pacific only), bicolor (Indian Ocean, Western Pacific and Hawaii), and incurva (India to northern Australia). They appear to have no ana- logues in other Recent seas. Of the pandemic groups of Pinna, species which extend through several faunal provinces, one is P. rudis Linne of the tropical Eastern Atlantic and Caribbean and its Eastern Pacific analogue, P. ru- gosa Sowerby. The latter may well be considered by some workers as a geographic subspecies of rudis. Another example is the Eastern Atlantic P. nobilis Linne and its Western Atlantic analogue, P. cornea Gmelin. These two species are certainly [53-507] 182 Embryonic Valves J. Rosewater Pinnidae specifically distinct but show marked similarities in details of the nacreous pattern. Members of the subgenus S ervatrina of the genus Atrina in distant parts of the world may closely re- semble each other. The shells of A. seminuda (La- marck) and A. serrata (Sowerby) of the Western Atlantic are almost inseparable from some speci- mens of A. pectinata (Linne) of the Indo-Pacific. It is of interest to note that while seminuda and serrata are easily separable in the Western Atlan- tic, phenotypes somewhat resembling both Atlantic species appear to intergrade in pectinata in the Indo-Pacific. Young specimens of Atrina mama (Sowerby) of the Eastern Pacific also resemble seminuda, serrata and pectinata but become more lamellosely sculptured as adults and quite distinct. Atrina fragilis Pennant of Europe and A. chau- tardi (Nickles) of West Africa are apparently the only members of Atrina s.s. present in the Eastern Atlantic. The species are quite convergent and both somewhat resemble A. squamifera (Sowerby) of South Africa. The three may have had a continuous distribution in the past (see Remarks under A. squamifera) . There is a greater degree of spinosity in squamifera and chautardi than in fragilis but the latter appears to attain a larger size. Specimens of chautardi have not been available for comparison; however, the nacreous patterns of the other two are similar. Plate 138. Diagrammatic sketch of gill of Pinna. Vertical section of outer demibranch, one-half plica in thickness (looking down on long axis of gill; after Ridewood, 1903, fig. 17). Atrina rigida (Solander) of the Western Atlantic may be considered the analogue of A. tuberculosa ( Sowerby ) of the Eastern Pacific, although the shell of the latter becomes larger and heavier. Both are closely related to A. vexillum (Born), a widely distributed Indo-Pacific species. All three are heavy, with dark pigmentation, either spinose or smooth, and exhibit a similar nacreous pattern with very large posteriorly protruding muscle scars. Certain features peculiar to each of these species, undoubt- edly brought about through isolation, serve to dis- tinguish them. The world distribution of living Pinnidae is sum- marized in Table 1. Plate 139. Embryonic valves of Indo-Pacific Pinnidae. Figs. 1-2. Pinna muricata Linne from Long Reef, New South Wales, Australia (0.5 mm. in length). Figs. 3-4. Atrina vexillum (Born), from Auau Channel, Hawaii (21-28 fathoms) (0.5 mm. in length). Figs. 5-6. Atrina pectinata (Linne), from Nagasaki, Japan (107 fathoms) (1.0 mm. in length ). [53 - 508] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinnidae 183 Table 1. World Distribution of Living Pinnidae Indo-Pacific East Pacific West Atlantic East Atlantic South Africa muricata rugosa ruclis rudis Pinna bicolor cornea nobilis incurva Atrina vexillum tuberculosa rigid a fragilis squamifera chant arch Servatrina pectinata zelandica maura serrata seminuda tasmanica Streptopinna saccata Origin and Distribution of Indo-Pacifc Pinnidae — It is probably impossible to determine the place of origin of the family Pinnidae because it is an old group whose most ancient ancestors have never been found. On the basis of the distribution of Re- cent Indo-Pacific species, it would appear that the Western Pacific might have been the place of ori- gin, as it is here that two-thirds of the species are found. However, this information may be mislead- ing. Now, and undoubtedly in the past, the largest number of species in most groups have thrived where their particular ecological requirements are met most efficiently. Given time, any species having a pelagic larva can probably reach any location in the oceans. However, conditions must be ideal for these migrants to become established. This, of course, limits many species which have narrow eco- logical requirements. Arguments of Ladd (1960) and Abbott (1960) respectively suggest a nearly mid-Pacific and ex- treme western Pacific origin for several groups of mollusks. It is probable that, through time, both areas have served this purpose. In recent geologic time the mid-Pacific islands have been nearly inun- dated, and rich mineral and organic constituents are lacking which could supply the necessary eco- logical requirements for some species. The western Pacific is a rich area ecologically and probably has been so for a very long time, supplying an abun- dance of nutrient materials for the food chain. This may explain the accumulation of species in the lat- ter area, as it no doubt attracts and holds species which arrive there. This does not eliminate more easterly islands as places of origin since, in the past, there is evidence that these also supported a large fauna. In the case of the Pinnidae, those species whose ranges extend from Africa to Polynesia (P. muri- cata; A. vexillum; and S treptopinna saccata) proba- bly have sufficiently broad ecological tolerances permitting them to live not only in rich continental seas but also on the shores of coral islands. Pinna muricata , however, seems to grow most rapidly and attain a larger size in the western Pacific ( Mariveles Bay, Philippines). The rest of the Indo-Pacific Pin- nidae are limited to continental areas or the larger islands. Pinna bicolor Gmelin is limited to the western Pacific and Indian Oceans. It is the only Pinna which is found in Victoria and South Australia, no other member of the genus extending any farther south than New South Wales. It inhabits the high islands from southern Japan to New Caledonia but does not inhabit the low coral atolls of the mid- Pacific. It is found westward from the high islands into the Indian Ocean to the east coast of Africa and the Red Sea. It was recently found in Hawaii. Young specimens were, perhaps accidentally, trans- ported there, and it is doubtful whether or not the species will survive in Hawaiian waters. The scanty locality data for Pinna incurva Gmelin, from the literature and specimens exam- ined, indicate that its range is relatively narrow in southeast Asia, extending from India to Queens- land, Australia, and north to the Philippines. Species of Atrina, other than the widespread vex- illum, have more limited ranges. Atrina pectinata (Linne) is the most ubiquitous of these remaining species, its range extending from the coast of India to Queensland and north to southern Japan. This species evidently has not invaded the low atolls of the mid-Pacific. Atrina pectinata zelandica (Gray) is limited in its distribution to New Zealand; A. tasmanica (Teni- [53 - 509] 184 Doubtful Species J. Rosewater Pinnidae son-Woods) to Tasmania, Victoria, South Australia and New South Wales; A. squamifera (Sowerby) to South Africa. It is of interest to note that these last three species have extended beyond the warm Indo-Pacific faunal provinces into cooler waters and occupy widely separated geographic positions. Al- though not properly members of the Indo-Pacific fauna, they are included here to complete the fam- ily for the area. Doubtful Species of Pinnidae Most of the specific names in the Pinnidae were treated in Winckworth’s catalogues (1929, 1936). He cited several nude names but the following were not included. 1931 Pinna striolata ‘Turton’ Sherborn, Index Ani- malium, section 2, part 25, p. 6204 [nomen nudum]; error for Anomia striolata Turton, 1822. 1933 Pinna italicus Sherborn, ibid., part 32, p. 842 [nomen nudum]. The doubtful species listed below in some cases provide additional information to that given by Winckworth. Hedley (1924) also listed erroneous species recorded from Australia. Pinna rotundata Linne, 1758 Range — Mediterranean? Remarks — Hanley (1855) and Winckworth ( 1929) considered rotundata unrecognizable; Dodge ( 1952 ) felt it suggested Pinna nobilis Linne of the Mediterranean. Gualtieri’s plate 79, fig. C, to which Linne referred is too vague for certain placement of this species. The immature shell figured could be that of any young Pinna. Synonymy — 1758 Pinna rotundata Linne, Systema Naturae, ed. 10, p. 707 (O. meridionali ) ; refers to Gualtieri, pi. 79, fig. C. Pinna digitiformis Linne, 1758 Range — Indo-Pacific? Remarks — Hanley (1855, p. 153) supposed this to be a pteropod; Winckworth ( 1929, p. 291 ) guessed it was a brachiopod. I concur with Dodge (1952, p. 230) that the species is unrecognizable and should be eliminated from further conjecture. Synonymy — 1758 Pinna digitiformis Linne, Systema Naturae, ed. 10, p. 708 (O. Indico). Pinna lobata Linne, 1758 Range — Indo-Pacific? Remarks — As in the case of P. digitiformis it is doubtful whether P. lobata is a bivalve. The spe- cies is unrecognizable (see Hanley, 1955; Winck- worth, 1929; Dodge, 1952). Synonymy — 1758 Pinna lobata Linne, Systema Naturae, ed. 10, p. 708 (O. Indico). Pinna pennaeea Linne, 1758 Range — Unknown. Remarks — This is the pen of a cuttlefish ( Cepha- lopoda). It was first described as a Pinna with reser- vations, but removed from this group (Linne, 1767) and placed in Sepia (see Hanley, Winckworth and Dodge ) . Synonymy — 1758 Pinna pennaeea Linne, Systema Naturae, ed. 10, p. 708 (no locality); 1767, ibid., ed. 12, p. 1090. Pinna sanguinea Gmelin, 1791 Range — Unknown. Remarks — Gmelin himself questioned the valid- ity of this species. Winckworth (1929, p. 295) sug- gested that P. sanguinea might be P. nobilis Linne of the Mediterranean. However, I agree with him that Gualtieri’s pi. 79, fig. B, which was cited by Gmelin, is a doubtful figure, and I consider san- guinea unrecognizable. Synonymy — 1791 Pinna sanguinea Gmelin, Systema Naturae, ed. 13, p. 3367 ( no locality given ) . Pinna bullata Gmelin, 1791 Range — Unknown. Remarks — The plate reference given for this spe- cies by Gmelin was Gualtieri, pi. 79, fig. C, which was also cited for P. rotundata Linne. In agreement with Gmelin (“an distincta a reliquis species?’’), Hanley (1855) and Winckworth (1929) considered P. bullata as unrecognizable. Reeve’s ( 1858, pi. 9, fig. 16) figure is probably P. bicolor from the Mo- luccas. Synonymy — 1791 Pinna bullata Gmelin, Systema Naturae, ed. 13, p. 3367 (no locality given); refers to Gualtieri, pi. 79, fig- C. [53 - 510] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinnidae 185 Pinna cleflecta Perry, 1811 Range — Unknown; Indo-Pacific? Remarks — The figure in Perry appears doubtful and could represent more than one Indo-Pacific Pinna. The species is unrecognizable. Synonymy — 1811 Pinna deflecta Perry, Conehology, pi. 61, Pinna, fig. 2 ( no locality given ) . Pinna marginata Lamarck, 1819 Range — Unknown. Remarks — Lamarck had no specimens of margi- nata, but referred to the same plate in Gualtieri (pi. 79, fig. C) cited for P. rotundata Linne and P. bullata Gmelin. Winckworth (1929) allied margi- nata with bullata Gmelin and called the latter a synonym of rotundata. Pinna rotundata was un- identified at that time and remains so. The species is unrecognizable. Synonymy — 1819 Pinna marginata Lamarck, Histoire Naturelle des Ani- maux sans Vertebres, vol. 6, part 1, p. 132 (no lo- cality given); refers to Gualtieri, pi. 79, fig. C, and to Pinna bullata Gmelin. Pinna sanguinolenta Reeve, 1859 Range — Unknown; Indo-Pacific? Remarks — The outward appearance and red col- oration of this species are reminiscent of P. carnea Gmelin. A photograph of the valve of the type in the British Museum (N.H.) shows an immature specimen with a nacreous pattern similar to that of P. bicolor. Because of the lack of locality data it seems best to leave this species in a doubtful cate- gory until proof is available of its identity. Synonymy — 1859 Pinna sanguinolenta Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 33, fig. 62 (no locality given). Acknowledgments Many individuals and institutions generously pro- vided photographs of type specimens, loaned mate- rial and supplied information to make this report more complete. Their help is gratefully acknowl- edged and credit given in appropriate places throughout the text. Thanks are due the following for helpful criticism and suggestions which aided this study greatly: Harakl A. Rehder and J. P. E. Morrison, USNM; Harry S. Ladd, USGS; R. Tucker Abbott and Robert Robertson, ANSP; William J. Clench and Ruth D. Turner, MCZ. The author has obtained permission to publish this manuscript from the Secretary of the Smith- sonian Institution. Financial support for the research involved was generously given by the Atomic En- ergy Commission under contract no. AT (30-1 )-2409. Bibliography Most references to the taxonomy of the Pinnidae will be found in the synonymies of the genera and species. Others from the text and those quoted for locality records may be found by consulting the Zoological Record for the year mentioned. The more recent and important references, those con- cerning biology, and others which might be diffi- cult to find, are listed below. Abbott, R. Tucker. 1960. The Genus S trombus in the Indo- Pacific. Indo-Pacific Mollusca, vol. 1, no. 2, p. 44 [09- 842], Calm, A. R. 1951. Clam Culture in Japan. Report No. 146, Natural Resources Section, General Headquarters Supreme Commander for the Allied Powers, pp. 1-103. Comfort, Alex. 1951. The Pigmentation of Molluscan Shells. Biological Reviews, vol. 26, pp. 285-301. Feen, P. J. v. d. (Jutting, W. S. S. v. B.). 1949. Byssus. Basteria, vol. 13, no. 4, pp. 66-71. Gabriel, C. J. 1936. Victorian Sea Shells. Handbook, Field Naturalists’ Club of Victoria. Brown, Prior, Anderson Pty. Ltd., Melbourne. 67 pp., 6 pis., text figs. Gilroy, G. C. 1845. The history of Silk, Cotton, Linen, Wool and other Fibrous Substances. Harper and Brothers, New York, 464 pp. Grave, B. H. 1909 [1911]. Anatomy and Physiology of the Wing-shell, Atrina rigida. Bull. Lb S. Bureau Fisheries, no. 29, Document 744, pp. 409-439, 3 pis. Gregoire, C. 1960. Further Studies on Structure of the Or- ganic Components in Mother-of-Pearl, especially in Pele- cypods ( Part 1 ) . Bull. Institut royal des Sciences na- turelles de Belgique, vol. 36, no. 23, Brussels, pp. 1-22, 5 pi. Haas, Fritz. 1955. Natural History of the Pearls. Comunica- ciones del Instituto Tropical de Investigaciones Cientificas de la Universidad de El Salvador, ( San Salvador, Centro America) Ano 4, nos. 3/4, pp. 113-126. Habe, T. 1953a. Pinnidae, Placunidae and Anomiidae in Japan. Illustrated Cat. Japanese Shells, vol. 1, no. 24, pp. 185-200. 1953b. Notes on the Type Specimens of Three Spe- cies of Japanese Molluscs Described by F. Jousseaume. Publ. Seto Marine Biol. Lab., vol. 3, (2), pp. 169-172, pi. 9, figs. 1-6. Hanley, Sylvanus. 1855. Ipsa Linnaei Conchylia, pp. 1-556, 5 pis. [53-511] 186 Bibliography J. Rosewater Pinnidae Holthuis, Dr. L. B. 1952. The Decapoda of the Siboga Ex- pedition, Part XI., The Palaemonidae. Monograph 39a10 Siboga-Expeditie, pp. 1-253, 110 figs. Iredale, T. 1929. Queensland Molluscan Notes. Memoirs Queensland Museum, Brisbane, vol. 9, p. 281. Jackson, R. T. 1890. Phylogeny of the Pelecypoda, the Avi- culidae and Their Allies. Memoirs Boston Society of Nat- ural History, vol. 4, no. 8, pp. 277-400, 8 pis., 53 text figs. Jameson, H. Lyster. 1902. On the Origin of Pearls. Proceed- ings Zoological Society of London, pp. 140-166, 4 pis. Kawaguti, Siro and Nariaki Ikemoto. 1957. Electron Micros- copy of the Smooth Muscle from the Adductor of Span- ish Oysters, Pinna attenuata and Atrina japonica. Biol. Journ. Okayama Univ., vol. 3, no. 4, pp. 248-268. Keen, A. Myra. 1958. Sea Shells of Tropical West America. Stanford University Press, Stanford, California, xii + 624 pp. Kemp, S. 1922. Pontoniinae. Notes on Crustacea Decapoda in the Indian Museum, XV. Records Indian Museum, vol. 24, pp. 113-288, text figs. 1-105, pis. 3-9. Kolm, A. J. 1956. Problems of Speciation in Marine Inverte- brates. Perspectives in Marine Biology. A Symposium, Scripps, pp. 571-588. Kubo, Ituo. 1940. Studies on Japanese Palaemonoid Shrimps. Journ. Imperial Fisheries Institute, vol. 34 (1), pp. 1-75. Ladd, H. S. 1960. Origin of the Pacific Island Molluscan Fauna. American Journal of Science, Bradley Volume, vol. 258-A, pp. 137-150. Lamy, E. 1938. Pinna de l’lndochine. Bulletin Museum D’Histoire Naturelle, Paris, (2) vol. 10, no. 4, pp. 396-400. Lister, M. 1685. Historiae sive Synopsis Methodieae Con- chyliorum. London. Nakazima, Masao. 1958. On the Differentiation of Stomach of Pelecypoda (I.). Venus, vol. 20, no. 2, pp. 197-207. Olsson, A. A. 1961. Panamic-Pacific Pelecypoda Paleonto- logical Research Institution, Ithaca, N. Y., 574 pp. [Pin- nidae: pp. 143-145]. Ota, Shigeru. 1961. Identification of the Larva of Pinna (Atrina) japonica (Reeve). Bull. Japanese Soc. Sci. Fish., vol. 27, no. 2, pp. 107-112, 3 figs., 1 pi. Patten, William. 1886. Eyes of Molluscs and Arthropods. Mittheilungen aus der Zoologischen Station zu Neapel, vol. 6, part 4, pp. 542-756, pis. 28-32. Purchon, R. D. 1957. The Stomach in the Filibranchia and Pseudolamellibranchia. Proceedings Zoological Society, London, vol. 129, part 1, pp. 27-60. Rawitz, Bernhard. 1890. Der Mantelrand der Acephalcn, Zweiter Teil, Arcacea, Mytilacea, Unionacea. Jenaischen Zeitschrift fur Naturwissenschaft, vol. 24, N. F. 27, pp. 1- 83, 4 pi. Reyne, A. 1947. On the structure of the shells and pearls of Tridacna squamosa Lam. and Hippopus hippopus (Linn.). Arch. Neerlandaises Zook, vol. 8, liv. 1, 2, p. 208 (foot- note ) . Ridewood, W. G. 1903. On the Structure of the Gills of the Lamellibranehia. Philosophical Transactions of the Royal Society of London, series B, vol. 195, pp. 147-284. Rogers, J. E. 1951. The Shell Book. Branford Co., Boston, new edition, pp. xxi + 503 pp. (p. 401). Salis von Marschlins, C. U. v. 1795. Travels through various Provinces of the Kingdom of Naples, in 1789; translated from the German by Anthony Aufrere. London, viii + 527 pp. Simmonds, P. L. 1879. The Commercial Products of the Sea. Griffith and Farran, London, pp. 306-310 (viii + 484 pp.). Smith, E. A. 1903. Marine Mollusca: In: Fauna and Geog- raphy of the Maidive and Laccadive Archipelagoes, vol. 2, part 2, pp. 589-630, pis. 35, 36. Edited by J. S. Gard- iner, Cambridge, England. 1906. On South African Marine Mollusca. Annals Natal Government Museum, vol. 1, part 1, pp. 60-61. Suter, H. 1913. Manual of the New Zealand Mollusca, with an Atlas of quarto plates. Wellington, xxiii + 1 120 pp. 1914. Revision of the Tertiary Mollusca of New Zea- land, part I. New Zealand Geological Survey Palaeonto- logical Bulletin no. 2, 64 pp., 17 pis. 1915. Ibid., part II. New Zealand Geological Survey Palaeontological Bulletin no. 3, 69 pp., 9 pis. Tesch, P. 1920. Jungtertiare und quartare Mollusken von Timor. In: J. Wanner, Palaontologie von Timor, Stuttgart, Lief 8, part 2, p. 91. Turner, R. D. 1960. Some Techniques for Anatomical Work. Annual Reports for 1959, The American Malacological Union, Bulletin no. 26, pp. 6-8. Turner, R. D. and Joseph Rosewater. 1958. The Family Pin- nidae in the Western Atlantic. Johnsonia, vol. 3, no. 38, pp. 285-326. Van der Schalie, Henry. 1953. Nembutal as a Relaxing Agent for Mollusks. American Midland Naturalist, vol. 50, no. 2, pp. 51 1 -512. Van der VIerk, I. M. 1931. Caenozoic Amphineura, Gastro- poda, Lamellibranchiata, Scaphopoda. Leidsche Geo- logische Mededeelingen, Leiden, deel 5, pp. 206-296. Vokes, II. E. [1951] Preliminary Classification of the Genera of the Pelecypoda. ( An unpublished mimeographed com- pilation. ) Wada, Koji. 1961. On the Relationship between Shell Growth and Crystal Arrangement of Nacre in Some Pelecypoda. Venus, Fukuyama, vol. 21, no. 2, pp. 204-211, 4 figs. Wilkins, G. L. 1953. Notes from the British Museum, I. Pinna. Proceedings Malacological Society of London, vol. 30, pp. 23-29, 1 pi. Will, J. G. F. 1844. LTeber die Augen der Bivalven und der Ascidien. Froriep’s Neue Notizen aus dem Gebiete der Natur- und Heilkunde, vol. 29, no. 622, pp. 80-87; no. 623, pp. 99-103. Winckworth, R. 1929. Marine Mollusca from South India and Ceylon, III. Pinna. With an Index to the Recent Spe- cies of Pinna. Proceedings Malacological Society of Lon- don, vol. 18, pp. 276-297. 1936. Ibid., X. Further Notes on Pinna. Ibid., vol. 22., pp. 20-23. Yates, J. 1843. Textrinum Antiquorum. Taylor and Walton, London, pp. 152-159. Yonge, C. M. 1953. Form and Habit in Pinna carnea Gmelin. Philosophical Transactions Royal Society of London, series B, vol. 237, pp. 335-374. Yoshida, H. 1956. Early Life History of Useful Bivalves in the Ariake Sea (I). Journal Shimonoseki College of Fish- eries, vol. 6, pp. 115-122, 4 figs. [53 - 512] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 187 Genus Pinna Linne, 1758 Type: Pinna rudis Linne, 1758 This genus includes all of the pen-shells which have the internal nacreous layer divided into two portions by a longitudinal sulcus. Its species are world-wide in distribution in tropical and subtropi- cal seas. Several generic names have been proposed for apparently different morphological and geographi- cal groups of Pinna. They are here considered syno- nyms of Pinna. The descriptions of the last three (see synonymy of Pinna) were prompted by the great degree of ecologically-caused phenotypic variation shown by the species of this group. Quantulopinna, Sabitopinna and Exitopinna, all Iredale, 1939, are “geographical genera” erected for the Australian forms of widespread Indo-Pacific species. Their types are assignable respectively to an arrested specimen of P. muricata ; a “normal” P. bicolor and an abnormal P. bicolor (“ Exitopinna deltodes ultra" Iredale). Cyrtopinna Morch is mono- typic for Pinna incurva Gmelin, an attenuate spe- cies from southeast Asia and, although this species is quite distinct, it does not warrant treatment in a separate genus with present available information. Three Recent and four Tertiary species from the Indo-Pacific are considered here. Description — Shell reaching a large size, up to 700 mm. (about 28 inches) in length in some spe- cies; wedge-shaped, generally fragile in structure and sculptured with radiating ribs which may bear spines or imbrications. Nacreous layer divided by a longitudinal sulcus into dorsal and ventral lobes. Posterior adductor muscle scar usually completely enclosed within dorsal lobe. Synonymy — 1758 Pinna Linne, Systema Naturae, ed. 10, p. 707. Type by subsequent selection (Children, 1823; also see Eames, 1951, Phil. Trans. R. Soe., London, vol. 235 (B), p. 339; Turner and Rosewater, 1958, pp. 301- 303): Pinna rudis Linne, 1758. 1791 Chimaera Poli, Testacea Utriusque Siciliae, vol. 1, p. 31. Type by subsequent selection, (Winckworth, 1929): Pinna nobilis Linne. 1795 Chimaeroderma Poli, Testacea Utriusque Siciliae, vol. 2, p. 259. Type by subsequent selection ( Turner and Rosewater, 1958, p. 301): Pinna nobilis Linne. 1806 Pinnarius Dumeril, Zoologie Analytique, pp. 169, 340; Iredale, 1939, Great Barrier Reef Expedition Scien- tific Reports, vol. 5, p. 309 [new name for Pinna Linne], 1815 Pinnula Rafinesque, Analyse de la Nature, ou tableau de l’Univers et des Corps Organises, Palerme, p. 147 [a substitute name for Pinna Linne]. 1853 Cyrtopinna Morch, Catalogus Conchyliorum Comes de Yoldi, part 2, p. 51. Type by monotypy : Pinna in- curvata Chemnitz [= P. incurva Gmelin]. 1939 Quantulopinna Iredale, Great Barrier Reef Expedition Scientific Reports, vol. 5, Mollusca, part 1, p. 310. Type by original designation: Quantulopinna delsa Iredale, 1939 [= Pinna muricata Linne]. Plate 140. Figs. 1-5. Pinna asakuraensis Nagao. Hoshuyama- mura, Asakura-gun, Chikuzen, Kyushu, Japan. Eocene. Fig. 3 is holotype (all reduced %, from Nagao, 1928, pi. 8, figs. 16, 18, 20, 20a, 21). Fig. 6. Pinna rembangensis Martin. Rem- bang, Java. Miocene, (nat. size, from Martin, 1910, pi. 51, fig. 73). Figs. 7-8. Pinna punjabensis Eames, Punjab and Kohat, Pakistan. Eocene, x Vz. ( Fig. 8. is P. shekhanensis Eames = punjabensis ; from Eames, 1951, pi. 11, figs. 39a, 40). [53 - 533] 188 Pinna J. Rosewater Pinnidae 1939 Subitopinna Iredale, ibid., p. 312. Type by original designation: Pinna menkei Reeve [= P. bicolor Gmelin]. 1939 Exitopinna Iredale, ibid., p. 315. Type by original designation: Exitopinna deltodes ultra Iredale [= P. bicolor Gmelin]. Pinna asakuraensis Nagao, 1928 (PI. 140, figs. 1-5) Range — From the Tertiary of Japan, Lower Eo- cene, Island of Kyushu. Remarks — This species is a true Pinna and is apparently more closely related to P. muricata in sculptural characteristics than to the other Recent species of Pinnidae. As mentioned by Nagao, P. blanfordi Boettger is “more slender and differently sculptured” and is here considered closer to P. bi- color Gmelin. Synonymy — 1928 Pinna asakuraensis T. Nagao, Science Reports of To- hokn Imperial University, Sendai, Japan, series 2 (Geology) vol. 12, no. 1, p. 31, pi. 8, figs. 16, 18, 20-22 [21] (Kawamagari Beds; Doshi and Hoshu- yama Mines, Hoshuyama-mura, Asakura-gun, prov- ince of Chikuzen ) . Pinna rembangensis Martin, 1910 (PI. 140, fig. 6) Range — Lower Miocene of Rembang, and West- Progo beds, Java, Indonesia (Van der Vlerk, 1931); also, questionably, from the Pliocene of Karikal, India (Cossmann, 1924, p. 93). Remarks — From the appearance of the partial internal mold figured by Martin, rembangensis is closest in relationship to Pinna muricata Linne. It is impossible to assign with certainty the fragment figured by Cossman to this species. Synonymy — 1910 Pinna rembangensis K. Martin, Sammlongen Des Geol- ogischen Reichs-Musenms In Leiden, Neue Folge, Bd. 1, 2 Abteilung, Heft 2, p. 357, pi. 51, fig. 73 (Rembang [Java]). ?1924 Pinna cf. rembangensis Martin, Cossmann, Journal de Conchyliologie, vol. 68, p. 92, figs. 21-22 (Pliocene, Karikal ) . Pinna punjabensis Eames, 1951 (PI. 140, figs. 7-8) Range — From the Eocene of Pakistan. Remarks — Pinna pun jabensis has sculptural char- acteristics similar to Pinna muricata Linne and may be ancestral to that species. In spite of the differ- ences pointed out by Eames in his description of P. shekhanensis, also from the Pakistan Eocene, it is probably a synonym of punjabensis. The figure of the former (Eames, 1951, pi. 11, fig. 40) represents a fossil probably consisting of portions of two valves and when so considered is much like punjabensis. Synonymy — 1951 Pinna punjabensis Eames, Philosophical Transactions Royal Society of London, Series B, vol. 235, no. 627, p. 339, pi. 11, fig- 39 a, b (Ghazij Shales; south of Nila Kund, Dera Ghazi Khan District, Pun- jab (Pakistan)). 1951 Pinna shekhanensis Eames, ibid., p. 340, pi. 11, fig- 40 ( Kohat area ( Shekhan Nala section): Lower Shek- han Limestone ( Pakistan ) ) . Pinna muricata Linne, 1758 (Pis. 139, 141-145) Range — East Africa to eastern Polynesia. Remarks — The Prickly Pen Shell owes its com- mon name to the fairly fine, sharp spines which may be present on the radiating ribs on the exterior of the shell. These are a rather poor distinguishing character as they may be absent in this species, and other members of the family may exhibit them similarly. This species may be readily distinguished from other members of the genus in the Indo-Pa- cific by its exceedingly narrow longitudinal sulcus which brings the dorsal and ventral nacreous lobes into very close proximity. A specific character pos- sibly related to the last is the overlapping of a small portion of the posterior adductor scar onto the ventral lobe of the nacreous area. This condi- tion is unique in the genus Pinna, the posterior ad- ductor scar usually being limited entirely to the dorsal nacreous lobe. The movement of the adduc- tor muscle, and thus its scar, onto the ventral lobe appears to be an age-jrrogressive phenomenon. Young specimens of P. muricata commonly do not exhibit it, while older ones usually do, even though they may be stunted and small. The number of radiating ribs, usually 12 to 26, may also be a distinguishing character; P. bicolor usually has 8 to 17, except that occasionally the shells of injured specimens of bicolor which have regenerated may exhibit a higher number of ribs jmsteriorly; in incurva the ribs are obsolete. The characteristic of alternating smooth and spinose ribs mentioned by some authors in describing muri- cata and its synonyms is present in some specimens and absent in others (see also Remarks under P. bicolor). Bruce (in litt., 1960) reported finding specimens of the shrimp Anchistus custos ( Forskal, 1775) in the mantle cavity of P. muricata from Mazizini Bay, Zanzibar. [53 - 534] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 189 Plate 141. Pinna muricata Linne, Hilo, Hawaii. Fig. 1. Ex- terior of right valve. Fig. 2. Interior of left valve with poste- rior adductor muscle scar outlined in hlack. (reduced ¥2, Smithsonian Institution photo. ) Casts of fossil Pinna made available for my in- spection by Dr. H. S. Ladd, U. S. Geological Sur- vey, bear a striking resemblance to P. muricata Linne. They were found on Guam Island in the Mariana Limestone and are probably Pleistocene in age. Dr. Ladd (personal communication, March, 1960) suggested that the fossils are probably refer- rable to a present day species. These specimens are very probably P. muricata. Habitat — Pinna muricata is most commonly found with only the posterior margin of the shell protruding from sand or silty mud among rocks, in eel grass sand Hats, and in sandy patches on coral reefs, in shallow water from a few feet to two fathoms in depth. It has also been dredged alive from depths of 20 to 33 fathoms from sand and mud bottoms. Ecological data indicate that its tol- eration for fresh water at the mouths of rivers is fairly great. Description — Shell reaching 311 mm. (about 12K inches) in length; attenuately triangular in shape, the posterior margin usually sharply truncate; mod- erately inflated, a specimen 155 mm. in length is about 25 mm. in thickness at the widest point; with a moderately weak longitudinal keel on the ante- rior half of the shell. Valves rather thin, appearing fragile and sculptured with radiating ribs. Shells translucent and usually light horn color, but often having few to many narrow, irregular, blotchy, radiating tan to dark reddish brown bands on the posterior half. Surface generally smooth and shin- ing anteriorly where imbedded in substrate, but dull posteriorly where exposed, and sometimes coated with fine sandy mud. Radial sculpture con- sists of from about 12 to 26 radiating ribs limited to the posterior slope which often bear relatively few upright spines which are open posteriorly; ven- tral slope smooth or with obsolete ribs. There are often minor ribs occurring between the major spine-bearing ribs, together totaling the mentioned number. Concentric sculpture consists of quite fine [53 - 5351 190 Pinna J. Rosewater Pinnidae incremental lines which are transverse on the poste- rior slope, turning rapidly anteriorly on the ventral slope where they may form semilunar ridges of ir- regular occurrence. Posterior margin showing signs of repeated breakage and repair, usually truncate. Dorsal margin nearly straight to concave. Ventral margin broadly concave to convex and sometimes convex posteriorly and concave anteriorly. Interior of valves light horn color to faint yellowish green and sometimes with tinges of pinkish orange; pos- terior portion usually with brown to purplish black patches, smooth and glossy. Nacreous layer hardly iridescent, instead with a silvery or whitish veil, occupying the anterior one-half to three-quarters of the shell and divided along most of its length into two lobes by an especially narrow longitudinal sul- cus. Dorsal lobe of nacreous layer extending farther posteriorly than ventral lobe. Both lobes rather squarely truncate to bluntly rounded posteriorly. Anterior adductor muscle scar small, subapical, sit- uated a short distance anterior to the end of the longitudinal sulcus. Posterior adductor muscle sear subterminal on dorsal nacreous lobe, usually ex- tending partially onto the ventral lobe. Primary hinge ligament thin, black, extending from the an- terior end of the shell to the posterior border of the nacreous layer. Secondary ligament not colored, but evident in intact specimens whose dorsal mar- gins are fused. Embryonic valve about 0.5 mm. in length, rounded oval, not inflated; the umbos prominent, directed medially and slightly posteriorly ( plate 139, figs. 1, 2). Posterior mantle margin grayish black with white markings in preserved specimens; “eyes of Will darkly pigmented, but not conspicuous, widely spaced and located deep in the fold be- tween the inner and middle mantle lobes (see In- troduction: Remarks on Anatomy) . Measurements (mm.) — length width 311 149 (large; Mariveles Bay, Philippines' 245 98 (large; Oahu, Hawaii) 215 115 (large; Hilo, Hawaii) 150 92 (average; Cocos-Keeling Atoll) 82 37 (small; Cagayan, Philippines) Synonymy — 1758 Pinna muricata Linne, Systema Naturae, ed. 10, p. 707, (in M. Mediterraneo [Indo-Pacific] ) ; refers to Rumphius Mus., pi. 46, fig. M., and others; not P. muricata ‘Linne’ Holmes, 1860 [= Atrina serrata Sowerby, Western Atlantic]. 1786 Pinna nebulosa Solander, Catalogue of Portland Mu- seum, pp. 16, 71 (nomen nudum); = P. muricata Linne, fide Dillwyn (1817) Descriptive Catalogue of Recent Shells, vol. 1, p. 238. 1786 Pinna tenera Solander, ibid., p. 61 (nomen nudum); P . papyracea Gmelin [= P. muricata Linne] fide Dillwyn, loc. cit., p. 331. 1791 Pinna papyracea Gmelin, Systema Naturae, ed. 13, p. 3367 ( Oceano indico ) ; refers to Chemnitz, vol. 8, pi. 93, fig. 786. 1837 Pinna semi-costata Conrad, Journal Academy of Nat- ural Sciences, Philadelphia, vol. 7, p. 245, pi. 20, fig. 1 1 ( Sandwich Islands ) . 1843 Pinna virgata Menlce, Molluscorum Novae Hollandiae Specimen, p. 36, (ad litus occidentale [Western Australia] ) . 1858 (April) Pinna rumphii ‘Hanley’ Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 5, fig. 9 (Moluccas); Hanley [July] 1858, Proceedings Zoological Society of London, p. 136; Clessin, 1891, Conchylien Cabi- net, vol. 8, part 1, Malleacea, p. 98, pi. 18, fig. 3. 1858 Pinna philippinensis ‘Hanley’ Reeve, ibid., pi. 11, fig. 20 (Philippine Islands); [attributed to Hanley in error]. 1858 Pinna zebuensis Reeve, ibid., pi. 14, fig. 26 (Island of Zebu, Philippines). 1858 Pinna semicostata Reeve, ibid., pi. 16, fig. 30 (Philip- pine Islands). 1861 Pinna trigonalis Pease, Proceedings Zoological Society of London, p. 242 (Kingsmill [Gilbert] Islands); figured in von Martens and Langkavel (1871) Donum Bismarckianum, p. 64, pi. 4, fig. 7. 1866 Pinna philippensis ‘Reeve’ von Martens, Annals and Magazine of Natural History (3) vol. 17, p. 87 [error for philippinensis Reeve]. 1866 Pinna philippinarum ‘Reeve’ von Martens, ibid, [error for philippinensis Reeve], 1880 Pinna aequilatera von Martens, Beitrage zur Meeres- fauna der Insel Mauritius und der Seychellen: Mo1- lusken, p. 317, pi. 22, fig. 4 (Mauritius); not P. aequilatera Weinkauft, 1867, Die Conchylien des Mittlemeeres, vol. 1, Mollusca acephala, p. 236 [= P. nobilis Linne]. 1896 Pinna cebuensis ‘Reeve’ Elera, Catalogo Sistem. Fauna de Pilipinas, vol. 3, p. 800 (Cebu) [error for ze- buensis Reeve]. 1932 Pinna (Atrina) strangei ‘Reeve’ Prashad (part) The Lamellibranehia of the Siboga Expedition, System- atic Part II.: Pelecypoda, p. 136, pi. 4, figs. 11-13 (southern Philippines and East Indies) [Prashad’s figures are P. muricata Linne]; not P. strangei Reeve, 1858 [= Atrina pectinate Linne], 1938 Pinna hawaiensis Dali, Bartsch and Rehder, Bernice P. Bishop Mus. Bull. no. 153, p. 73, pi. 17, figs. 8-11 (dredged off Kaanapali, Maui, 4-8 fathoms [Ha- waii] ) . 1938 Pinna exquisite Dali, Bartsch and Rehder, ibid., p. 75, pi. 17, figs. 1, 2 (Albatross Sta.: 3965, near Laysan Island [Hawaii] in 147-116 fathoms on coral sand bottom ) . 1939 Quantidopinna delsa Iredale, Great Barrier Reef Ex- pedition Scientific Reports, vol. 5; Mollusca, part 1, p. 311, pi. 4, fig. 16 ( Low Isles, Queensland). 1939 Quantidopinna delsa howensis Iredale, ibid., p. 311 (Lord Howe Island [New South Wales]). Nomenclature — Linne (1758) listed as the type locality for Pinna muricata, “in M. Mediterraneo.” Turner and Rosewater (1958, pp. 302-303) consid- ered this problem and referred to pertinent discus- sions in the literature which show that this species is limited in its distribution to the Indo-Pacific. A possible explanation for Linne’s error in assigning muricata to the Mediterranean may be traced to the reference under the original description in the [53 - 536] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 191 Systema Naturae to Lister, pi. 370, fig. 215 (1685). On consulting Lister, no such combination of table and figures can be found. Figure 215 (pi. 374) is a representation of Pinna nobilis Linne, duly inscribed by Lister: “mar med.” Apparently, the figure to which Linne had intended to refer was 210 (pi. 370; the lower specimen, with the words, quoted by Linne, “Pinna tenuis striata muricata”) which shows a reasonable likeness of Pinna muricata from the Indo-Pacific. The locality, Jamaica, given by Lister, for the figured specimen is also in error. 3 his may account, also, for the several references to muricata in the West Indies which appear in the literature. bear transverse series of prickly, short, arched spines; with straight dorsal and ventral margins; the posterior margin being oblique and arcuately truncate. Although this description is not as clear as could be hoped and Menke’s types are not ex- tant (Keen, 1958, p. viii), this species is almost cer- tainly Pinna muricata Linne, whose range does not extend to South Australia, according to presently available records. The reddish lines, prickly valves and truncate posterior margin represent a combi- The holotype of Pinna exquisita Dali, Bartsch and Rehder is a unique specimen of this species described from a rather badly worn and partially broken shell dredged in Hawaiian waters. When considered apart from the range of morphological variation of the shell of P. muricata it may seem rather distinct. However, it is a young specimen which resembles certain other individuals of muri- cata of the same age. The unusually large number of ribs in this specimen are also present in other young muricata examined. The ribs may become indistinct as these individuals increase in size. With some doubt, Winckworth (1929) consid- ered Pinna virgata Menke to be a synonym of P. atropurpurea Sowerby (=P. bicolor G melin). Previ- ously Hedley (1924) had proposed that virgata was a valid species and that its range extended to South Australia. Later, Winckworth (1936) ap- peared to follow this suggestion. Critical reading of Menke’s original description reveals certain clues regarding the identity of virgata. The species was Plate 143. Pinna muricata Linne. Holotype of Quant uto- pinna delsa Iredale, Low Isles, Queensland. Fig. 1. Exterior of right valve. Fig. 2. Interior of left valve. (% nat. size, Australian Museum photo. ) [53 - 537] 192 Pinna J. Rosewater Pinnidae nation of characters not present in P. bicolor, but commonly found in specimens of muricata. Winckworth (1929, p. 284) included P. aeqiii- latera von Martens in the synonymy of P. atropur- purea (= P. bicolor). The figure (von Martens, 1880, pi. 22, fig. 4) clearly shows an average specimen of P. muricata, lacking spines but with characteristic coloration and a concave dorsal shell margin. Von Martens noted that P. muricata was similar to his species and gave the reference: Chem- nitz, vol. 6 [8], fig. 779, which is also muricata. Similar results were noted upon examination of photographs from the British Museum (N.H.) of the holotypes of Pinna philippinensis and zebuensis Reeve, i.e., formerly considered synonyms of atro- purpurea by Winckworth, upon critical study they were found to be synonyms of P. muricata. Pinna rumphii Reeve was considered by Winck- worth to be a synonym of the apparently rare P. incurva Gmelin. A photograph of the holotype in the British Museum (N.H.) shows rumphii to be a young specimen of P. muricata. T ypes — Since the whereabouts of the type speci- men of Pinna muricata is apparently unknown, Rumphius pi. 46, fig. M is here selected as the type figure. A discussion of the type locality of P. muri- cata is given above under Nomenclature and is also thoroughly discussed by Dodge (1952, pp. 227-228), Plate 144. Pinna muricata Linne, an example of the splen- did, large phenotype named philippinensis by Reeve; Mari- veles Ray, Bataan, Philippines. Fig. 1. Exterior of right valve. Fig. 2. Interior of left valve, the posterior adductor muscle scar outlined in black. (% nat. size, Smithsonian Institution photo. ) establishing the latter as an Indo-Pacific species. The type locality is here restricted to Amboina, Moluccas Islands, Indonesia, where the specimen figured by Rumphius was undoubtedly collected. The location of the type of P. papyracea Gmelin is not known. Chemnitz, vol. 8, pi. 93, fig. 786 is here selected as the type figure. The type of P. semi- costata Conrad may be in the Academy of Natural Sciences, Philadelphia, although it could not be found during a recent search there. Types of spe- cies described by Reeve are in the British Museum (N.H.). The type of Pinna virgata Menke is pre- sumed lost as is the balance of that author’s type material (Keen, 1958, p. viii). The type locality of P. virgata is here restricted to La Grange Bay, Western Australia, a locality from which specimens are known to have been collected. The type of P. aequilatera von Martens is probably in the Berlin Museum. The holotypes of Pinna hawaiensis and P. exquisita Dali, Bartsch and Rehder are in the United States National Museum. Types of Quantu- lopinna delsa and Q. delsa howensis Iredale are in the Australian Museum, Sydney. Lectotype Selection: Pinna trigonalis Pease was Plate 145. Pinna muricata. Lectotype of P. trigonalis Pease, Kingsmill Islands, Gilbert Islands, Micronesia. Fig. 1. Exte- rior of right valve. Fig. 2. Interior of left valve. ( reduced about Vz, Smithsonian Institution photo. ) [53-538] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 193 Plate 146. Pinna blanfordi Boettger. West Coast of Suma- tra, Indonesia. Lower Eocene. ( reduced Vi, from Boettger, 1880, pi. 3, figs. 4-6; Smithsonian Institution photo.) described from specimens collected in the Kings- mill [Gilbert] Islands, Micronesia. Although many of the Pease types are in the M.C.Z. the holotype of trigonalis is not. A syntype of trigonalis in the U.S.N.M. collection, received from Pease from the type locality, is here selected as lectotype; U.S.N.M. Catalogue number 41455; length 225 mm., width 82 mm. (plate 145, figs. 1, 2). Selected Records (see accompanying map, pi. 142). Solid dots; specimens examined; open circles: literature records — SOUTH AFRICA: Port Alfred (MCZ). MOZAMBIQUE: Inhaca Id., Delagoa Bay (J. K. Howard, MCZ; ANSP). TANGANYIKA: Dar-es-Salaam (MCZ). ZANZIBAR: Chumbe Id. ( NSF). RED SEA: Suez. GULF OF ADEN: Djibouti; Aden (all Lamv, 1928, p. 353). MADAGASCAR: Nossi Be (MCZ). SEYCHELLES: Astove Icl; West North and Menai Islands, Cosmoledo Atoll ( all A. J. Kohn, Pea- body Mus., Yale Univ. ); Mahe Id. (MCZ; A. J. Kohn and W. D. Hartman, Peabody Mus., Yale Univ.); Frigate Id. (Peabody Mus., Yale Univ.). MALDIVE ISLANDS: Doni- kolu Id., South Malosmadulu Atoll (A. J. Kohn, Peabody Mus., Yale Univ.) CEYLON: Hikkaduwa (G. F. Kline, ANSP; MCZ); Kacheri, Powder Bay, Trincomali (G. F. Kline, ANSP). JAPAN: Shikoku ( Habe, 1953a, p. 186). RYUKYU ISLANDS: Naga, Okinawa Id. (MCZ; USNM; ANSP). PHILIPPINES: many localities (see map; ANSP; USNM; MCZ). INDONESIA: Kilsium, off west coast Kei Ids. (MCZ, ex. Siboga Expedition); Koeta Bay, Bali (MCZ). COCOS-KEELING ISLANDS: Cocos-Keeling Atoll (USNM). AUSTRALIA: False Cape Bossuit, La Grange Bay, Western Australia (ANSP); Capricorn Group, Queens- land, Long Reef [north of Sydney], New South Wales (both J. Kerslake; USNM). NEW GUINEA: Mios Workbondi Ids., Schouten Ids. (NSF; MCZ). NEW CALEDONIA: Plage de Poe, Bourail (NSF). FIJI: Suva, Viti Levu (USNM). MARIANA ISLANDS: Saipan (USNM); Guam (NSF). PALAU IDS.: Babelthaup Id. (NSF). CARO- LINES: Ifaluk Atoll (F. M. Bayer, USNM); Kutu Id. Sata- wan Atoll (USNM). MARSHALLS: Bikini Atoll and others (see map) (USNM). GILBERTS: Apemama (USNM). HA- WAII: Kaneohe Bay, Oahu (C. M. Burgess, USNM). LINE ISLANDS: Palmyra Id. (MCZ; USNM). PHOENIX IDS.: Hull Id. (USNM). SAMOA: Tutuila Id. ( USNM L SOCI- ETY IDS.: west of Mount Tahara. District of Mahina, Ta- hiti (R. Robertson, ANSP). TUAMOTU IDS.: south end Oneroa Id., Raroia Atoll (J. P. E. Morrison, USNM); Hao and Vahitahi Atolls (both Dautzenberg and Bouge, 1933, p. 434). Fossil Records — Yokoyama (1923, Japanese Joum. Geol. and Geogr., vol. 2, no. 3, p. 57, pi. 6, fig. 15) reported a “fragment of a beak-portion with several distinct radiating riblets’’ which is probably referrable to P. muricata: Lower Pliocene, coast of Takinai, Kii, Japan. Fossils of this species have also been found on Guam Island in the Mariana Lime- stone and are of Pleistocene age (see Remarks) . Pinna blanfordi Boettger, 1880 (PI. 146) Range — Lower Eocene of Sumatra, Indonesia. Remarks — This species was considered to be dis- tinct from Atrina vexillum by Boettger, and it does seem to be a true Pinna. It is probably more closely related to P. bicolor than to either tnnricata or incurva. Synonymy — 1880 Pinna blanfordi, O. Boettger, In Verbeek, R. D. M., O. Boettger und von Fritsch, K., Palaeontographica, Supplement III., Lief. 8-9, Teil 1, p. 48, pi. 3, figs. 4-6 ( untereocanen Plattenkalken von Loerah Tarn- bang bei Boekiet Bessi [west coast, Sumatra]). Pinna bicolor Gmelin, 1791 (Pis. 147-1.53) Range — East Africa to Melanesia, including southern Japan, the Philippines and Australia; Hawaii. Remarks — It may be surprising to find the name P. atropurpurea included in the synonymy of P. bi- color. However, examination of as large a series of shells as possible from different localities through- out the range of these “species” has convinced me that they merge. The two forms are represented in plates 147 and 148. “Typical” bicolor seems to be more prevalent in the western Indian Ocean with the “atropurpurea” form becoming more common further east, although mixtures seem to occur. Ex- amination of large series from single localities (“ menkei ” from Australia; “atropurpurea" from the Philippines) shows a uniformity of characters in specimens from any one place. This may indicate that the species has a short larval life and oppor- tunities for exchange of genetic material between far flung localities is nearly prohibited. If this is so, it may serve as a partial explanation for the appar- ent considerable variation between different popu- lations of Pinna bicolor. Due to the effects of the physical environment which subject the shell to breakage, often followed by abnormal regrowth and repair, the substrate upon which the animal settles and the degree to which it is imbedded, a relatively broad or narrow [53 - 539] 194 Pinna J. Rosewater Pinnidae Plate 147. Pinna bicolor Gmelin, northeast end Maroepi Id., Ambai Group, northwest Dutch New Guinea. Specimen is 375 mm. (15 inches) in length. Fig. 1. Exterior of left valve. Fig. 2. Interior of right valve. (Smithsonian Institu- tion photo. ) shell with a variety of sculptural patterns may re- sult. The coloration of the shell is also highly vari- able sometimes being dark brownish purple but often light horn color or of alternating bands of both colors— possibly reflecting differences in avail- able food, chemical composition of the sea water or minor genetic variation. A degree of morphological stability is evident in Pinna bicolor amid the array of shell variation in this species. The animal within a shell looking much like “atropurpurea,” from New South Wales, is nearly identical to one from Hong Kong, the lat- ter having a shell more similar to the narrower. light horn form of bicolor. Furthermore, both speci- mens possess large “eyes of Will” protruding from their mantle edges. Pinna bicolor differs from muricata in having a rounded posterior margin, rather than a truncate one; in having the ventral nacreous area rounded, rather than somewhat truncate; in usually having 8 to 17 radiating ribs, rather than 12 to 26; and in usually being bluish black with radial rays, rather than being light horn to tan. The “eyes of Will” are conspicuous and closely spaced in the mantle of bicolor, while in muricata they are inconspicuous and widely spaced. Pinna bicolor may also be distinguished from P. incurva Gmelin on the basis of shell characters: shape, sculpture, color, transverse growth lines and the nacreous patches (see descriptions). Holthuis (1952) from quoted sources noted the [53 - 540] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 195 presence of the palaemonoid shrimps, Anchistus custos (Forskal) and Conchodytes biunguiculatus (Paulson) in the mantle cavity of P. bicolor from the Andaman Islands (see Kemp, 1922). Anchis- tus custos was also noted in P. mad id a Reeve [= P. bicolor] from Bowen, Queensland. Bruce (in litt., 1960) found specimens of the shrimp, Paranchistus ornatus Holthuis, in the mantle cavity of Pinna bi- color from Mazizini Bay, Zanzibar. For some reason, perhaps related to ecological tolerances, bicolor appears to be limited in its dis- tribution to the shores of larger land masses or to the islands in close proximity to continental areas. In this respect it differs from muricata whose dis- tribution includes oceanic islands. The sporadic ap- pearance of P. bicolor in Hawaii is probably the result of chance introduction. Habitat — Imbedded in muddy sand and reef Hats, in shallow water of 1 to 2 fathoms in depth. Description — Shell reaching 495 mm. (about 19 inches) in length; broadly to attenuately triangular in shape, the posterior margin varying from nearly truncate to more or less evenly convex-rounded and sometimes dorsally convex and sloping ventrally; moderately inflated, a specimen 255 mm. in length is about 35 mm. in thickness at its widest point; with a moderately strong longitudinal keel on the anterior half of shell. Valves varying from rather heavy and thick in large specimens to rather thin and fragile in smaller individuals, sculptured with radiating ribs. Shells translucent and varying from light horn color to dark purplish brown, often with radiating bands of alternating dark and light color (particularly conspicuous near the posterior border in older specimens) sometimes interrupted, giving the color pattern a concentric appearance. Surface of valves generally smooth and shining anteriorly where imbedded in substrate, but dull posteriorly and often encrusted with coralline algae, coral, other organisms and debris. Radial sculpture con- sists of from about 8 to 17 fairly well defined radi- ating ribs which may become nearly obsolete and which posteriorly may bear a few flat to tubular spines, in some specimens approaching a lamellose condition; spines usually nearly obsolete except near the posterior margin. Concentric sculpture consisting of fine growth lines which are bowed posteriorly and then slope anteriorly toward the ventral margin. Posterior margin often showing signs of repeated breakage followed by rough re- pair presenting a greatly thickened margin; al- though this may be entirely absent in specimens which apparently grow in sheltered areas; outline as stated above. Dorsal margin usually straight, but may be only slightly convex to moderately concave. Ventral margin often duplicating dorsal margin as the arms of an inverted isosceles triangle, but some- times convex posteriorly and concave anteriorly, almost forming an upright but flattened sigmoid curve. Interior of valves light smoky horn to dark purplish brown in color, often with the radiat- ing bands of alternating dark and light showing through; surface smooth and glossy. Nacreous area iridescent, roughly occupying the anterior half of the shell and is divided along most of its length by a longitudinal sulcus. Dorsal and ventral lobes of the nacreous area moderately well separated, the interlobe distance especially varying posteriorly. Dorsal lobe of nacreous area usually extends farther posteriorly than ventral lobe near the sulcus, its posterior margin truncate to slightly oblique. The ventral lobe may extend obliquely farther posteri- orly near the ventral margin or is sometimes ( aber- rantly ) unevenly truncate and shorter than the dor- sal lobe. Anterior adductor muscle scar small to Plate 148. Pinna bicolor Gmelin, Ambariotelo, between Nossi Be and Nossi Komba, Madagascar. Fig. 1. Exterior of left valve. Fig. 2. Interior of right valve; actual posterior border of ventral nacreous lobe outlined by solid black ir- regular line; broken line indicates probable normal appear- ance. ( % nat. size, Smithsonian Institution photo. ) [53-541] 196 Pinna J. Rosewater Pinnidae Plate 149. Geographical distributions of Pinna bicolor Gme- lin (solid dots and circles) and Pinna incurva Gmelin (tri- angles ) . moderate in size; subapical, located just anterior to end of longitudinal sulcus. Posterior adductor scar moderately large, subterminal, on ventral half of dorsal lobe; never extending onto ventral lobe. Pri- mary hinge ligament moderately thick, black; ex- tending from the anterior end of shell to near the posterior border of the nacreous layer. Secondary hinge ligament not colored, but evident in intact specimens whose dorsal margins are fused. Ovary of ripe female a vivid pinkish red; diges- tive gland green (Personal communication D. F. McMichael, Australian Museum); upon examina- tion of preserved animal collected from Sydney Harbor, N.S.W., in February, 1960, the ovary was found to be full of mature eggs. Posteriorly the mantle margin is grayish black with white vermicu- lations in preserved specimen; “eyes of Will” usu- ally large, conspicuous, darkly pigmented, closely spaced and sometimes protruding from between inner and middle mantle lobes (see Remarks on Anatomy). Measurements (mm.) — length width 495 180 ( large; Philippines ) 400 170 (large; Dutch New Guinea) 252 118 (average; Trincomali, Ceylon) 119 58 ( small; Tanganyika ) Synonymy — 1791 Pinna bicolor Gmelin, Systema Naturae, ed. 13, p. 3366 (in rnari rubro); refers to Chemnitz, vol. 8, pi. 90, fig. 780. 1819 Pimm dolabrata Lamarck, Animaux sans vertebres, vol. 6, p. 133 (les mers australes?) Gulf St. Vincent [South Australia] locality restricted by Cotton and Godfrey (1938, p. 84); refers to Chemnitz, vol. 8, pi. 90, fig. 780. 1825 Pinna atropurpurea Sowerby, A Catalogue of the shells of the Earl of Tankerville, Appendix, p. v ( no local- ity given); Wilkins, 1953, Proceedings Malacologi- cal Society of London, vol. 30, p. 24, pi. 6. 1843 Pinna deltodes Menke, Molluscorum Novae Hollandiae Specimen, p. 37 (prope Victoria river [Northern Territory, Australia] ) topotype sent by Menke to Reeve, figured in Conchologia Iconica, 1858, pi. 21, fig. 40, fide Winckworth ( 1929) and Iredale ( 1939). 1852 Pinna trigonium Dunker, Zeitschrift fur Malakozool- ogie, vol. 9, p. 60 ( Indiae orientales ) ; figured in Dunker (1858) Novitates Conchologicae Mollusca Marina, p. 27, pi. 8. 1858 Pinna electrina Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 14, fig. 25 (Moluccas). 1858 (May) Pinna fumata ‘Hanley’ Reeve, ibid., pi. 15, figs. 27, 28 (San Nicholas, Island of Zebu, Philippines); Hanley [Nov.] 1858, Proc. Zool. Soc. London, p. 227. 1858 Pinna madida Reeve, ibid., pi. 17, fig. 31 (Port Es- sington. New Holland ) . 1858 Pinna mutica Reeve, ibid., pi. 18, fig. 33 (Island of Negros, Philippines). 1858 (June) Pinna menkei ‘Hanley’ Reeve, ibid., pi. 18, fig. 34 (no locality given); Hanley [Nov.] 1858, Proc. Zool. Soc. London, p. 228 (Portus Jacksonicus [New South Wales] ). 1858 (June) Pinna euglypta ‘Hanley’ Reeve, ibid., pi. 20, figs. 37, 38 (Amboyna); Hanley [Nov.] 1858, Proc. Zool. Soc. London, p. 228. 1858 Pinna vespertina Reeve, ibid., pi. 23, fig. 44 (locality not given); not P. vespertina ‘Reeve’ Clessin, 1891, Conehylien Cabinet, vol. 8, part 1, Malleacea, p. 90, pi. 39, fig. 2 [= Atrina pectinata Linne; Clessin by error transposed Reeve’s figs. 43 and 44]. 1858 Pinna attenuata Reeve, ibid., pi. 24, fig. 46 ( Moluccas ). 1858 Pinna angustana ‘Lamarck’ Reeve, ibid., pi. 27, fig. 51 (Moluccas); not P. angustana Lamarck, 1819, from the Mediterranean [probably = P. nobilis Linne], 1858 (August) Pinna regia Hanley, Reeve, ibid., pi. 30, fig. 56 (Amboyna); Hanley [Nov.] 1858, Proc. Zool. Soc. London, p. 227. 1858 Pinna rostellum Hanley, Proceedings Zoological Soci- ety of London, p. 227 (Insulae Indicae). 1859 Pinna firnbriatula Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 33, fig. 63 (Japan). 1859 Pinna stutchburii Reeve, ibid., pi. 33, fig. 64 (Moreton Bay, Australia ) . 1887 Pinna inermis Tate, Transactions of the Royal Society of South Australia, vol. 9, p. 71, pi. 4, fig. 5 (from Eucla to the South-East [Western and South Aus- tralia] ) . [53 - 542] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 197 ?1891 Pinna cumingii Hanley’ Clessin, Conchylien Cabinet, vol. 8, part 1, Malleacea, p. 55, pi. 23, fig. 1 (Aus- tralien); not P. cumingii ‘Hanley’ Reeve, 1858 [= Atrina maura Sowerby, from Eastern Pacific], 1891 Pinna rollei Clessin, ibid., p. 77, pi. 30, fig. 1 (locality unknown ) . 1891 Pinna molluccensis Clessin, ibid., p. 82, pi. 33, fig. 1 (Moluccas) [new name for P. angustana Reeve, f858, not P. angustana Lamarck, 1819]. 1891 Pinna atrata Clessin, ibid., p. 83, pi. 32, fig. 2 (locality unknown). 1894 Pinna epica Jousseaume, Le Naturaliste, Revue Illus- tree Des Sciences Naturelles, Paris, 16th Annee (8th Annee de la 2nd serie) no. 182, p. 229 (Japon [Japan]); type figured by Habe, 1953a, Publ. Seto Mar. Biol. Lab., vol. 3, pi. 9, figs. 1, 2. 1901 Pinna cochlearis H. Fischer, Journal de Conchyliologie, vol. 49, p. 126, pi. 4, fig. 13 ( Djibouti [French Somaliland] ) . 1906 Pinna natalensis Smith, Annals Natal Government Mu- seum, vol. 1, p. 60, pi. 8, fig. 9 (Durban [South Africa] ). 1924 Pinna isosceles Hedley, Records of the Australian Mu- seum, vol. 14, p. 145, pi. 19, fig. 1 ( New South Wales: Port Jackson, North Harbour). 1924 Pinna menkei caviterga Hedley, ibid., p. 147, pi. 20, fig. 8 (Queensland: Fraser’s Island). 1924 Pinna scapula Hedley, ibid., p. 148, pi. 19, figs. 6, 7 (Northern Territory: Darwin). 1932 Pinna densecostata Turton, Marine Shells of Port Al- fred, p. 219, pi. 56, no. 1520 (Port Alfred [South Africa] ) . 1939 Exitopinna deltodes ultra Iredale, Great Barrier Reef Expedition Scientific Reports, vol. 5, Mollusca, part 1., p. 315 ( Batt Reef; Low Isles; Keppel Bay, Queensland ) . Nomenclature — It is no great wonder that gen- eral confusion exists concerning the species extant in the family Pinnidae. When one adds to their variability the various and sundry misconceptions which have accumulated over the years regarding Plate 150. Holotype of Exitopinna deltodes ultra Iredale [= a stunted and repeatedly broken specimen of P. bicolor Gmelin] Low Isles, Queensland. Fig. 1. Interior of left valve. Fig. 2. Exterior of left valve. ( about Vt nat. size, Australian Museum photo. ) the species the confusion becomes compounded. Both Winckworth and Iredale agreed that at least a topotype of the species, Pinna deltodes Menke, was available to Reeve (1858, pi. 21, species 40). Although easily mistaken for a specimen of Atrina vexillum, the figure, when carefully studied, reveals itself as representing an extremely badly broken and many times repaired specimen of Pinna bicolor. Hedley apparently misunderstood this species and placed it in the genus Atrina, Iredale ( 1939 ) con- sidered deltodes a valid species and even named a subspecies, ultra (pi. 150, figs. 1 and 2). In the same section he referred Pinna scapula Hedley ( 1924, pi. 19, figs. 4, 5 [6, 7] ) to this species. Iredale (pp. 315, 316) suggested, on the basis of the misshapen del- todes ultra, that vexillum Born is also a member of the “pinnoid” group and not an Atrina. Fortunately Born’s plate (1780, pi. 7, fig. 8) conclusively shows an Atrina with the postero-ventrally lobate shell which is characteristic of vexillum thus establishing the latter as a valid species belonging to the sub- genus Atrina s.s. The large number of synonyms included under Pinna bicolor testify to the variability of this spe- cies. The development of the narrow, elongate forms, such as attenuate Reeve, may be the result of the effect on the species when it is living near Plate 151. Pinna bicolor. Syntype of P. dolabrata Lamarck, South Australia. Fig. 1. Interior of left valve. Fig. 2. Exte- rior of left valve. Specimen is 215 mm. (8.6 inches) in length. (Photograph courtesy of E. Binder, Museum D’His- toire Naturelle, Geneva.) [53 - 543] 198 Pinna J. Rosewater Pinnidae the edge of its range (Japanese waters), although light-colored, narrow individuals resembling this phenotype occur with the more typical form in other parts of the range of the species ( stutchburii Reeve; angustana ‘Lamarck’ Reeve). When attempts are made to separate these forms it is found that intergrades exist. For this reason, at present, it seems best to include all of these apparent varia- tions under one name. Pinna fimbriatula Reeve was considered a dis- tinct species by Winckworth (1929) and by Habe ( 1953a. ) who placed it in the genus Atrina stat- ing: “This species has never been rediscovered . . . and . . . resembles a form of Atrina vexillum.” Photographs of the holotype of fimbriatula (B.M. 1952.8.29.22) including a view of the interior of a valve, show it to be a true Pinna and most cer- tainly a synonym of P. bicolor. Habe reprinted Reeve’s figure 63 of fimbriatula without securing a view of the interior. His reference to vexillum is thus understandable because there is a strong re- semblance, until the characteristic nacreous layer is examined. This short, wide form of bicolor was also described by Hedley as scapula, by Menke as clel- tocles, and Iredale later named a subspecies, del- todes ultra (pi. 150, figs. 1, 2). As mentioned by Hedley this broad shell form probably results from the animal being shallowly imbedded, perhaps in a little sand over rock, the resultant growth produc- ing a more broadly fan-shaped and less elongate shell. Correlated with the wider shell is the greater distance between the nacreous lobes. This was men- tioned as a characteristic of bicolor by Winckworth, and supposedly served to distinguish bicolor from atropurpurea. The unreliability of this character in this species is shown through examination of a photograph of the holotype of Pinna mutica Reeve (pi. 152, figs. 3, 4). Included in the synonymy of atropurpurea by Winckworth (1929, p. 283), the internal view shows the wide posterior separation of the nacreous lobes which that author attributed to bicolor. Habe (1953b, pi. 9, figs. 1, 2) figured the type of Pinna epica Jousseaume, including an internal view of the valve which reveals this species to be a true Pinna and a synonym of P. bicolor. Habe earlier ( 1953a, pi. 25, fig. 8 ) figured a specimen collected from Tosa Bay in 70 fathoms which he also called epica, placing it in the subgenus Servatrina. An in- ternal view of Habe’s epica was not given in the latter case. The generic misplacement is under- standable because the external sculpture of the Plate 152. Pinna bicolor Gmelin. Figs. 1-2. Lectotype of Pinna fumata Reeve (erroneously labeled “Holotype”) [= P. bicolor], San Nicolas, Island of Zebu, Philippines. (241 mm. or 9.6 inches in length ) . Figs. 3-4. Holotype of Pinna mu- tica Reeve [= P. bicolor], Island of Negros, Philippines. (215 mm. or 8.6 inches in length). Fig. 3. Interior of left valve showing wide posterior separation of nacreous lobes sup- posedly characteristic of bicolor in a form more typical of “atropurpurea” . (British Museum (N.H.) photos.) [53 - 544] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 199 shell is similar to that of an Atrina, the specimen being somewhat anomalous. Ti/pes — The location of the holotype of Pinna bicolor Gmelin from the Red Sea is unknown. The type figure here selected is Chemnitz, vol. 8, pi. 90, fig. 780. A syntype of Pinna dolabrata Lamarck is in the Museum D’Histoire Naturelle, Geneva, and was figured by Hedley (1924, pi. 21, figs. 14, 15). A neotype of P. atropurpurea Sowerby was selected and figured by Wilkins (1953, pi. 6) and is in the British Museum (N.H.). The neotype locality is Trincomali, Ceylon. The holotype of P. deltocles Menke is presumed lost (Keen, 1958, p. viii), how- ever, Reeve’s fig. 40 of the syntype sent to him by Menke is here selected as the type figure of this species (see Winckworth, 1929, p. 291; Iredale, 1939, p. 315). Types of species described by Reeve and Hanley are in the British Museum (N.H.) as is the type of P. natalensis Smith. A syntype of P. inermis Tate is in the South Australian Museum, Catalogue No. 14607, according to B. C. Cotton (in litt., 1960). A holotype was never designated by Tate. The location of the types of Clessin is un- known. Some of them were said to be in the Rolle collection which was undoubtedly dispersed. The types of P. conchlearis Fischer and P. epica Jous- seaume are in the Museum D’Histoire Naturelle, Paris. The types of P. isosceles, P. caviterga and P. scapula, all described by Hedley, and Exitopinna deltodes ultra Iredale are in the Australian Mu- seum, Sydney. The type of P. densecostata Turton Plate 153. Pinna bicolor Gmelin. Lectotype of Pinna eug- lypta Reeve (erroneously labeled “Holotype”), from Am- boina. Specimens 177 mm. (7 inches) in length. Fig. 1. In- terior of left valve. Fig. 2. Exterior of left valve. ( British Museum (N.H.) photo.) is in the Oxford University Museum. 1. Lectotype Selection: Reeve (1858, figs. 37, 38) figured two specimens of Pinna euglypta. Wilkins ( 1953, p. 27 ) listed these and another specimen as syntypes of this species, not selecting a lectotype because euglypta was considered a synonym of P. atropurpurea. Since atropurpurea is, in turn, here considered a synonym of bicolor it is probably best to fix the name euglypta upon a single specimen at this time. The specimen depicted in Reeve’s fig. 37 is here selected as lectotype of Pinna euglypta ‘Hanley’ Reeve: B.M. 1952.8.29.19; length 177 mm., width 90 mm.; type locality: Amboyna (plate 153, figs. 1, 2; photograph courtesy of British Museum (N.H.)). 2. Lectotype Selection: A similar case exists re- garding Pinna fumata ‘Hanley’ Reeve, 1858 (figs. 27, 28) which is also a synonym of P. bicolor. The specimen portrayed by Reeve (fig. 28) is a dark form; that in figure 27 displays the color pattern often seen in bicolor. The former specimen ( fig. 28, the more mature but dark form ) is here selected as lectotype of Pinna fumata ‘Hanley’ Reeve: B.M. 1952.8.29.23; length 241 mm., width 110 mm.; type locality: San Nicolas, Island of Zebu (Cebu), Phil- ippines (plate 152, figs. 1, 2; photograph courtesy of the British Museum (N.H.) ). Selected Records (see map. Pi. 149) — SOUTH AFRICA: Durban ( E. A. Smith, 1906). MOZAMBIQUE: Inhaca Id., Delagoa Bay (ANSP); Santa Carolina Id., Bazaruto Bay (MCZ). TANGANYIKA: Mboa Magi, 9 miles south of Dar- es-Salaam (USNM). ZANZIBAR: Mnazi Moja, Zanzibar City (USNM); 1 mile north of Chuckwani (NSF). RED SEA: near Hodeida, Yemen ( R. E. Kuntz, USNM); Suez (USNM). PERSIAN GULF: Ras Tanura, Saudi Arabia (ANSP; MCZ; USNM); Kuwait [Kuwait?] (USNM). MADAGASCAR: Ambariotelo, between Nossi Be and Nossi Komba (A. G. Humes, MCZ; USNM). MAURITIUS [?] (USNM). INDIA: Calcutta (USNM). CEYLON: Trinco- mali (MCZ; USNM). NICOBAR ISLANDS (ANSP). THAILAND: Sattahip (G. M. Moore, MCZ). MALAYA: Singapore (MCZ; USNM). CHINA: Chip-bee, Amoy (USNM); Port Shelter, Sharp Id. and Rocky Harbour, Tai She Wan, Hong Kong (both A. J. Staple, ANSP). JAPAN: Awaji (MCZ; USNM); Imaizumi, Kakoshima Bay (USNM). PHILIPPINES: many localities (see map; USNM; ANSP; MCZ). INDONESIA: Po Bui Icb, Sandakan, North Borneo (USNM); Amboina (USNM; MCZ); Kleine Kombius, Diokjkarta Bay, Java (MCZ). AUSTRALIA: Yallingup (USNM); mouth False Cape Creek, La Grange Bay (ANSP); Broome (MCZ; NSF; ANSP) all Western Aus- tralia; Arafura Sea, near Darwin, Northern Territory ( A. R. Cahn, ANSP); Hamilton Id., Cumberland Group ( J. K. Howard, MCZ); Hervey Bay ( J. Kerslake; U S N M ) both Queensland; Gunnamatta Bay, Port Hacking, south of Syd- ney, New South Wales (D. F. McMichael, Aust. Mus.; USNM); Outer Harbor. St. Vincent Gulf, South Australia (W. Old. USNM). NEW GUINEA: Moroepi Icb, Ambai Grou- iNSF; MCZ; USNM); Merauke (MCZ). NEW CALEDONIA (USNM); HAWAII: off Barbers Point, Oahu Id. ( C. M. Burgess, 1961, USNM). Fossil Records — Chapman (1920, Proc. Roy. Soc., Vic- toria, vol. 32, p. 229 ) recorded P. inermis Tate [= P. bicolor] from the older Pleistocene deposits of the Ooldea District, Victoria, Australia. [53 - 545] 200 Pinna J. Rosewater Pinnidae Pinna incurva Gmelin, 1791 (Pis. 149, 154, 155) Range — East coast of India, Burma and the East Indies ( Winckworth ) ; Nicobar Islands (Chem- nitz); the Philippines and northern Australia. Remarks — This species appears to be rather rare in collections. I have seen only three specimens: two from Australia, only one of which is in fair condition; a third from the Philippines is young (pi. 155, figs. 1, 2). Winckworth (1929) stated that P. incurva was “a form so distinct from others Plate 154. Pinna incurva Gmelin. Fig. 1. Type figure (from Chemnitz, 1785, vol. 8, pi. 90, fig. 778). Figs. 2-3. Specimen from Keppel Bay, Queensland. Specimen 290 mm. (11.4 inches) in length. Fig. 2. Exterior of right valve. Fig. 3. In- terior of left valve; borders of nacreous lobes and posterior adductor muscle scar outlined in black. ( Smithsonian Insti- tution photos. ) [53 - 546] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Pinna 201 of this group that Morch made a genus Cyrtopinna for it . It is easily separable from bicolor and muriccita because of its narrowly attenuate shape, apparently fairly uniformly light horn color, and the characteristic shapes of the posterior margins of the lobes of the internal nacreous layer which slope sharply obliquely toward dorsal and ventral shell margins. The name incurva applied by Gmelin to the species as here restricted probably makes reference to the posterior margin of the shell which slopes ( incurves ) gradually toward the dorsal mar- gin from a point far down the ventral margin. This character also is responsible for the generic name “Cyrtopinna” erected by Morch for this species (Cyrto: derived from the Greek, kyrtos, a combin- ing form meaning “anything curved”). Habitat — Probably lives imbedded deeply in sandy-muddy bottom, below the low tide mark; “dredged ... in 9-12 fathoms in Port Curtis” [east coast of Queensland, Australia] (Iredale, 1939). Description — Shell reaching 290 mm. (about 11 inches) in length, narrowly attenuate and wedge- shaped in outline; posterior margin extremely arcu- ate; subinflated; with a relatively weak longitudinal keel on the anterior half of the shell. Valves thin, fragile and sculptured with nearly obsolete radiat- ing ribs. Shell translucent to nearly transparent, light reddish brown to dilute yellowish horn color. Surface smooth and shining. Radial scnlpture hardly observable; consists of about 6 ribs limited to the posterior slope. Concentric sculpture of broadly arcuate growth lines arising on the dorsal margin, turning slightly posteriorly for a short dis- tance, then rapidly arching antero-ventrally to the ventral border; often forming distinct semi-lunar ridges on anterior part of ventral slope. Dorsal mar- gin straight; only occasionally subconvex to sub- concave. Ventral margin also straight. Interior of valves light horn to reddish brown in color, smooth and shining. Nacreous layer iridescent, occupying most of the anterior half of the shell and divided along most of its length into two lobes by a narrow, longitudinal sulcus. Both dorsal and ventral lobes with the posterior margins obliquely truncate; slop- ing from snleus toward dorsal and ventral shell margins; forming with this equal posterior exten- sion a wide and deep “V.” Posterior margins of both lobes may show slight embayments. Anterior adductor muscle scar moderately small, subapical, the sulcus nearly reaching it. Posterior adductor muscle scar medium sized, subterminal on ventral portion of dorsal nacerous lobe. Primary hinge liga- ment very thin, black, appearing not to extend to posterior tip of dorsal nacreous lobe. Embryonic valves and animal unknown. Measurements ( mm.) — length width 290 78 (average?; Keppel Bay, Queensland) 98 33 (small; Iloilo, Philippines) Synonymy — 1791 Pinna incurva Gmelin, Systema Naturae, ed. 13, p. 3366 ( in Oceano indico ) ; refers to Chemnitz, vol. 8, pi. 90, fig. 778; and others; 1818, Wood, Index Testaceologicus, London, p. 60 (Amboyna); 1825, Wood, ibid., pi. 13, fig. 15. 1825 Pinna incurvata Sowerby, A Catalogue of Shells of Earle of Tankerville, p. 23 (no locality given); 1858, Reeve, Conchologia Iconica, vol. il. Pinna, pi. 5, fig. 8 (Moluccas); both refer to Chemnitz, vol. 8, pi. 90, fig. 778; neither are Pinna incurvata Born, 1778 [= P. nobilis Linne]. Plate 155. Pinna incurva Gmelin. Young specimen from Iloilo, Panay, Philippines. Fig. 1. External view, right valve. Fig. 2. Internal view, left valve; the borders of nacreous lobes and posterior adductor muscle scar outlined in black, (slightly enlarged, Smithsonian Institution photo.) [53-547] 202 Pinna J. Rosewater Pinnidae Nomenclature — The several plate references cited by Gmelin for his species, incurva, present an unclear concept of this species. This is due to the fact that the Bonanni figures are of Pinna nobilis, and the Rumphius, Petiver and Klein figures are of misshapen specimens of P. muricata. Subsequent authors ( Sowerby and Reeve ) have limited their references to incurva of Chemnitz, pi. 90, fig. 778, which is done here also. This figure shows an at- tenuate Pinna with the arcuate margin of incurva and what appears to be sufficient of the growth line pattern to fix the identity (pi. 154). Although not an excellent likeness of the species, it will do. The name Pinna attenuate has apparently been applied to any elongate pinnoid shell. It is quite often used for the Japanese form which, however, has a heavier shell than incurva and is a narrow variety of P. bi- color. Types — The location of the holotype of Pinna incurva is unknown. Gmelin referred to several authors' figures (see above). Of these, Chemnitz, pi. 90, fig. 778 is here selected as the type figure (see pi. 154, fig. 1). The original type locality given by Gmelin: “in Oceano indico” (Nicobar Is- lands: Chemnitz) is here restricted to Keppel Bay, Queensland, Australia, a locality from which this species is known to have been collected and from which I have seen material. Pinna incurvata Sowerby, 1825, is apparently the earliest valid in- troduction of that name for the species incurva, that of Chemnitz being unavailable as it was intro- duced in a work not consistently binomial. Chem- nitz’ figure 778 must also be considered the type figure for this species. Born’s earlier introduction of the name P. incurvata appears to be synonymic with P. nobilis of the Eastern Atlantic; the figures to which he refers are the latter species. Reeve’s account of incurvata also refers to the Chemnitz figure, although apparently that author had a speci- men which he figured. Locality Records (A triangles, pi. 149) — INDIA: Ennur, near Madras (Winckworth, 1929). PHILIPPINES: Iloilo, Panay (USNM). INDONESIA: Amboina (Wood, 1818, p. 60). ALtSTRALIA: Keppel Bay ( H. Bernard, Australian Museum; USNM); Yepoon (J. Kerslake, Australian Mu- seum; LTSNM), both Queensland. [53 - 548] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 203 Genus Atrina Gray, 1842 Atrina is world-wide in distribution and reaches slightly more northerly and southerly latitudes than does Pinna. The reason for this may lie, in part, in its more generalized physiology and resultant wider ecological tolerances. Two subgenera are here recognized in Atrina. Atrina ( sensu stricto ) is world-wide in distribution. Servatrina is apparently absent from the Eastern Atlantic (see Table 1). Description — Shell reaching large size, up to 480 mm. (about 19 inches) in length, wedge, wing- shaped to subglobular in outline. Thin to rather heavy in structure and sculptured with spinose to imbricate, sometimes nearly obsolete, radiating ribs. Nacreous layer not divided by a longitudinal sul- cus, occupying entire surface of anterior two-thirds to three-fourths of the inner side of valves. The posterior adductor muscle scar located subcentrally. Subgenus Atrina Gray, 1842 Type: Atrina vexillum (Born, 1778) Externally, shells of the members of the sub- genus Atrina have the characteristics of the genus Atrina (sensu lato). The subgeneric difference is based on an internal character, the protrusion of the posterior adductor muscle scar beyond the pos- terior border of the nacreous layer. There are two Recent species in this subgenus in the Indo-Pacific and related areas, Atrina vexillum (Born) and A. squamifera (Sowerby). One fossil species from the Miocene of Ceylon is provisionally included here. Synonymy — 1840 Atrina Gray, Synopsis Contents of the British Museum, ed. 42, p. 151 [nomen nudum]; 1842, ibid., ed. 44, p. 83 [described but no species listed]. Type by subsequent selection ( Gray, 1847, Proc. Zool. Soc. London, p. 199 ) : Pinna nigra Dillwyn [= Atrina vexillum (Born, 1778)]. 1853 Pennaria Browne’ Morch, Catalogus Conchyliorum Comes de Yoldi, p. 51; based on Patrick Browne, Civil and Natural History of Jamaica, London, 1756, p. 412; also Browne, 1789 [fide notes in Division of Mollusks, U. S. National Museum: not binomial and the former is prelinnaean]. Type by subsequent se- lection (Turner and Rosewater, 1958, p. 310): Atrina rigida ( Solander, 1786); not Pennaria Oken, 1815, not de Blainville, 1818. Atrina (Atrina?) pachyostraca (Davies, 1923) Range — From the Miocene of Ceylon. Remarks — As indicated by Davies, pachyostraca may be similar to Atrina vexillum. However, this fossil species is based on fragments representing only portions of valves, and its relationship to other fossil or Recent Pinnidae is somewhat obscure. Synonymy — 1923 Pinna pachyostraca Davies, Quarterly Journal Geo- logical Society, London, vol. 79, p. 593, pi. 29, figs. 3-4 (Miocene ( Vindobonian) : Minihagalkanda, Southern Province ( Ceylon ) ) Plate 156. Atrina vexillum (Born). Figs. 1-2. Syntype of Indies] [= A. vexillum], (reduced Vi, photograph courtesy of Pinna nigrina Lam., “L’Ocean des grandes Indes” [East E. Binder, Museum D’Histoire Naturelle, Geneva.) [53 - 569] 204 Atrina J. Rosewater Pinnidae Atrina (Atrina) vexillum (Born, 1778) (Pis. 139, 156-158) Range — Central east Africa to eastern Polynesia. Remarks — The Flag Atrina received its name due to the fact that the specimen which Born de- scribed exhibited a condition often present in this species: an especially lobate ventral margin which somewhat presents the appearance of a flag blow- ing in the breeze. This condition is quite noticeable in medium sized and large specimens and occa- sionally develops in smaller individuals. The dark coloration of the interior and exterior of the valves is also present rather uniformly in mature speci- mens, but younger ones are often lighter and may approach the coloration of the Atrina squamifera or Servatrina pectinata groups. The posterior adduc- tor muscle scar bulges prominently beyond the pos- terior border of the nacreous area in mature Atrina vexillum. However, as discussed by Turner and Rosewater ( 1958, pp. 313-314 and pi. 159 ) this subgeneric character is actually progressively cor- related with the size and age of individual speci- mens: mature A. vexillum having the character well-developed, but immature forms often being quite difficult to distinguish from specimens of Servatrina pectinata. In A. vexillum, the posterior adductor muscle scar is usually contiguous with the posterior border of the nacreous layer in immature specimens before bulging beyond, while in S. pecti- nata it is always located well within the posterior nacreous border. Atrina (Atrina) vexillum is heavier, broader and more darkly colored than Atrina (Atrina) squa- mifera. A. vexillum is usually smooth, squamifera grossly spinose, although younger specimens of vexillum and the anterior portions of the shells of older ones commonly bear the characteristic pat- tern of low sculpture ( see Description ) . In south- east Africa vexillum is apparently replaced by squamifera. Atrina vexillum is probably one of the more eco- nomically important members of the family in the Indo-Paeific area. Because it attains a large size, the posterior adductor muscle can be used for food and is said to be quite delicious (personal communica- tion, H. A. Rehder, 1960; “gut zu essen,” von Mar- tens, 1880, p. 317). In Polynesia, at least, the black shell of this species is intricately carved by the natives and used to fashion lamps, other decorative articles and as plates for food (personal communi- cation, C. E. Cutress, 1960). Jameson (1902, p. 142) mentions finding pearls in Puma nigrina Lamarck ( = A. vexillum ) in New Guinea. Although black pearls produced by this animal are very beautiful they fracture easily and are soon destroyed because of their radial prismatic structure (see Haas, 1955; Turner and Rosewater, 1958). Holthuis (1952) noted, from quoted sources, the presence of the palaemonoid shrimp, Anchistus custos (Forskal) in the mantle cavity of Atrina vex- illum ( also reported in Pinna nigra ‘Chemnitz’ Dill- wyn ( = A. vexillum). Bruce (in litt., 1960) also re- ported finding A. custos with A. vexillum from Mazizini Bay, Zanzibar. This species does not seem to be at all common in Hawaii. The few records of its presence are based on dredged fragments and a single living in- dividual taken in 120 feet off Waikiki, Oahu. It may live there only in deeper water or perhaps only occasionally invades these comparatively northern islands. Habitat — This species is usually collected in sandy mud, or in sandy eel grass patches on reefs. It is usually in water from 1 to 2 fathoms in depth, although a specimen from the Philippines was dredged dead, but in good condition, from a depth of 230 fathoms (USNM: 248688). It was collected alive in 20 fathoms in Hawaii. Description — Shell reaching 480 mm. ( about 19 inches) in length, triangular to subglobular to hatchet- (flag-) shaped in outline; inflated; rather heavy to thick in structure and sculptured with Plate 157. Atrina vexillum (Born). Lectotype of Atrina gouldii banksiana Iredale [= immature A. vexillum]. Low Isles, Queensland. ( reduced about %, Australian Museum photograph. ) [53 - 570] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 205 occasionally spinose radiating ribs. Shell semitrans- lucent, ranging in color from dark, sometimes red- dish, brown to black, but when viewed with trans- mitted light, a rich reddish purple shines through. Surface of valves usually dull; faintly shining in young specimens. Radial sculpture consists of from about 10 to 17 major, but sometimes obsolete, radi- ating ribs which become crowded and too numer- ous to count on the ventral slope; ribs often imbri- cate anteriorly, and posteriorly occasionally bear large, upright, often twisted spines which are open posteriorly. There are often from 1 to 3 rows of minor ribs between major spine-bearing ribs total- ing from about 13 to 25 in addition to the men- tioned number. Concentric sculpture consists of fine growth lines which are convex posteriorly. Posterior margin truncate to very broadly oval in outline. Dorsal margin usually nearly straight to definitely convex in flag-shaped specimens. Ventral margin broadly convex posteriorly and concave near the umbo. Interior of valves dark brown to black, but a lighter greenish brown mottling is sometimes present. Occasionally younger specimens have alternate bands of reddish purple and brown, especially conspicuous near the posterior margin; smooth and shining. Nacreous layer iridescent, occupying the anterior half of the interior of the valve. Posterior adductor muscle scar large, sub- circular, protruding beyond the posterior border of the nacreous layer in mature specimens; not pro- truding in immature specimens. Extent of pro- trusion varies and increases with the age of the specimen. Anterior adductor muscle scar small to moderate in size, located at the anterior tip of the nacreous area. Posterior border of nacreous area with embayment which forms a broadly obtuse angle ventral to the posterior adductor muscle scar; the contour of the ventral border follows the ven- tral margin of the shell; dorsal border contiguous with dorsal shell margin. Primary hinge ligament thick, black, extending from umbo to the posterior border of the nacreous area; secondary hinge liga- ment not colored but evident in intact specimens whose dorsal margins are fused. Embryonic valve about 0.5 mm. in length, broadly triangular, subin- flated, umbos directed posteromedially; sculptured with closely spaced, concentric, incised lines of growth (pi. 139, figs. 3, 4). Measurements ( mm.) — length width 425 295 (large; Indo-Pacific, USNM ) 210 145 ( average; Sangr i Sanga Id., P. I. ) 158 105 ( immature; off Waikiki, Hawaii) 123 75 ( immature; Suv a, Fiji) Synonymy — 1767 Pinna rudis var. B Linne, Systema Naturae, ed. 12, p. 1159; refers to Rumphius, pi. 46, fig. L; Gmelin, 1791, Systema Naturae, ed. 13, p. 3363 (in indico et mari rubro). 1778 Pinna vexillum Born, Index Rerum Naturalium Mus. Caes. Vindob., p. 118; 1780, Born, Testacea Musei Caesarei Vindobonensis, p. 134, pi. 7, fig. 8 (no locality given); [1956, Rutsch, Nautilus, vol. 69, p. 78], 1786 Pinna nigricans Solander, Catalogue of the Portland Museum, p. 147, species 3242 (Otaheite) [nomen nudum]. 1791 Pinna exusta Gmelin, Systema Naturae, ed. 13, p. 3366 (in Oceano indico australi); refers to Seba, pi. 91, fig. 2; and Chemnitz, vol. 8, pi. 91, fig. 782 (a young specimen ) . 1798 Pinna gubernaculum Roding, Museum Boltenianum, p. 159 (no locality given); refers to Gmelin, 1791, “Pinna vexillum’’ [p. 3366] species 15; and Chem- nitz, vol. 8, pi. 91, fig. 783 (a young specimen). 1817 Pinna nigra Dillwyn, Descriptive Catalogue of Shells, vol. 1, p. 325 ( Inhabits the coasts of Amboyna ( Rumphius ) ; Red Sea ( Chemnitz ) ; Otaheite ( Hum- phry) ); refers to Chemnitz, vol. 8, pi. 88, fig. 774. 1817 Pinna adusta Dillwyn, ibid., p. 328 (Inhabits the coasts of Manilla (Chemnitz); New Zealand (Hum- phry)); refers to Chemnitz, vol. 8, pi. 91, fig. 782 ( a young specimen ) . 1819 Pinna nigrina Lamarck, Animaux sans vertebres, vol. 6, p. 135 ( l Ocean des grandes Indes); refers to Chemnitz, vol. 8, pi. 88, fig. 774; 1825, Sowerby, The Genera of Recent and Fossil Shells, part 26, pi. 103. 1858 (May) Pinna gouldii ‘Hanley’ Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 11, fig. 21 (locality not given); Hanley [Nov.] 1858, Proc. Zool. Soc. Lon- don, p. 255 ( “Amboyna” ) . 1858 Pinna hystrix Hanley, Proc. Zool. Soc. London, p. 226 (Amboyna); figured in Reeve, 1859, Conchologia Iconica, vol. 11, Pinna, pi. 32, figs. 60, 61. 1938 Pinna oahua Dali, Bartsch and Rehder, Bernice P. Bishop Museum Bull. no. 153, p. 75, pi. 18, figs. 5, 6 (Oahu, Hawaiian Ids., 33-50 fathoms). 1938 Atrina (Atrina) recta Dali, Bartsch and Rehder, ibid., p. 76, pi. 17, figs. 3, 4 (Albatross Station 3850, off south coast of Molokai [Hawaii] 33-50 fathoms). 1939 Atrina gouldii banksiana Iredale, Great Barrier Reef Expedition Scientific Reports, vol. 5, Mollusca, part 1, p. 316, pi. 4, fig. 18 (Low Isles [off Port Douglas] Queensland ) . 1953 Atrina (Servatrina) tenuis Habe, Illustrated Catalogue of Japanese Shells, vol. 1, p. 193, pi. 24, fig. 3 (Tosa Bay, Shikoku ) . 1953 Atrina (Servatrina) linguafelis Habe, ibid., p. 194, pi. 25, fig. 7 ( Tosa Bay, Shikoku, about 200 fathoms in depth ) . Nomenclature — As noted by R. F. Rutsch in 1956 (Nautilus, vol. 69, no. 3, p. 78) the earliest reference for Rorn’s Pinna vexillum is 1778, and not 1780. Atrina is a feminine noun (from Latin: ater, atra = black colored). It would be expected that the endings of specific adjectival names should agree in gender with this genus. However, the name vexil- lum ( Latin = flag) is a substantive noun, not an adjective, and must retain its neuter ending and cannot be spelled “vexilla." The figures referred to by Gmelin for Pinna ad- [53-571] 206 Atrina J. Rosewater Pinnidae Plate 158. Geographical distribution of Atrina vexillum ( Born ) . usta, by Reeve for gouldii, Roding for g ubernacu- lum and Dillwyn for P. adusta may be questioned as to their specific identity with Atrina vexillum. They all represent the shells of fairly young indi- viduals of this species which are admittedly diffi- cult to distinguish from Atrina pectinata. However, examination of a geographic series of vexillum in which are found representatives of both young and old specimens yields shells similar to those figured or referred to by the above authors and proves their identity. Pinna hystrix Hanley was included in the syn- onymy of P. strangei by Winckworth. As strangei is here considered a synonym of Atrina pectinata, it would seem simplest to treat hystrix similarly. However, the situation is more complex. Wilkins ( 1953, p. 27 ) found three “paratypes” of hystrix in the British Museum collection. The holotype in the Museum Hanley was not located and is presumed lost. Reeve ( 1859, figs. 60, 61 ) figured two of the paratypes and a photograph of the third (courtesy of the British Museum) is in my hands. The para- types figured by Reeve apparently represent a mid- dle-aged ( fig. 60 ) and a young ( fig. 61 ) specimen of Atrina (Atrina) vexillum. To judge from the growth form, these specimens probably grew rather rapidly under favorable conditions, their sculpture being well-developed. The secondary ribbing char- acteristic of vexillum may be noted especially in the smaller specimen. The third paratype of which photographs have been made is a specimen of Atrina pectinata. From all available information. Pinna hystrix is a composite species representing both pectinata and vexillum. This mixture may pos- sibly account for the East African ( Red Sea ) litera- ture records cited by various authors for hystrix and strangei (see Lynge, 1909; Lamy, 1928) which are probably traceable to variations of Atrina vexil- lum. As here understood, the range of Atrina pecti- nata does not extend farther west than the coast of India, although additional records may change this. Because the usual interpretation of P. hystrix is based on the figures of Reeve which are Atrina vex- illum it seems best to consider hystrix a synonym of vexillum (see Types). As noted by Dodge (1952, p. 225) and earlier by Hanley (1855, pp. 148-149) the variety B men- tioned by Linne under P. rudis in the 12th edition of the Systema Naturae, and by Gmelin in the 13th, is without doubt referrable to Atrina vexillum Born. Linne’s reference to Rumphius, pi. 46, fig. L is vex- illum, and Gmelin’s to Rumphius, pi. 46, fig. 11 [?L] ; Gualtieri, pi. 81, fig. A; and Chemnitz, pi. 88, fig. 774 are also this species. Roding (1798, p. 160) cited for his species P. ferruginea: Gmelin’s P. rudis var. B, but gave as the plate reference Chemnitz, pi. 88, fig. 773, which is the true Pinna rudis Linne. Although ferruginea has been considered a syno- nym of rudis by Winckworth (1929) and Turner and Rosewater (1958) it would be reasonable to suspect that Roding’s reference to Chemnitz’ fig. 773 was an error for fig. 774 which would then place the name in the synonymy of vexillum. On the preceding page (159) Roding cited P. rudis var. B, but gave Chemnitz pi. 88, fig. 774 as refer- ence (= A vexillum). Therefore, it would seem that Roding did not separate rudis and its var. B, that his reference to fig. 773 under ferruginea was pur- poseful, and that that name must stand as a syno- nym of P. rudis from the Eastern Atlantic and West Indies. Types — The holotype of Pinna vexillum Born is in the Austrian Museum of Natural History, Vienna. Types of species described by Hanley and Reeve are in the British Museum (N.H.), London, ac- cording to Wilkins ( 1953). In view of the confusion [53 - 572] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 207 concerning the true identity of Pinna hystrix, it may be valuable to restrict the interpretation of this spe- cies to Atrina vexillum Born; the type figure here selected: Reeve, vol. 11, Pinna, fig. 60, thus remain- ing consistent with previous usage. As will be noted, the name P. hystrix (part) is also cited in the syn- onymy of Atrina pectinata. A syntype of Pinna nigrina Lamarck is in the Museum of Natural His- tory, Geneva, Switzerland (pi. 156, figs. 1, 2). Lectotype Selection: Atrina gouldii banksiana Iredale, 1939, was described from several Great Barrier Reef specimens. A sketch was given of one of these from Low Isles [off Port Douglas, Queens- land] and it was more fully described than the others, although never specifically designated as holotype. This specimen is here selected as lecto- type of Atrina banksiana Iredale: Australian Mu- seum Catalogue number C. 62033, length 110 mm., width 60 mm. (pi. 157, figs. 1, 2). Types of Atrina recta and Pinna oahua Dali, Bartseh and Rehder are in the United States Na- tional Museum. The locations of the types of the following are unknown to me: Pinna nigricans Solander, Pinna exusta, Gmelin, Pinna gubernacu- lum Roding, and Pinna nigra and adusta Dillwyn. The types of Servatrina tenuis and linguafelis Habe are probably at the Zoological Institute, Kyoto Uni- versity, Kyoto, Japan. Selected Records (see map, pi. 158); solid dots: speci- mens examined; open circles: from the literature — TAN- GANYIKA: Mboa Magi, 9 miles south of Dar-es-Salaam (USNM); Dar-es-Salaam (MCZ). ZANZIBAR: Pange Id.; Kiwengwa (both NSF); Mnazi Moja, Zanzibar City (USNM). KENYA: Diani Beach, Mombasa (MCZ). RED SEA: Massaua, Eritrea; Djibouti, French Somaliland. GULF OF ADEN: Aden (all Lamy, 1928). MADAGASCAR: Nossi Be (MCZ). MAURITIUS (ANSP). CEYLON: Hikkaduwa; south shore Ft. Frederick, Trincomali (both G. F. Kline, ANSP). ANDAMAN ISLANDS: Port Blair (MCZ). THAI- LAND: Prochuap; Mutapone Id., Chumphon (both G. M. Moore, MCZ); Koh Tao; Koh Phangan (both USNM). MALAYA: Singapore ( R. D. Purchon, USNM; MCZ; ANSP). CHINA: Rocky Harbour, Datum Point, Hong Kong (A. L Staple, ANSP); Hong Kong (USNM). JAPAN: Shikoku (Habe, 1953a, p. 190). RYUKYU ISLANDS: (MCZ; ANSP; LISNM), Yaeyama [Retto] (Mus. Zool. U. Mich.). PHILIPPINES: Many localities (see map). INDO- NESIA: Salajar Id., off southern tip of Celebes (ex Siboga Expedition, MCZ); Amboina, Moluccas (MCZ; USNM); Bantam, Java (USNM). COCOS-KEELING ISLANDS (USNM). AUSTRALIA: Hayman Id., northwest of Hook Id., Whitsunday Passage, Queensland (MCZ). NEW GUINEA: Rouw Id., Aoeri Ids., Geelvink Bay (MCZ; NSF); China Strait, Papua (USNM). SOLOMON IS- LANDS: Kieta, Bougainville Id. (ANSP). NEW CALE- DONIA: Bourail (NSF). FIJI ISLANDS: Yasawa Ids. (USNM). Suva, Viti Levu (USNM). PALAU ISLANDS: 1 mile south of West Passage, Babelthaup Id. (NSF); Malakal Harbor, Koror Id. (MCZ; NSF; USNM). CAROLINE IS- LANDS: Ponape (MCZ; USNM). MARSHALL ISLANDS: Arno Atoll ( R. W. Hiatt. USNM). GILBERT ISLANDS: Kingsmill Group (ex A. Garrett, MCZ). HAWAII: Midway Id. (MCZ); off Waikiki, Oahu (Cliff Weaver! ex C. M. Bur- gess); south coast Molokai Id., 43-66 fathoms (USNM); Auau Channel between Maui and Lanai Ids. (USNM). LINE ISLANDS: Palmyra Id. (ANSP; USNM). SOCIETY ISLANDS: Raititi Point, Bora Bora (R. Robertson); Raiatea; Baie de Maroe, Huahine (both USNM); Tahiti (H. A. Pils- bry! ANSP; MCZ). Fossil Records — Martin ( 1910, p. 357) recorded A. vexil- lum as a fossil from Tegal, Java, Indonesia (Upper Miocene, Tjilanang beds and Pliocene) fide Van der Vlerk (1931, p. 269); also see Tesch (1920, p. 91). Martin (1879, Die Ter- tiarschichten auf Java, p. 120, pi. 19, figs. 5-6) figured a fragment of the fossil he called vexillum. Beets ( 1950, p. 299) recorded fossil vexillum from the Tertiary or Quater- nary of the island of Mandul, East Borneo. Atrina (Atrina) squamifera (Sowerby, 1835) (Pis. 159, 160) Range — Known only from the east coast of the Union of South Africa. Remarks — Atrina squamifera is apparently lim- ited in its distribution to the southeastern portion of the Union of South Africa. It cannot easily be confused with any other species since there is nothing like it within its range. The other member of the subgenus Atrina in East Africa, A. vexillum Born, seems not to occur so far south. A. squa- mifera has a much more narrow and less thick shell than vexillum; the number of radiating ribs in squamifera is about 12 or less, although 19 to 20 may be counted on the specimen figured by Krauss (1848, pi. 2, fig. 8). In vexillum there may be from 10 to 30 ribs, but they are usually smooth, although occasional spinose individuals occur. Shell colora- tion in squamifera is light tannish brown, while in vexillum it ranges from reddish brown to black. There is usually an embayment in the posterior border of the nacreous area which forms an acute or nearly 90° angle ventral to the posterior adduc- tor muscle scar in squamifera; the same structure in vexillum is usually broadly obtuse. There are superficial similarities between Atrina (Atrina) squamifera Sowerby and Atrina (Serva- trina) tasmanica Tennison-Woods. However, the two species belong to different subgenera and their ranges are separate ( see Remarks under tasmanica) . Nickles ( 1953, Bull. Institut roy. Sci. nat. Bel- gique, vol. 29, no. 13, p. 1, pi. 1, figs. 1, 2) described Pinna [Atrina s.s.?] chautardi from West Africa, assigning a range from Mauritania to Angola. The figures show a specimen which appears very simi- lar to A. squamifera. It is possible that the range of squamifera extends from West Africa to southeast Africa. However, until specimens are available for comparison, it seems best to consider them distinct species. There is, as yet, little basis for uniting West and South African faunas (see Stephensen, 1947, Ann. Natal Mus., vol. 11, part 2). 153 - 573] 208 Atrina J. Rosewater Pinnidae Obvious similarities also exist between squamif- era and chautardi, and fragdis of European waters. As pointed out by Tomlin (1922, Jour. Conch., vol. 16, pp. 255-262) present oceanic currents hinder dispersal of northern East Atlantic species toward tropical West African regions. However, the pres- ence of counter-currents and earlier geological dis- tribution patterns must be taken into consideration. These three forms may be the remnants of a former continually distributed species now broken-up into isolated populations which merit specific rank (see Kohn, 1956, p. 572). Habitat — The development of the large, nearly tubular spines suggests that this species lives in quiet water, probably in bays and other sheltered places. It has also been collected in the lower por- tions of rivers where the water has a tendency to be brackish. Description — Shell reaching about 368 mm. ( 14/2 inches, Sowerby, 1904, Marine Investig. South Af- rica, vol. 4, p. 3) in length, triangularly wing- shaped in outline; moderately inflated; rather thin and fragile in structure and sculptured with strongly spinose radiating ribs. Shell nearly transparent, light tannish brown in color, sometimes with a few re- peated transverse bars of violet evident between the ribs. Surface of valves dully shining. Radial sculpture consists of from about 10 to 12 fairly prominent radiating ribs becoming extremely crowded on the ventral slope giving the area a highly rugose appearance; ribs usually have small spines anteriorly and large, semitubular, and often Plate 159. Atrina squamifera (Sowerby). Figs. 1-2. Holo- Museum (N.H.) photo.) Figs. 3-4. Atrina squamifera. Port type of Pinna squamifera Sby., Cape of Good Hope, South Alfred, South Africa. ( Museum Comparative Zoology photo. ) Africa. Fig. I . Interior of left valve; border of nacreous layer All reduced about Vz nat. size, outlined with black. Fig. 2. Exterior of left valve. (British [53 - 574] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 209 twisted spines posteriorly. Concentric sculpture of fine, often irregular growth lines showing signs of past injury. Posterior margin usually truncate; occa- sionally rounded in stunted specimens (as in A. alfredensis Bartsch). Dorsal margin usually straight to slightly convex. Ventral margin slightly convex posteriorly, curving inward anteriorly and becom- ing somewhat concave. Interior of valves light tan- nish brown, the violet bars showing through; smooth and highly glossy in some specimens. The rather thin nacreous layer iridescent, occupying about an- terior one half to two thirds of interior of valve. Posterior adductor muscle scar medium sized, rounded-oval, protruding beyond posterior border of nacreous area. Posterior border of nacreous area with small embayment usually forming an acute angle ventral to posterior adductor muscle scar; ventral border following ventral margin of shell in contour; dorsal border contiguous with dorsal shell margin. Primary hinge ligament moderately thin, black, extending from umbo to posterior border of nacreous layer; secondary hinge ligament not col- ored but evident in intact specimens whose dorsal margins are fused. Measurements (mm.) — length width 290 102 (large; Port Alfred, Cape of Good Hope) 245 96 (large; Port Alfred, Cape of Good Hope) 75 40 (small; Port Alfred, Cape of Good Hope) 67 48 ( stunted; Port Alfred, Cape of Good Hope) Synonymy — 1835 Pinna squamifera Sowerby, Proceedings Zoological So- ciety of London, p. 85 ( ad Caput Bonae Spei [Cape of Good Hope, Union of South Africa] ). 1835 ?Pinna afra Sowerby, ibid., p. 85 (ad Caput Bonae Spei [Cape of Good Hope, Union of South Africa] ) . 1848 Pinna capensis Krauss, Die Siiafrikanischen Mollusken, p. 27, [line 33] ( In limo arenoso ad otium Hum, Knysna [Cape of Good Hope, Union of South Africa]; capensis not figured; squamifera figured, pi. 2, fig. 8. 1858 Pinna kraussii Hanley, Proceedings Zoological Society of London, p. 226 (Natal [Union of South Africa]). 1915 Atrina alfredensis Bartsch, Bull. United States National Museum, No. 91, p. 183, pi. 40, fig. 3 (Port Alfred [Union of South Africa]). 1932 Pinna segmenta Turton, Marine Shells of Port Alfred, p. 217, pi. 56, no. 1510 (Port Alfred). 1932 Pinna whitechurchi Turton, ibid., p. 218, pi. 56, no. 1512 (Port Alfred). 1932 Pinna rufanensis Turton, ibid., p. 218, pi. 56, no. 1513 (Port Alfred). Nomenclature — With the exception of Pinna densecostata ( =P . bicolor), and P. anomioides and similis (both = S. saccata Linne), all of the new names of Pinnidae introduced by Turton (1932) are synonyms of Atrina squamifera. His figures, al- though mostly of immature shells, are recognizable. The specimens called Pinna afra Sowerby and men- tioned by Turton as having been identified by Dr. Paul Bartsch are indeed fragments and are repre- sented in the United States National Museum col- lection (catalogue nos. 187160, 250991) from the Port Alfred area. It is difficult to assign these frag- ments generically and less specifically. It is proba- Plate 160. Geographical distribution of Atrina squamifera (Gray) (New Zealand); and A. tasmanica ( Tenison- Woods) (Sby. ) (South Africa); A. pectinata pectinata (Linne) ( Tasmania and Southeastern Australia) . (South Asia and Western Pacific); A. pectinata zelandica [53 - 575] 210 Atrina J. Rosewater Pinnidae ble that they are fragments of squamifera, badly beachworn. The true identity of P. afra is still un- clear and the types apparently lost. Considering the type locality and general description of the species, I must agree with Winckworth ( 1929, p. 290 ) that afra is probably a synonym of squamifera. Types — The holotype of Pinna squamifera Sowerby is in the British Museum (N.H.). It is here figured, pi. 159. Locations of the holotypes of P. afra Sowerby, P. capensis Krauss and P. kraussii Hanley are unknown. The Turton types are in the Oxford University Museum. The holotype of Atrina alfredensis Bartsch is in USNM, no. 227815. Records (see map, plate 160) — SOUTH AFRICA: Cape of Good Hope (ANSP); Jeffreys Bay ( MCZ; ANSP; Mus. Zool. U. Mich.); Port Elizabeth (MCZ); Kowie River, Port Alfred (MCZ); Port Alfred (MCZ; USNM); Natal (Han- ley, 1858). [53 - 576] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 211 Subgenus Servatrina Iredale, 1939 Type: Atrina pectinata (Linne, 1767) Externally the shells of the members of the sub- genus Servatrina have the characteristics of the genus Atrina (s.l. ). The subgeneric difference is based on an internal character: the posterior ad- ductor muscle scar is enclosed completely within the nacreous area rather than protruding beyond its posterior border as in Atrina (s.s. ). This subgenus is world-wide in distribution in tropical and warmer temperate seas; however, it appears to be absent from the Eastern Atlantic. There are 3 Recent species or subspecies in this subgenus in the Indo-Pacific area: S. pectinata pec- tinata Linne, pectinata zelandica Gray and S. tas- manica Tenison-Woods. In addition, 3 fossil species from the mid-Tertiary of southern Australia are in- cluded here. Synonymy — 1939 Servatrina Iredale, Great Barrier Reef Expedition Scientific Reports, vol. 5, Mollusca, part 1, p. 317. Type by original designation: Pinna assimilis Reeve [= Atrina pectinata Linne, 1767]. Atrina (Servatrina) pectinata (Linne, 1767) Within its relatively narrow range in the Indo- Pacific, Atrina (Servatrina) pectinata fulfills the pre- requisite of a species of the Pinnidae by exhibiting maximum variation. The species ranges from the pale, translucent, subinflated olivaceous form of south India to the large, tumid, dark “ japonica ” of southern Japan. Throughout this range, however, distribution is probably continuous, thus affording an opportunity for the exchange of genetic material among the several forms. It is doubtful that free genetic exchange occurs between the East Indian and Australian populations of pectinata and those of New Zealand. If such does rarely occur, it would be in the southern direction. For this reason it is convenient to consider the New Zealand form of pectinata as a subspecies. Servatrina pectinata pectinata is limited in its dis- tribution to southern and southeast Asia, the East Indies, northern Australia and Melanesia. The sub- species zelandica inhabits the islands of New Zea- land. With the exception of these two forms the only other representative of the subgenus Serva- trina present in the Indo-Pacific is S. tasmanica Tenison-Woods from northern Tasmania and south- eastern Australia. Atrina pectinata subspecies pectinata (Linne, 1767) (Pis. 139, 160-163) Range — Southeast India and Ceylon to western Melanesia. Remarks — The Comblike Atrina probably de- rived its name from the spines which commonly Plate 161. Atrina pectinata pectinata (Linne). Fig. 1. Ex- 3-4. Internal and External views of Holotype of Pinna serra ternal view of specimen from Fukura, Awaji, Japan, of large Reeve [= A. pectinata ], Moreton Bay, Queensland, (about size, probably cultivated for food. Fig. 2. Interior of same Vz nat. size, British Museum (N.H.) photos.) specimen (% nat. size, Smithsonian Institution photos.) Figs. [53 - 597] 212 Servatrina J. Rosewater Pinnidae Plate 162. Atrina pectinata pectinata (Linne). Holotype of Pinna lurida Reeve. Philippine Islands. Fig. 1. Exterior of right valve. Fig. 2. Interior of right valve. (V2 nat. size, Brit- ish Museum ( N.H. ) photos. ) protrude from the dorsal-most rib of the shell and resemble the teeth of a comb. These spines may be present even when the remainder of the valve is quite smooth. They are not evident in Linne’s ref- erence, Gualtieri, plate 79, fig. A., but are present in similar specimens. Atrina pectinata cannot be easily confused with any other Indo-Pacific species when the range of its variation is understood. It varies from nearly smooth to finely imbricate in sculpture, and, in the northern part of its distribution, attains a fairly large size. The subgeneric characteristic of an in- ternal posterior adductor muscle scar is constant and is shared only with Servatrina pectinata ze- landica and S. tasmanica. This species has quite often been dredged in rela- tively deep water. In fact, dredgings by the United States Bureau of Fisheries Steamer Albatross in the Philippines in 1908 yielded large numbers of very thin, broken valves of pectinata from depths as great as 300 fathoms ( Albatross Station 5189). A small living specimen was collected from 170 fath- oms at station 5267 off Matocot Point, west Luzon. It has generally been believed that the Pinnidae are a shallow-water group. In the case of A. pecti- nata, however, this appears to have been a miscon- ception, at least partially. It is probable that the usual habitat of pinnas is in shallower water than [53 - 598] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 213 that just mentioned. The young living specimen col- lected in nearly 200 fathoms may have settled there and woidd not have survived much longer. The quantities of valves collected at 300 fathoms may have been washed to this depth by ocean currents. It seems apparent, however, that pectinata does often occur in other than shallow water in certain parts of its range. The species lives in fairly shallow water in the northern portion of its range on the coast of Japan but its vertical distribution ranges to deeper waters here also. Cahn ( 1951 ) reports that Atrina japonica Reeve (= pectinata Linne) is found in depths of from 8 to 60 meters ( about 4.4 to 33 fathoms). It is reported from Tosa, Japan, from 100 fathoms. Some of the largest specimens seen are from Japanese waters, reaching nearly 37 centi- meters (about 14M inches) in length. The species is used commercially there for food and fertilizer and is planted intertidally and actively cultivated for subsequent harvest. A shrimp, Conchodytes nipponensis (DeHaan) (identified by Dr. L. B. Holthuis of Leiden, Hol- land), was found living in the mantle cavity of a specimen of pectinata from Japen Island, Dutch New Guinea. This species of shrimp has been re- ported in pectinata previously by Kubo ( 1940 ) . Cahn ( 1951 ) reported that pectinata is dioecious; spawning occurs from June to September in Ariake, Japan; the optimum critical temperature for spawn- ing is between 24 and 27° C.; sexual maturity is reached at about one year. Yoshida (1956) reported on a study of the early life history of pectinata in Japan and figured stages in the development of the veliger. Veligers of pecti- nata were found to be of larger size than any other species of Pinnidae studied. In the present study, the embryonic valves of this species were found to be about twice the size of other species available for study (see pi. 139, figs. 1-6, and the descriptions of pectinata, vexillum and muricata ). Yoshida found young (post veligers) of pectinata buried in bot- tom mud and attached to apparatus used in collect- ing the spat of the ark clam, Anadara subcrenata. Comparisons made between young of pectinata and “ Pinna (Atrina) japonica” (of Yoshida) [= Pinna bicolor Gmelin?] showed the former to differ from the latter in the shape of the posterior margin and in the lack in pectinata of the pinnoid sulcus and dark coloration in the valves. Habitat — Cahn (1951) has reviewed the Japa- nese literature concerning the ecology of Atrina (Servatrina) japonica Reeve (= pectinata Linne). The species lives in sand or sandy mud and is buried almost completely during the summer, but works up so that only the anterior tip is buried in winter. Orientation is with the ventral ( open ) por- tion of the shell facing toward the current. The species survives a range of temperatures from 39° C. to 1° C., and is resistant to sea water with low- ered salinity. The major foods consumed by pecti- nata in Japan were found to be diatoms, copepods, and protozoans. Description — Shell reaching 370 mm. (about 14/2 inches) in length; triangular-wedge-shaped in out- line and moderately to strongly inflated; rather thin and fragile to only moderately heavy in structure; sculptured with radiating ribs which may be nearly spineless, or bear minute imbrications ranging to a few short, upright spines which are open posteri- orly. Shell translucent, usually olivaceous tan in color, approaching dark brown in some specimens. Surface of valves faintly shining. Radial sculpture consists of from 15 to 30 radiating ribs on the pos- terior slope; in young specimens ribs may all be fine and closely spaced; sculpture on the ventral slope becomes crowded and impossible to count. Ribs often smooth but usually ranging from finely imbricate to distinctly spinose. Concentric sculp- ture of fine lines of growth, convex posteriorly and sometimes wavy. Posterior margin usually truncate; the junction of posterior and ventral slopes pro- jecting farthest posteriorly. Dorsal margin usually nearly straight, often with a series of short spines protruding from the most dorsal rib (from which the name “ pectinata ” is derived). Ventral margin forms a gentle sigmoid curve: convex posteriorly, concave anteriorly. Interior of valves the same oli- vaceous tan, ranging to dark brown or nearly black in Japanese specimens; smooth and shining. Nacre- ous layer iridescent, occupying the anterior two- thirds to three-quarters of the valve. Posterior adductor muscle scar medium to large in size, sub- circnlar, located well within the posterior border of the nacreous area, but never protruding beyond. Anterior adductor muscle scar small, located at tip of nacreous area. Posterior border of nacreous layer broadly rounded with no distinct embayment. Pri- mary hinge ligament fairly thin, black and extend- ing from the anterior end of the shell to the pos- terior border of the nacreous layer; secondary hinge ligament not colored but evident in intact speci- mens whose dorsal margins are fused. Embryonic valves about 1 mm. in length, broadly triangular in outline, inflated; the umbos directed postero-medi- ally (pi. 139, figs. 5, 6). [53 - 599] 214 S ervatrina J. Rosewater Pinnidae Measurements (mm.) — length width 370 185 (large; Japan [Calm, 1951]) 350 220 (large; Fukura, Awaji, Japan) 185 107 (average; Lem Sing, Thailand) 90 40 (small; Biliran Id., Philippines) 48 19 (small; Biliran Id., Philippines) Synonymy — 1767 Pinna pectinata Linne, Systema Naturae, ed. 12, p. 1160 (in India); refers to Gualtieri, pi. 79, fig. A. ?1798 Pinna inflata Roding, Museum Boltenianum, Ham- burg, pt. 2, p. 159 (nomen nudum, see Winck- wortli, 1929). 1798 Pinna vitrea Roding, ibid., p. 159; refers to Gmelin s Pinna pectinata var. B, p. 3364, and Chemnitz, pi. 87, fig. 771 (not P. vitrea Gmelin, 1791 = Strepto- pinna saccata L. ) 1817 Pinna inflata Dillwyn, Descriptive Catalogue of Recent Shells, London, p. 326 ( Inhabits the coasts of the Nicobar Islands); refers to Chemnitz, vol. 8, pi. 87, fig. 771; not P. inflata Phillips, 1836, a fossil. 1823 Pinna cancellata Mawe, Linnaean System of Conehol- ogy, p. 77, pi. 17, fig. 2 (Ceylon). 1841 Pinna chinensis Deshayes (in Cuvier) Le Regne Ani- mal (Disciples’ edition), Mollusques, pi. 85 (no lo- cality given). 1858 Pinna chemnitzii Hanley, Proc. Zool. Soc. London, p. 136 (Insulae Philippinae ) ; refers to Chemnitz, vol. 8, pi. 87, fig. 770; figured in Reeve, 1859, vol. 11, Pinna, pi. 1, figs, a, b. 1858 Pinna hystrix Hanley (in part), ibid., p. 226 (see synonymy of vexillum). 1858 Pinna hanleyi Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 8, fig. 15 (Amboyna). 1858 Pinna lurida Reeve, ibid., pi. 13, fig. 24 (Philippine Islands ) . 1858 Pinna penna Reeve, ibid., pi. 21, fig. 39 (Philippine Islands ) . 1858 Pinna serra Reeve, ibid., pi. 23, fig. 43 ( Moreton Bay [Australia] ). 1858 Pinna japonica ‘Hanley’ Reeve, ibid., pi. 25, fig. 47 (Japan) [attributed to Hanley in error]. 1858 (August) Pinna strangei ‘Hanley’ Reeve, ibid., pi. 27, fig. 52 (Moreton Bay [Australia]); Hanley, [Nov.] 1858, Proc. Zool. Soc. London, p. 254. 1858 (August) Pinna assimilis ‘Hanley’ Reeve, ibid., pi. 31, fig. 59 (Raines’s Island, Torres Straits); Hanley [Nov.] 1858, Proc. Zool. Soc. London, p. 255. 1891 Pinna lischkeana Clessin, Conchylien Cabinet, vol. 8, part 1, Malleacea, p. 73, pi. 28, fig. 1 ( Japan, Yoko- hama ) . 1891 Pinna vespertina ‘Reeve’ Clessin, ibid., p. 90, pi. 39, fig. 2 [= Atrina pectinata Linne; Clessin by error transposed Reeve’s figs. 43 and 44], 1922 Pinna japonica ‘Hanley’ Yokoyama, Joum. College of Science, Imperial Lhiiversity of Tokyo, vol. 44, Art. 1, p. 185, pi. 15, fig. 8 ( Shi to; Oji in Musashi); 1925, ibid., vol. 45, Art. 5, p. 28, pi. 6, fig. 7 ( Shirado Pliocene ) . 1953 Atrina (Servatrina) teramachii Habe, Illustrated Cata- logue of Japanese Shells, vol. 1, p. 192, pi. 24, fig. 1, pi. 25, fig. 5 (Tosa Bay, Shikoku). 1953 Atrina (Servatrina) kinoshitai Habe, ibid., p. 193, pi. 24, fig. 2 [not fig. 4 as in text] (off Wakayama Pref., Honshu). 1961 Atrina (Servatrina) lamellata Habe, Coloured Ills. Shells of Japan (II), Osaka, p. 117, pi. 52, fig. 5; appendix, p. 37. N omenclature — A discussion of the nomencla- torial history of Servatrina pectinata Linne was given by Dodge (1952, pp. 225-226). I agree with his analysis and can only repeat that it is the great degree of variation in this species which has re- sulted in the large number of names which must be synonymized with it. Winckworth (1929) considered Pinna penna Reeve to be a synonym of Atrina (Servatrina) ser- rata Sowerby of the Western Atlantic and not from the Philipjhnes as stated by Reeve. Winckworth ( 1936 ) later retracted this statement because Pra- shad (1932, p. 137 and plate 4, figs. 14, 15) claimed to have found Atrina penna Reeve in the Philip- pines. The specimen figured by Prashad, as well as that figured by Reeve, are young Servatrina pecti- nata Linne. The imbricate sculpture which is often so evident in the young shells may be seen in many older specimens where it becomes considerably worn away on the anterior portion of the shell but often remains to form a roughened ventral slope. The confusion generated by Hanley (1855, p. 149) regarding the similarity between Atrina fragilis Pennant and pectinata Linne may be easily dis- persed. Not only are the two species geographically isolated, the former in the Eastern Atlantic and the latter in the Indo-Pacifie, but each is included in a different subgenus on the basis of its nacreous layer: fragilis in Atrina and pectinata in Servatrina. Therefore the type locality “in India” cited by Linne is correct and pectinata is without doubt an Indo-Pacific species. The name inflata is credited by both Dodge (1952) and Winckworth (1929) to Wood, 1818. As noted earlier by Sherborn ( 1927 ) and later by Winckworth (1936, p. 122) P. inflata was described by Dillwyn, 1817. Pinna [Atrina (Servatrina)] cumingii ‘Hanley’ Reeve, (May) 1858 was assigned the locality “Aus- tralia” by Reeve. Hanley (Nov., 1858) cited “Pe- ruvia” as the type locality. The species is indeed from the Eastern Pacific and a synonym of Atrina (Servatrina) maura Sowerby (also see Winckworth, 1929, p. 291). Pinna minax Hanley, 1858, whose type locality was cited by Hanley as: “Nov. Guinea? Mexico?” is also very probably synonymous with maura. Winckworth (1929) tentatively considered Pinna (Atrina) strangei Reeve a valid species but stated that he had seen no specimens from India or Cey- lon, the areas with which he was seriously con- cerned at the time. Photographs of the holotype of strangei (not published here) show this species to be in all probability Atrina (Servatrina) pectinata [53 - 600] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 215 Linne. Internally, a large oval posterior adductor muscle scar is present located well within the pos- terior border of the nacreous layer. Externally the sculpture is characteristic of pectinata, there being none of the secondary radiating ribs often visible in Atrina (Atrina) vexillum. There is also the obso- lete sculpture on the ventral slope which is seen so clearly in pectinata, although in the holotype the sculpture appears to be rather worn overall. The appearance of the type is that of a mature to older specimen, the shell of which has suffered repeated breakage and repair posteriorly (also see Pinna hystrix Hanley under the Nomenclature section of Atrina vexillum). Types — The location of the holotype of Pinna pectinata is unknown. Linne referred to Gualtieri, plate 79, fig. A, which is here selected as the type figure. The type locality was given by Linne as “in India.” Types of species described by Reeve and Hanley are in the British Museum (Natural His- tory). The type of P. lischkeana was said by Clessin to be in the Rolle collection which was probably dispersed. The types of Servatrina teramachii and kinoshitai, described by Habe are in the Zoological Institute, Kyoto University, Kyoto, Japan. The type figure here selected of Pinna inflata Dillwyn is Chemnitz, vol. 8, pi. 87, fig. 771. The locality of the type specimen of P. cancellata Mawe from Cey- lon is unknown to me, as is that of P. chinensis Deshayes. Records (see map, pi. 160) - CEYLON (MCZ). THAI- LAND: Trang (USNM); Praehuap Khiri Khan ( G. M. Moore, MCZ). MALAYA: Singapore (ANSP). KOREA: southern tip of Korea ( Cahn, 1951). CHINA: Hong Kong (ANSP). JAPAN: western Honshu (Cahn, 1951); Tokyo Harbor (ANSP; USNM); Tosa, Shikoku (ANSP); Kago- shima Gulf, Kyushu (USNM). PHILIPPINES: many lo- calities (see map, MCZ; ANSP; USNM). INDONESIA: Amboina, Moluccas (MCZ). AUSTRALIA: Broome, West- ern Australia (V. Orr, ANSP; MCZ); Torres Straits (USNM); Buchan’s Point, 17 miles north of Cairns (J. Kers- lake; Austra. Mus.; USNM); Brampton Reef, Bowen; South- port (both W. Old, Jr., USNM) all Queensland. NEW GUINEA: Samberbaba, Japen Id. (NSF); Merauke (MCZ). Fossil Records — Yokoyama (1922, 1925, see Synonymy ) recorded Pinna japonica Hanley ( probably = Atrina pectinata Linne) from the Pleistocene, Shito formation, of Oji in Musashi, Japan; and from the Shirado Pliocene. Atrina pectinata subspecies zelandica (Gray, 1835) (Pis. 160, 164) Range — New Zealand: North, South Islands, Stewart Island and The Snares (Powell, 1957). Remarks — This is the only member of the family Pinnidae known to inhabit New Zealand waters and therefore its identification is certain with relia- Plate 163. Atrina pectinata pectinata (Linne). Figs. 1-2. Pinna japonica Reeve, Japan. (96 mm. in length). Figs. 5-6. Syntype of Pinna assimilis Reeve, from Raines’s Island. Holotype of Pinna penna Reeve, Philippine Islands. (79 mm. Torres Straits. (135 mm. in length). Figs. 3-4. Holotype of in length). (British Museum (N.H. ) photos.) [53-601] 216 Servatrina J. Rosewater Pinriidae ble data. It is probable that pelagic young of Serva- trina pectinata reached New Zealand from Aus- tralia via the East Australian current or that young specimens were carried on floating debris. Since its arrival and the later establishment of isolated breed- ing populations in New Zealand, the subspecies zelandica has become distinct in this area. Servatrina zelandica may be distinguished from S. pectinata by the much more uniformly distrib- uted sculpture of semitubular spines adorning the ribs of the former species; it is usually more nar- rowly triangular, and the coloration in zelandica is usually lighter overall, occasionally with radiating bars of brownish purple, while in pectinata usually darker olivaceous hues predominate and no striping has been noted. Habitat — “Common on mud-flats; buried” (Pow- ell, 1957, Plate 15, fig. 16 caption). Description — Shell reaching 258 mm. ( about lOM inches) in length; elongate-triangular in outline and only moderately inflated; rather thin and frag- ile in structure; sculptured with radiating ribs which in unworn specimens are usually uniformly covered with short, upright to occasionally recurved semi- tubular spines. Shell translucent to nearly trans- parent; ranging from light to darker tan in color with only a trace of olive; occasionally with radiat- ing bars of purplish brown between ribs. Surface of Plate 164. Atrina pectinata zelandica (Gray). Neotype of Pinna zelandica Gray, New Zealand. Note the dark stripes posteriorly and generally uniform spinosity, characteristic of this subspecies. Specimen 205 mm. ( 8 inches ) in length. (British Mhseum (N.H.) photos.) [53 - 602] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 217 valves dully shining. Radial sculpture consists of from 14 to 30 radiating ribs which are easily dis- tinguished on the posterior slope, but which be- come nearly obsolete and very crowded on the ventral slope. Ribs on posterior slope usually with uniformly distributed spines which also become minute imbrications on ventral slope. Concentric sculpture of fine growth lines, convex posteriorly. Posterior margin truncate, projecting farthest pos- teriorly at junction of posterior and ventral slopes. Dorsal margin usually straight. Ventral margin nearly straight to only slightly concave anteriorly. Interior of valves light to darker, sometimes oliva- ceous, tan; with the purplish brown bars showing through. Nacreous layer iridescent occupying an- terior three-quarters of valve. Posterior adductor muscle scar moderately large, circular, located within posterior border of nacreous layer, never protruding beyond. Posterior border of nacreous layer usually rounded, but occasionally with a pos- teriorly directed pointed peak; with no distinct em- bayment. Primary hinge ligament thin, black, ex- tending from anterior end of shell to posterior border of nacreous layer on the dorsal margin; sec- ondary hinge ligament not colored, but evident in intact specimens whose dorsal margins are fused. Measurements (mm.) — length width 258 110 (large; New Zealand) 173 78 (medium; Hutt, New Zealand) 87 40 (small; Manukau Harbor, North Island) Synonymy — 1835 Pinna zelandica Gray [in Yate] New Zealand, p. 310 (New Zealand); not originally figured; neotype fig- ured in Reeve, Conchologia Iconica, vol. 11, Pinna, pi. 7, fig. 13 (Wilkins, 1953, p. 25). 1850 Pinna senticosa Gould, Proceedings Boston Society of Natural History, vol. 3, p. 312 (New Zealand); not figured; Gould, ( 1852, U. S. Exploring Exped., vol. 12, Mollusca and Shells, pp. 448-449) referred to fig. 574 in the Atlas ( 1856), a figure never included. 1873 Pinna zelandica Gray, Hutton, Catalogue of the Ter- tiary Mollusca and Echinodermata of New Zealand, p. 26 (Tertiary: Waganui ( U. ) ; Shakespeare Cliff; Awatere ) . 1873 Pinna lata Hutton, ibid., p. 26 (Tertiary: Cobden). 1873 Pinna plicata Hutton, ibid., p. 26 (Tertiary: Culver- den(?), in blue clay) [“a fan-shaped Fucoid”, Suter, 1914, p. 10; 1915, p. 53], 1873 Pinna distans Hutton, ibid., p. 26 (Tertiary: Caver- sham ) . Types — The holotype of Pinna zelandica Gray was apparently lost. A neotype (lectotype?) from the Cuming Collection, from New Zealand, was selected by Wilkins (1953, p. 25) and is in the Rritish Mu- seum (Natural History): B.M. 1952.9.16.24. The Plate 165. Fig. 1. Atrina cordata (Pritchard), Barwon River, near junction with Native Hut Creek, Victoria, Aus- tralia. Eocene. Holotype. (110 mm.) (from Pritchard, 1895, pi. 12, fig. 4.). Figs. 2-4. Atrina janjukiensis Crespin, Lakes Entrance Oil Shaft, Gippsland, Victoria, Australia. Middle Eocene. (Figs. 2-3. Holotype (140 mm.); Fig. 4. Paratype; from Crespin, 1950, pi. 17, figs. 18, 19, 20). Fig. 5. Atrina tateana Hedley, Adelaide and Aldinga Bay, South Australia. Miocene. Holotype. (130 mm.), (from Tate, 1886, pi. 12, fig. 9). (All copies of original plates; Smithsonian Instit. photos. ) [53 - 603] 218 Servatrina J. Rosewater Pinnidae specimen was figured by Reeve (1858), Concholo- gia Iconiea, vol. 11, Pinna, pi. 7, fig. 13. The loca- tion of the type specimen of Pinna senticosa Gould is unknown and it was presumably lost (personal communication, R. I. Johnson, 1960, who has in preparation a catalogue of the types of species de- scribed by A. A. Gould). Types of fossil species described by Hutton, P. lata, plicata and distans, are in the Colonial Museum, Wellington, New Zea- land. Records (see map, pi. 160) — NEW ZEALAND: North Island: beach near Devonport; Auckland (both ANSP); Manukau Harbor (A. W. B. Powell, MCZ; USNM) all Auckland Province; Hutt, Wellington Province (USNM). South Island: Pelorous Sound, northern Marlborough Prov- ince (MCZ). Forsterian Marine Province: [Otago, Stewart Id. and the Snares] (Powell, 1957, p. 77). Fossil Records — Hutton (1873, p. 26) described 3 new species of Pinnidae from the Tertiary of New Zealand. One of these, P. plicata, was later discovered to be “a fan-shaped Fucoid ( Suter, 1914, p. 10; 1915, p. 53). Although the types of Hutton’s species were not figured, if the remaining two are Pinnidae, they are undoubtedly part of Atrina pec- tinate zelandica Gray. According to Suter (1913, p. 893), zelanclica appears as a fossil in the Eocene, Miocene and Pliocene of New Zealand. Atrina (Servatrina) cordata (Pritchard, 1895) (PI. 165, fig. 1) Range — Eocene of Victoria, Australia. Remarks — It is probable that this species is closely related to Servatrina tasmanica Tenison- Woods. The original figure 4, here reproduced (pi. 165) shows a fossil having a shape similar to the Recent tasmanica. Synonymy — 1895 Pinna cordata Pritchard, Proceedings Royal Society of Victoria, n.s., vol. 7, p. 228, pi. 12, figs. 4-5 ( Eocene sandy limestones, Barwon River, near its junction with the Native Hut Creek. J. Betheras, collector). Atrina (Servatrina) janjukiensis (Crespin, 1950) (PI. 165, figs. 2-4) Range — From the Middle Miocene (Janjukian Stage) of Victoria, Australia. Remarks — Like the preceding species, this seems closest to Servatrina tasmanica Tenison-Woods and both may be merely earlier forms of the latter. However, neither fossil is sufficiently complete to assure its definite assignment to tasmanica. Synonymy — 1950 Atrina janjukiensis Crespin, Proceedings Royal Society of Victoria, n.s., vol. 60, p. 150, pi. 17, figs. 18, 19, 20 (Middle Miocene (Janjukian Stage): The Lakes Entrance Oil Shaft, Gippsland, Victoria). Atrina (Servatrina) tateana (Hedley, 1924) (PI. 165, fig. 5) Range — Tertiary (Miocene), South Australia, Australia. Remarks — The figure of Atrina tateana here re- produced (pi. 165) indicates that this species is also closely related to Atrina tasmanica Tenison- Woods. Synonymy — 1886 Pinna semicostata Tate, Transactions and Proceedings and Report Royal Society of South Australia (for (for 1884-1885) vol. 8, p. 122, pi. 12, fig. 9 (Older Tertiary: Oyster banks, Adelaide and Aldinga Bay); 1899, Transactions Royal Society of South Australia, vol. 23, part 2, p. 276 ( Miocene-Aldinga Bay). 1924 Atrina tateana Hedley, Records of Australian Museum, vol. 14, no. 3, p. 143 [new name for Pinna semi- costata Tate, not Pinna semicostata Conrad (=P. muricata Linne, a recent species)] Atrina (Servatrina) tasmanica (Tenison-Woods, 1876) (Pis. 160, 166, 167) Range — Southern and southeastern Australia and northern Tasmania. Remarks — The Tasmanian Atrina apparently is limited in distribution to the north coast of Tas- mania and southern and southeastern Australia. Superficially it appears nearest in its relationship to Atrina (Atrina) squamifera Sowerby of South Af- rica. However the subgeneric characters separate these species. In Servatrina tasmanica the posterior adductor muscle scar is placed well within the pos- terior border of the nacreous layer, but in A. squa- mifera the scar bulges beyond. As in other mem- bers of the Pinnidae, a superficial similarity is observed in the external sculpture of these two species. In both, well-defined tubular spines may develop on the radial ribs; the sculpture on the ventral slope becomes crowded, but small spines often persist, giving a prickly appearance to this portion of the valve. Other species of Atrina with which tasmanica might be confused are vex ilium, pectinata pectinata and p. zelandica. In pectinata and zelandica the numbers of rows of radiating ribs on the posterior slope are greater than in tasmanica (14,15-30 rather than 10-14). The subgeneric dif- ference easily separates mature specimens of Atrina [53 - 604] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Atrina 219 (Atrina) vexillum from Atrina (Servatrina) tasma- nica. Also, these species apparently do not occur within the range of tasmanica. It appears, there- fore, that Servatrina tasmanica is a valid and dis- tinct species which is endemic in the South Aus- tralian Region. Habitat — Lives in fairly deep water: 15-45 fath- oms (Hedley, 1924, pp. 152-153); probably buried in sandy mud with the posterior portion of the shell protruding from the substrate. Description — Shell reaching 250 mm. ( nearly 9/2 inches; [14 inches; Gabriel (1936)]) in length; tri- angular-wedge shaped in outline; moderately in- flated; rather fragile in structure and sculptured with radiating ribs which usually bear strong semi- tubular spines. Shell translucent, tannish brown in color; surface of valves shining. Radial sculpture consists of from 10-14 moderately prominent radi- ating ribs on the posterior slope; in older indi- viduals ribs may become nearly obsolete; sculpture on the ventral slope becomes crowded and impossi- ble to count. Ribs often smooth, but usually bear large, upright, occasionally twisted spines which are open posteriorly. Concentric sculpture of pos- teriorly convex lines of growth which sometimes overlap loosely giving the valves a scaly appear- ance. Posterior margin usually broadly rounded. Dorsal margin nearly straight to slightly convex. Ventral margin convex posteriorly, concave anteri- orly. Interior of valves tannish brown, with occa- sional dark brown blotches; smooth and shining. Nacreous layer iridescent, occupying anterior three fourths the length of valve. Posterior adductor muscle scar medium sized, subcircular, located well within posterior border of nacreous layer, never protruding beyond. Anterior adductor muscle scar smaller, located at anterior tip of nacreous layer. Posterior border of nacreous layer somewhat peaked centrally and sloping toward dorsal and ventral borders. Primary hinge ligament moderately thin, Plate 166. Atrina tasmanica (Tenison- Woods), Phillips Is- 2. Interior of right valve. Specimen 191 mm. (7.5 inches) in land, Victoria, Australia. Fig. 1. Exterior of right valve. Fig. length. (National Museum of Victoria photos.) [53 - 605] 220 S ervatrina J. Rosewater Pinnidae black and extending along dorsal margin from an- terior end of shell to posterior border of nacreous layer; secondary ligament not colored but evident in intact specimens whose dorsal margins are fused. Embryonic valves and animal unknown. Measurements (mm.) — length width 240 112 (large; San Remo, Victoria, Australia) 191 99.5 ( average; Phillip Island, Victoria, Australia ) 142 63 (small; North West Tasmania; lectotype) 133 72 (small; Adelaide, South Australia) Measurements of the center two specimens were kindly supplied by Dr. J. Hope Macpherson, Cura- tor of Molluscs, National Museum of Victoria, Vic- toria, Australia. Synonymy — 1876 Pinna tasmanica Tenison-Woods, Proceedings Royal Society of Tasmania [for 1875], p. 161 (on the north coast only [of Tasmania] W. Legrand ) [not figured], 1924 Atrina tasmanica var. dumosa Hedley, Records of Aus- tralian Museum, vol. 14, p. 153 (South Australia: Tapley Shoal, St. Vincent Gulf, 15 fathoms (Mathews and McDougall)) [not figured]. 1938 Atrina dumosa Hedley, Cotton and Godfrey, The Mol- luscs of South Australia, pt. 1, Peleeypoda, p. 85, fig. 71. Types — Lectotype selection: Pinna tasmanica Tenison-Woods, 1875. Tenison-Woods indicated that the specimens of Pinna tasmanica upon which his description was based were collected by Le- grand. A designated type has not been found in the National Museum of Victoria, the South Australian Museum, nor the Australian Museum, Sydney. In- quiries kindly made for me by Dr. J. Hope Mac- pherson have failed to locate the type of tasmanica in Tasmania. It appears that Tenison-Woods never designated a type and that the Legrand collection has been lost or dispersed. However, the National Museum of Victoria has in its collection three specimens from within the type locality, “on the north coast only,” received in August 1876, follow- ing the publication in March (fide Hedley, 1924, p. 152) of the description of tasmanica. It is possible that these specimens were among the material ex- amined by Woods and therefore may be consid- ered as syntypes. One of these is here selected as lectotype of the species Pinna tasmanica Tenison- Woods: National Museum of Victoria, Catalogue no. F21384A; length 142.5 mm., width 63 mm., depth 24 mm.; from “North West Tasmania”; re- ceived August, 1876; it is here figured, pi. 167, figs. 1, 2. It is a young specimen and fulfills the portion of Tenison-Woods’ description: “ribs sometimes . . . subnodose.” A more mature and “typical” speci- men is also figured, pi. 166, figs. 1, 2 (Nat. Mus. Victoria Cat. no. F21385; photographs courtesy of Dr. J. Hope Macpherson and Nat. Mus. of Victoria). The type locality of Pinna tasmanica is here re- stricted to Circular Head, Wellington County, [northwestern] Tasmania, a locality from which specimens of this species have been collected (see Tate and May, 1901). The location of the holotype of Atrina tasmanica var. dumosa Hedley, 1924 is unknown. A lectotype has been selected by Cotton ( South Australian Mollusca ( in press ) ) . Notification of his selection was given (in lift., 1960) and is here quoted with his permission: “The ‘subspecies’ Atrina tasmanica dumosa was described by Hedley from Australian Museum specimens and if a holotype were chosen it should be there. I have selected the specimen fig- ured here, (Peleeypoda 1938, fig. 71) measuring height 132 mm., length 75 mm., from Tapley Shoal, Gulf St. Vincent, as Lectotype, D. 14160, S.A. Mu- seum.” Records (see map, pi. 160) — AUSTRALIA; King Island; Circular Head, Wellington County; Port Sorrell, Devon County, all Tasmania (all Hedley, 1924). Norali Head, Northumberland County, New South Wales (Hedley, 1924). San Remo, near Melbourne, Victoria (ANSP). Tapley Shoal, St. Vincent Gulf (Hedley, 1924); Adelaide (ANSP), both South Australia. Plate 167. Atrina tasmanica. Lectotype of Pinna tasmanica Tenison-Woods, northwest Tasmania. Specimen 142.5 mm. (about 5V2 inches) in length. (National Museum of Victoria photo. ) [53 - 606] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 S treptopinna 221 Genus Streptopinna von Martens, 1880 Type: Streptopinna saccata ( Linne, 1758) This unique genus is found only in the Indo- Paeific faunal region where it is monotypic includ- ing the single Recent species Streptopinna saccata Linne. Von Martens (1880) introduced Strepto- pinna as a subgenus of Pinna and was followed in this use by Winckworth (1929). However, the dif- ferences between Streptopinna, Pinna and Atrina are considerable and each is accorded full generic status here. Major generic characters appear to be best demonstrated in the shell. They are: the crowded nacreous layer which occurs only in the dorsal anterior portion of the shell; the appearance of the small anterior adductor muscle scar on an elevated shelf; and the peculiar twisted condition of the posterior portion of the valves. A more com- plete description follows in the treatment of the species, saccata. Unfortunately the embryonic valves of Strepto- pinna are unknown. This is unusual, considering the hardy nature of the umbonal area of this group. It would be of extreme interest to determine their structure especially in the light of a similarity in the configuration of the nacreous layers between Streptopinna and Crenulata. The two groups are either extremely convergent in this feature or else they may be related through common ancestral stock. One southeastern Australian Tertiary fossil possi- bly belonging to this subgenus is included here. Synonymy — 1880 Streptopinna von Martens, Beitriige znr Meeresfauna der Insel Mauritius und der Seychellen p. 318. Type by monotypy: Streptopinna saccata (Linne, 1758). Streptopinna? reticosa (Chapman, 1912) (PI. 168) Range — Tertiary; King Island, Tasmania. Remarks — Chapman (1912, p. 48) compared reticosa with P. cordata Pritchard, but the latter is probably a Servatrina related to tasmanica. From the figure given by Chapman (pi. 168), if the fossil is a pinnid, it appears to be a small saccata- like form. However, this placement is doubtful. Synonymy — 1912 Pinna reticosa Chapman, Memoirs National Museum, Melbourne (Victoria, Australia), no. 4, p. 47, pi. 6, fig. 8 (Tertiary: polyzoal limestone of Seal River, King Island). Plate 168. Streptopinna ? reticosa (Chapman), Seal River, King Island, Tasmania. Tertiary. Holotype, x 4. (from Chap- man, 1912, pi. 6, fig. 8). (Smithsonian Institution photo.) Streptopinna saccata (Linne, 1758) (Pis. 136, 169-171) Range — East Africa to eastern Polynesia. Remarks — Streptopinna saccata Linne is widely distributed in the Indo-Pacific ( see E. A. Smith, 1903, p. 597). The immature forms of this species are hardly distinguishable exteriorly from young, rather obsoletely sculptured members of the genus Atrina. This condition continues through about the first third of the growth of shell as compared to the normal eventual adult size. Thereafter, growth pro- ceeds erratically and the shell of adult specimens may assume an almost unbelievable degree of con- tortion. There is apparently no predictable form which may be assumed, this lack of uniformity be- ing a basic character for the species. Certain speci- mens, apparently living under favorable conditions may, indeed, appear quite normally proportioned and identification is questionable until the defini- tive characteristics of the internal nacreous layer are observed (see Description) . The reason for the grotesque appearance of saccata has its basis in the ecology of the species. It is reported to live under [53-627] 222 Streptopinna J. Rosewater Pinnidae and between rocks in rocky tide pools and there- fore the growtli form of the shell is dictated by tbe curvature of the particular rocks between or under which the young saccata settles and commences its normal existence. The viscera are crowded into the dorsal portion of the anterior normal third of the shell. Posterior to the rather small posterior adduc- tor muscle the mantle must extend the additional two thirds the length of the shell to build and main- tain the shell and bring its edges into contact with the environment to form the incurrent feeding-re- spiratory and excurrent waste canals. The valves of S. saccata are often so twisted and their growth along the ventral margin so thickened that they actually appear to be fused here as well as along the dorsal margin. There is formed by this fusion of the ventral margin a veritable shell sack in which the animal is contained and, of course, the name “ saccata ” is derived from this fact (also see Jack- son, 1890, pp. 385-386). Although the species is widespread and is found in a great variety of shapes there is no basis for the establishment of more than one species. This is a valid and striking case of environmentally-caused, phenotypic variation in mollusks. Von Martens ( 1880, p. 318 ) remarked that Streptopinna saccata lacks a byssus. This is contra- dicted by the presence of a moderately large byssus in preserved specimens examined. Attachment of the byssal threads to coral and shell debris indi- cates that this species does protrude the byssus and anchors itself to bottom objects as do other species of the Pinnidae. Plate 169. Lectotype of Streptopinna saccata inusitata Ire- dale, Michaelmas Cay, off Cairns, Queensland. Specimen 97 mm. (about 3% inches) in length. Fig. 1. Exterior of right valve. Fig. 2. Interior of right valve. (Australian Museum photo. ) Holthuis ( 1952, p. 109 and the last page of in- dex) noted from the literature the presence of the palaemonoid shrimp, Anchistus custos (Forskal), in the mantle cavity of Streptopinna saccata from the Red Sea. Winckworth ( 1929, p. 282 ) and Dodge ( 1952, p. 229) stated that S. saccata does not move upward during growth as evidenced in other species of Pinnidae by the series of regular septa laid down anterior to the anterior adductor muscle scar. Obvi- ously, the body of the animal does enlarge posteri- orly, and lines of the former position of the pos- terior adductor muscle scar have been noted in specimens examined. Such lines are also in evi- dence on the anterior adductor scar, and much crowded septa have been noted in complete speci- mens. It is quite true that the umbonal area seems to survive more nearly intact in this species, whereas it is often worn away in others, perhaps because of its way of life and the lesser degree of its burrow- ing activities. Habitat — Under and between rocks in rocky tide pools; also, in cavities of coral (von Martens, 1880, P-318). Description — Shell reaching 235 mm. (9/i inches; not allowing for curvature) in length; roughly tri- angular in outline: sometimes elongate, but often broad and usually severely contorted posterior to the anterior one third which is normally pinnoid; predominant total flexure may be either to right or left, often producing inequivalve condition, the convex valve often being slightly larger; attaining moderately heavy structure and sculptured with radiating ribs. Shell translucent, occasionally trans- parent in young specimens; ranging in color from grayish white through tan to dark reddish brown; often light tan anteriorly with the darker color pos- teriorly. Surface of valves dully shining. Radial sculpture consists of from 5 to 12 radiating ribs on the posterior slope ( in some specimens posterior border may have double the number of ribs which show anteriorly, possibly due to injury); ventral slope without ribs, roughened. Ribs usually smooth, but in occasional specimens may bear a few coarse lamellate spines. Concentric sculpture of irregular lines of growth which in some specimens are spaced rather regularly. Posterior margin variable, fre- quently fractured and truncate, sometimes convex posteriorly. Dorsal margin variable, often gro- tesquely twisted. Ventral margin also variable pos- teriorly; anteriorly ventral margin greatly bulges just posterior to umbos; umbonal area usually sud- [53 - 6281 September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Streptopinna 223 denly considerably narrower than posterior portion of valves. Interior of valves grayish white to dark brown; anterior portion often lighter than posterior; surface very irregular, spaces between the external ribs quite clearly defined as “internal ribs”; inte- rior surface shining. Nacreous layer smoky white, crowded into anterior, dorsal one third of valve forming a small triangle with its base toward pos- terior border of valve. Anterior pallia] retractor muscle scar delimited from rest of nacreous layer, small, circular and located centrally in valve ven- tral to main part of nacreous area. In some speci- mens additional small patches of irregular nacreous material may appear ventral to nacreous layer, but this appears to be abnormal. Posterior adductor muscle scar small to medium sized, located in pos- tero-ventral corner of nacreous layer. Anterior ad- ductor muscle scar small, located on an elevated shelf at anterior tip of valve and touching anterior major portion of nacreous layer only at its postero- dorsal angle. Posterior border of nacreous layer usually with an embayment dorsal to posterior ad- ductor scar; nacreous area widest here. Ventral border proceeding from widest portion to a narrow tip which touches anterior adductor scar. Dorsal border contiguous with dorsal shell margin. Pri- mary hinge ligament thin, black, extending from umbos to posterior border of nacreous layer; sec- ondary hinge ligament not colored but evident in intact specimens whose dorsal margins are fused. Ventral margins of valves often so thickly produced and incrusted with organic material and debris that they present a virtually fused appearance forming a sack-like shell. Embryonic valves missing in all specimens examined. Pallial organ particularly long and vermiform. “Eyes of Will” fairly conspicuous, darkly pigmented, located in fold between middle and inner mantle lobes (see Introduction, Remarks on Anatomy) . Measurements (mm.) (not allowing for curvature of valves) — length width 235 78 (large; Keokea, Hilo, Hawaii) 181 98 (large; Suva, Fiji) 124 51 (average; Torres Straits) 112 63 (average; Kahaluu, Hawaii) 78 18 (average; Marongas Id., Philippines) 36 20 (small; Kwajalein Atoll) Synonymy — 1758 Pinna saccata Linne, Systema Naturae, ed. 10, p. 707 (in M. Mediterraneo, Indico); refers to Rumphius, pi. 46, fig. N; and to Gualtieri, pi. 79, fig. F [not P. saccata Linne Chemnitz, 1785, pi. 90, fig. 779 -P. muricata Linne]. 1786 Pinna lubrica Solander, Catalogue of the Portland Museum, pp. 61, 139 (nomen nudum); fide Dill- wyn, 1817, Descriptive Catalogue, vol. 1, p. 331: = vitrea Gmelin [= Streptopinna saccata Linne], 1791 Pinna vitrea Gmelin, Systema Naturae, ed. 13, p. 3366 (in Oceano indico); refers to Chemnitz, vol. 8, pi. 87, fig. 772 (= S. saccata L.); also Gualtieri, pi. 78, fig. C, and pi. 79, fig. 1 (both unrecognizable); [not Pinna vitrea Roding, 1798 = Atrina pectinata L.]. 1837 Pinna nuttallii Conrad, Journal Academy of Natural Sciences, Philadelphia, vol. 7, p. 244, pi. 19, fig. 4 ( Inhabits muddy marshes in the Sandwich Islands ) . 1858 Pinna elongata Reeve, Conchologia Iconiea, vol. 11, Pinna, pi. 4, fig. 6 (no locality given); Winckworth (1929) Proceedings Malacological Society of Lon- don, vol. 18, pp. 289, 292; [not P. elongata Roding, 1798 (=P. rudis Linne)]. Plate 170. Geographical distribution of Streptopinna sac- cata ( Linne ) . [53 - 629] 224 Streptopinna J. Rosewater Pinnidae Plate 171. Streptopinna saccata (Linne)- Figs. 1-2. Speci- men from Keokea, Hilo, Hawaii. (210 mm. or about 8V4 inches in length). Figs. 3-4. Immature specimen from Yaku Shima, Osumi, Japan. Specimen 60 mm. (about 2V2 inches) in length. Figs. 2 and 4. Internal views of valves with nacre- ous areas, anterior pallial retractors and anterior adductor muscle scars outlined in black. ( Smithsonian Institution photos. ) 1927 Streptopinna saccata inusitata Iredale, Australian Zool- ogist, vol. 4, p. 333, pi. 46, figs. 9-11 (Caloundra, Moreton Bay, Queensland [Michaelmas Cay, off Cairns, Queensland; see Types]). 1932 Pinna saccata var. similis Turton, Marine Shells of Port Alfred, p. 218, pi. 56, fig. 1518 (Port Alfred [South Africa] ). 1932 Pinna anomioides Reeve’ Turton, ibid., p. 218, no. 1519 (South Africa). ?1932 Pinna aenigmatica Turton, ibid., p. 219, pi. 56, fig. 1521 (Port Alfred) [probably a species of Malleus]-, YVinckworth (1936) Proc. Mai. Soc. London, vol. 22, p. 21. Nomenclature — Because of the wide variety of forms which may be assumed by this species it is surprising that a larger number of synonyms has not come into being. Surprisingly there are a rela- tively small number and some of these are ques- tionable as to identity. Pinna elongata Reeve is listed by Winckworth (1929) as P. saccata variety. It is doubtful that Reeve intended to create a varie- tal name in this case, and that the word “elongata” is a descriptive Latin diagnosis for the form P. nut- tallii Conrad. Pinna aenigmatica Turton is called a “gruesome piece of beach refuse” by Winckworth (1936) who placed it in the synonymy of saccata, “if it be a Pinna at all.” It quite probably is not a member of this family. The strong ridge “down the inside of one of the valves” which has the “hinge at the mid- dle of the wider end” (Turton, 1932) strongly sug- gests, as does the plate, a species of Malleus. Pinna nuttallii Conrad was the specific name as- signed the Hawaiian form of saccata ; Streptopinna saccata inusitata Iredale was named as a Queens- land variety. As Pinna saccata is a very widespread species ranging from east Africa to Polynesia it is doubtful that these names have any standing even on a subspecific basis. Types — The type specimen of Pinna saccata Linne, 1758 was said to be in the Linnaean collec- tion according to Hanley ( 1855 ) and Winckworth (1929). Dodge (1952) does not mention having found the type, and, therefore, its presence may be questioned. Linne referred to Rumphius, pi. 46, fig. N; and Gualtieri, pi. 79, fig. F. Of the two the Rumphius reference is the better since it represents [53 - 630] September 28, 1961 INDO-PACIFIC MOLLUSCA, vol. 1, no. 4 Streptopinna 225 a specimen which is more mature and character- istic of the species. Rumphius, pi. 46, fig. N is here selected as the type figure. The type locality origi- nally given by Linne, “in M. Mediterraneo, Indico,” is here restricted to Amboina, Moluccas, from which the specimen figured by Rumphius undoubtedly came. Streptopinna saccata is not known from the Mediterranean. The type figure of Pinna vitrea Gmelin here selected is Chemnitz, vol. 8, fig. 772; the figures of Gualtieri, pi. 78, fig. C, and pi. 79, fig. E, are unrecognizable as any distinct species of Pinnidae. The types of species described by Tur- ton: P. similis and aenigmatica are in the Oxford Museum according to Turton. The type of Pinna nuttallii Conrad should be in the Academy of Natu- ral Sciences, Philadelphia, although it could not be found during a visit there. As P. elongata Reeve was probably not actually proposed by Reeve as a species but only used descriptively, there are no types available. Lectotype Selection: Streptopinna saccata inusi- tata Iredale was described from three specimens: two from Michaelmas Cay [off Cairns], North Queensland, and one from Caloundra [north of Brisbane], South Queensland. The three specimens were figured by Iredale without strict indication of a holotype. The lectotype here selected of Strepto- pinna saccata inusitata is the syntype specimen shown by Iredale (1927, pi. 46, fig. 11 ), Australian Museum Catalogue number C.53671; approximate measurements: greatest length 97 mm., width 59 mm.; type locality: Michaelmas Cay, off Cairns, Queensland [erroneously stated by Iredale to come from Caloundra, fide D. F. McMichael, Austr. Mus.]; here figured, pi. 169, figs. 1, 2 (photograph courtesy of the Australian Museum, Sydney). Records (see map, pi. 170) solid dots: specimens exam- ined; open circles: trom the literature — SO llTH AfRICA: Durban (MCZ). MOZAMBIQUE: Inhaca Id., Delagoa Bay (MCZ). ZANZIBAR: outer reef, Kiwengwa (NSF). GULF OF OMAN: Muscat, Oman (ANSP)'. MADAGASCAR: Nossi Be (ANSP; MCZ). SEYCHELLES; MAURITIUS; RE- UNION (all Von Martens, 1880). CEYLOiX: Galle ( Winck- worth, 1929). JAPAN: Shikoku; Kyushu; Honshu (all Habe, 1953a); Yaku Shima, Osurni (ANSP; USNM). FOR- MOSA (Habe, 1953a). PHILIPPINES: 26 miles southwest of Corregidor, Luzon Id. (USNM); Magallanes Bay, north end Mactan Id., eastern Cebu Island (ANSP); Bongao Channel, southwest end Sanga Sanga Id., Sulu Archipelago (ANSP). INDONESIA: Amboina (ex. Siboga Expedition, MCZ). Koeta Beach, Bali (MCZ). COCOS-KEELING IS- LANDS (USNM). AUSTRALIA: Torres Strait (USNM); Long Reef, New South Wales (Australian Museum; USNM). NEW GUINEA: IV2 miles southwest of Biak, Schouten Ids. (NSF; USNM); Abroeki Isle, Maransabadi Id., Aoeri Ids., Geelvink Bay (NSF). FIJI ISLANDS: entrance to Suva Harbor, Viti Levu (USNM). PALAU ISLANDS: Ngadarak Reef, north of mouth of Malakal Pass (USNM). CARO- LINES: Yap Id. (USNM). MARSHALLS: lagoon, north end Eniwetok Atoll (J. B. Burch, University of Michigan); Namu Id., Bikini Atoll ( J. P. E. Morrison, USNM); Ine Vil- lage, Arno Atoll ( R. \V. Hiatt, USNM). GILBERT IS- LANDS: Kingsmill Group (MCZ). HAWAII: Pearl and Hermes Reef (MCZ; USNM); Kahaluu, north Kona, Island of Hawaii ( C. M. Burgess, USNM). LINE ISLANDS: Flint Id. (ANSP). SAMOA: Vaoto, Vailele Bay, Upolu Id. (NSF). SOCIETY ISLANDS: Port Du Bourayne, southeast of Vaiorea Id., outer reef, Huahine ( Bredin-Smithsonian In- stitution Expedition, USNM). TLTAMOTUS: Marutea du Sud ( Dautzenberg and Bouge, 1933, p. 434 ) . INDEX TO PINNIDAE NAMES IN VOL. 1, NO. 4 adusta Dillwyn, 205 aenigmatica Turton, 224 aequilatera von Martens, 190 aequilatera YVeinkauff, 190 afra Sowerby, 209 alfredensis Bartsch, 209 ampla Sowerby, 178 angustana Lamarck, 196 angustana ‘Lam.’ Reeve, 196 anomiodes ‘Reeve’ Turton, 224 asakuraensis Nagao, 188 assimilis Reeve, 211, 214 atrata Clessin, 197 Atrina, 203 atropurpurea Sowerby, 196 attenuata Reeve, 196 Aviculipinna, 178 banksiana Iredale, 205 bicolor Gmelin, 193-199 bifida Rafinesque, 178 blanfordi Boettger, 193 bullata Gmelin, 184 cancellata Mawe, 214 capensis Krauss, 209 carnea Gmelin, 178 caviterga Medley, 197 cebuensis Elera, 190 ehautardi Nickles, 178 chemnitzii Hanley, 214 Chimaera, 187 Chimaeroderma, 187 ehinensis Deshayes, 214 cochlearis H. Fischer, 197 cordata Pritchard, 218 cumingii Clessin, 197 cumingii Reeve, 197 Curvula, 178 Curvulites, 178 Cyrtopinna, 187 deflecta Perry, 185 delsa Iredale, 190 deltodes Menke, 196 densecostata Turton, 197 digitiformis Linne, 184 distans Hutton, 217 dolabrata Lamarck, 196 dumosa Hedley, 220 electrina Reeve, 196 [53-631] 226 Index J. Rosewater Pinnidae elongata Reeve, 223 elongata Roding, 223 epica Jousseaume, 197 euglypta ‘Hanley’ Reeve, 196 Exitopinna, 188 exquisita Dali, B. and R., 190 exusta Gmelin, 205 fimbriatula Reeve, 196 Habella Hall, 177 flexicostata McCoy, 177 fragilis Pennant, 178 fumata ‘Hanley’ Reeve, 196 gouldii ‘Hanley’ Reeve, 205 granulata Sowerby, 178 gubernaculum Roding, 205 hanleyi Reeve, 214 hawaiensis Dali, B. and R., 190 howensis Iredale, 190 hystrix Hanley, 205 incurva Gmelin, 200-202 incurvata Born, 201 incurvata Sowerby, 201 inermis Tate, 196 inflata Dillwyn, 214 inflata Phillips, 214 inflata Roding, 214 inusitata Iredale, 224 isosceles Hedley, 197 italicus Sherborn, 184 janjukiensis Crespin, 218 japonica ‘Hanley,’ 214 japonica Reeve, 214 kinoshitai Habe, 214 kraussii Hanley, 209 Laevipinna, 178 lata Hutton, 217 linguafelis Habe, 205 lischkeana Clessin, 214 lobata Linne, 184 lubrica Solander, 223 lurida Reeve, 214 madida Reeve, 196 marginata Lamarck, 185 maura Sowerby, 178 menkei ‘Hanley’ Reeve, 196 molluccensis Clessin, 197 muricata Linne, 188-193 mutica Reeve, 196 natalensis E. A. Smith, 197 nebulosa Solander, 190 nigra Dillwyn, 205 nigricans Solander, 205 nigrina Lamarck, 205 nobilis Linne, 178 nodosus Lycett, 178 nuttallii Conrad, 223 oahua Dali, B. and R., 205 Oxysma, 178 pachyostraca Davies, 203 Palaeopinna, 177 papyracea Gmelin, 190 pectinata Linne, 211-215 penna Reeve, 214 pennacea Linne, 184 Pennaria ‘Browne’ Morch, 203 philippensis von Martens, 190 philippinensis Reeve, 190 Pinna, 187 Pinnarius, 187 Pinnigena, 178 Pinnula, 187 plicata Hutton, 217 prisca Muenster, 178 punjabensis Eames, 188 Quantulopinna, 187 recta Dali, B. and R., 205 regia ‘Hanley’ Reeve, 196 rembangensis Martin, 188 reticosa Chapman, 221 rigida Solander, 178 rollei Clessin, 197 rostellum Hanley, 196 rotundata Linne, 184 rudis Linne, 178, 223 rufanensis Turton, 209 rugosa Sowerby, 178 rumphii ‘Hanley’ Reeve, 190 saccata Linne, 221-225 sanguinea Gmelin, 184 sanguinolenta Reeve, 185 scapula Hedley, 197 segmenta Turton, 209 semi-costata Conrad, 190 semicostata Reeve, 190 semicostata Tate, 218 seminuda Lamarck, 178 senticosa Gould, 217 serrata Sowerby, 178 Servatrina, 211 shekhanensis Eames, 188 similis Turton, 224 spatula McCoy, 178 squamifera Sowerby, 207-210 Stegoconcha, 178 strangei ‘Hanley’ Reeve, 214 strangei Prashad, 190 Streptopinna, 221 striata Rafinesque, 178 striolata Sherborn, 184 stutchburii Reeve, 196 Subitopinna, 188 Sulcatipinna, 177 tasmanica T. -Woods, 218-220 tateana Hedley, 218 tenera Solander, 190 tenuis Habe, 205 teramachii Habe, 214 Trichites, 178 trigonalis Pease, 190 trigonium Dunker, 196 tuberculosa Sowerby, 178 ultra Iredale, 197 vespertina Reeve, 196 vespertina ‘Reeve’ Clessin, 214 vexillum Born, 204-207 virgata Menke, 190 vitrea Gmelin, 223 vitrea Roding, 214 whitechurchi Turton, 209 zebuensis Reeve, 190 zelandica Gray, 215-218 INDEX TO COMMENSAL CRUSTACEA IN PINNIDAE Anchistus custos, 188,195,204,222 Commensalism, 176 Conchodytes biunguiculatus, 195 Paranchistus ornatus, 195 Conchodytes nipponensis, 213 Pinnotheres, 176 [53 - 632]