Natural History, ^m.tyiiu&

University of

Collation, Revision,

and Additions







Published by the University of Illinois Press under the auspices of the Graduate School. by the University of Ulinois Press. Copyright. Manufactured in the United States of America . Permission for rejiroduction in any form mav be obtained from the Publisher. 1945.

eighty species and races of the family have been examined. There are some shell characteristics. the Basommatophora are still in a condition of more or less chaos as regards classification. represented by more than 725 specimens.PREFACE and the economic significance from a medical standpoint. large number of species represented. For the anatomical studies recorded in this volume. affording many cases of parallel development. THE WIDE DISTRIBUTION. which are of value in classification when used in conjunction with anatomical features. It is the largest and most conspicuous of the families of Basoramatophorous pulmonate snails. The shell alone cannot be relied upon for this purpose because some of the characteristics are often misleading. however. of a favorable nature. Pilsbry and his co-workers. The study of the internal organization of the family Planorbidae has shown that. as the presence of lamellae within the aperture. That it is still in general use after an interval of more than thirty years is evidence that its reception has been. part I on the anatomy and classification of the family. constitute the family Planorbidae as one of the most important groups of fresh-water Mollusca. The anatomy of six additional species has been taken from the literature. on the whole. . In 1911 the writer published a monograph of the Lymnaeidae of North and Middle America. which feature is found in a number of groups which are unrelated anatomically. and other groups might follow the publication of the Lymnaeidae volume if it should meet with the approval of the conchological public. Helisomatinae 41 Planorbulinae 10. Physa. as intermcfliate hosts of flukes or trematode worms. and part II* on descriptions of the species of the family inhabiting North and South America and the West Indies. The condition of this large group as far as classification is concerned is little better than that of the land snails seventy-five years ago. the anatomy gives the only true key to a natural classification. A large majority of the known genera and subgenera are represented and it is confidently believed that additional anatomical study will not greatly alter the classification herein set forth. basing the classification on characteristics of the genitalia and radula. Unlike the terrestrial pulmonates (Stylommatophora Pulmonata) which have been brought to a high state of precise classification from the anatomical studies of Dr. The present volume on the Planorbidae was begun some twenty-five years ago and has been in active preparation for the past ten years. Henry A. In that work it was suggested that studies on Planorbis. As the work progressed it became evident that adequate treatment of the subject could not be given in a volume similar to that on the Lymnaeidae and so the work has been divided into two parts. Segmentininae 11. as in the case of the land snails. The number of species examined has been divided among the four subfamilies as follows: Planorbinae 19. all of the monographs and many of the local studies being based wholly or partly on characteristics of the shell with little regard for relationships which might be gained by a study of the organization of the animal. ix . *See title page of Part II (page 213) for statement regarding change in scope of this part.

are included. To have done this would have lengthened the work to a large degree without adding greatly to its value. The list of species included in each genus or subgenus has no pretension to completeness. Only the most important citations. Every generic or group name has been given careful study to ascertain its true relationship. It does not at present appear to be of any advantage to recognize the terms section or subsection for any of the groupings. Some of the finer details have been filled in by hand on the camera lucida figure. but in the majority of the figures the author has endeavored to portray the form of the organs as they appeared to him under the microscope. Some of the figures are diagrammatic. The radulae have all been examined under a Zeiss microscope with magnification up to about 1000 diameters. In order that the features of the different organs might be fully known they have been separated and otherwise dissected to bring out all relationships. As additional material is examined some of the species listed may be shifted to other groups and additional species may be included.X The Molluscan Family Planorbidae The anatomical drawings in this work have all been made by the author with the aid of the camera lucida. In the matter of bibliographic citations it has not been deemed necessary to include all references to works in which a genus or other group is listed or described. and the outlines and much of the detail of the figures are correct from the standpoint of dimensions. Until a greater number of species have been examined for the internal organization this division appears best. which have been ignored or placed in the synonymy by previous authors. In many cases sections have been made through organs or groups of organs the better to show structural conditions. including the first one in which the generic name appears. none as sections. only those species being listed which have either been examined anatomically or appear from the form of the shell to be related. It will be noted that many groups. have been recognized in this work. Some of the groups herein considered genera may later be placed as subgenera. Only the shells of the type species of the different genera and subgenera are figured in the plates prepared for Part I. January. All groups described in this work are arranged under the heading of a genus or of a subgenus. This is particularly true of Asiatic groups which have been little known anatomically. 1942 . All dissections were made with a Spencer binocular microscope with magnification up to seventy-five diameters.

Louis. Albany Dr. Johns Hopkins University. School of Tropical ^ledicine. Louis Botanical Gardens. Miss Ruth E. Johns Hopkins University. A. Dallas. Tulane University. Bequaert. LaRocque. Cahn. Curator of Recent and Cenozoic Invertebrates. Both of these gentlemen have also contributed much anatomical material. Puerto Rico Mr. University Museum.C. Litsey. Redlands. S. many of the series of species being incomparable in size and completeness. California Academy of Science. Hoffman. A. Leslie Hubricht. William A. Tok3-o. Japan Dr. formerly of the Pacific Biological Station. Harvard University Medical School. University of British Columbia. John B. B.C. Oregon Dr. Canada Mr. New Orleans Mr. supplemented the National Museum material by the loan of thousands of specimens. William J. Missouri Mr. St. G. Stillman Berry. Thural Dale Foster. The authorities of the United States National Museum at Washington. Ann Arbor Dr. BuRRiNGTON Baker. Berry. Philadelphia John C. Brooks. Andrews. University of Pennsylvania. A. Ernest C. B. Professor of Zoology. C. Vancouver. G. R.* University of Illinois. through the interest of Dr. Baltimore Dr. Curator of the Department of Mollusca. Paul Bartsch. E. University of lUinois. San Francisco Professor Junius Henderson. it would have been imjiossible to understand the relationships of the American species of the family. Nanaimo. Southern IMethodist Universit}'. Texas *Deceased. Adams. Clench. Clemens. Edwardsville. Boulder Dr. Professor of Zoology (Retired). Faust. Dr. Cheatum. Coats. J. Curator of Lower Livertebrates. Carnegie ]\Liseum. P. Now Professor of Zoology. State Museum of New York. Poland Mr. C. California Dr. Cambridge Edward W. Baltimore Dr. D. probably the largest assemblage of species of this family extant. William A. ]\Iuseum of Comparative Zoology of Harvard University. xi . San Juan. Assistant Curator of Paleontology. Warsaw. A. L. Calvin Goodrich. ]Mr. Hanna. Professor of Zoology. Dr. Hertlein. Professor of Paleontology.ACKNOWLEDGMENTS INbeen due to the the successful preparation of this work has help and encouragement of many American conchologists and zoologists. The list of those rendering assistance is given below: LARGE ^MEASURE Dr. Director. Illinois Dr. Urbana Dr. Firkins. San Francisco Mr. have freely lent the collections of Planorbidae for study. Urbana Mr. Pittsburgh Dr. Curator of ]\Ioilusca. H. St.* Zoological Institute. Canadian Geological Survey. Jankowski. Ottawa. Other individuals and institutions have lent or given specimens for both anatomical and dry study. Shintoro Hirase. University of Michigan Museum. University of Colorado. California Academy of Science.* Curator Emeritus. Dallas Dr. Professor.* School of Troi)ical Medicine. C. Without this magnificent material. Stanley T. Tellamoole. formerly Associate Professor of Zoology. E. Curator of Paleontology. To all of these the writer is under the greatest obligation.

Johns Hopkins University. Academy of Natural Sciences. Buffalo Dr. State Museum. Canada Harold A. Washington. D. Washington. Dr. China Baini Prashad. American Museum of Natural History. S. in charge of U. Dr. New Brunswick Dr. The U. Philadelphia Mrs. The libraries lending these books include the John Crerar Library. Florida The writer is highly appreciative of the help given him by the University of Illinois Library. Department of Living Invertebrates. Philadelphia C. Berlin H. William B. Ping. D.C. Ann Arbor Mr. D.M. India D. Professor of Zoology. Washington. Peiping. Dr. Katharine Hill Paul. but also for its aid in securing through the interlibrary loan system many rare volumes not in our own library. Professor of Zoology. Calcutta.Y.N. Harvard University. Van Cleave. artist in the Department of Zoology. Wright. Rao. Calcutta. Louis Miss Elberta C. and Lakeland. Zoological Survey of India. U. Florida Roy W. formerly Assistant Curator of Mollusca (Retired). of Illinois Press here records its appreciation of the assistance of Professor Harley J. University of Illinois. Curator.S. Dr. in addition to writing the 'Memorial' and compiling the bibliography attached thereto. Baltimore. St. Henry A. Rodeck. Van Cleave.S. Iowa City.M. including the legends. T. Without this assistance this volume could not have been completed. Harley J. Curator.. under the direction of Mr. Boulder Mr. and has given many helpful suggestions on the typographical and scientific arrangement of the data. Boulder Mr.S. Department of Science. Cambridge. Assistant Curator. Hugo G.C. Boynton. Curator.* Detroit Dr. Geological Survey. Paul R. Indian Museum. St. Assistant Secretary. Frank Collins Baker *Deceased. Florida Dr.C. Smith. Smithsonian Institution. N. Curator of Mollusca. Division of Mollusca. W. Rehder. Carmichael. tThe University — .f xii The Molluscan Family Planorbidae Mr. Henry Van der Schalie. Fan Memorial Institute. Dr.C. Bureau of Fisheries Mr. Indian Museum. Alexander Wetmore. Bryant Walker. Horace G. not only for the volumes consulted in its stacks. New York City Mr. University of Colorado Museum. Caribou. U. VAN Hyning. Pilsbry. India H. University of Saskatchewan.S. Maine Dr.C. Dr. H. Urbana Dr. University of ^Michigan Museum. Shay. S. Academy of Natural Sciences.N. Normandy High School. has rendered valued service in making numerous alterations of the original drawings to ensure satisfactory engravings. University of Michigan. has carefully checked the copy and the plates. H. Atlantic Coast Biological Station.. Robertson. The photographs of the shells and the copies from author's works were all made in the Photographic Laboratory of the University of Illinois. of the Graduate School of the University of Illinois. Mr. Ann Arbor. To jNIiss Fanny Dunlap the author is deeply indebted for this help.N. Manager of the Laboratory. Andrews.E. Miner. W. Paul McGinty. who has greatly aided in the preparation of the work by providing several grants of funds for the making of the photographic plates. Ray R.M.* Penn Yan. Hamm. Buffalo Museum of Science. Richards. and the Congressional Library of AVashington. Olof 0. Gainsville.. Imogene C. F. D. Nylander. University of Colorado Museum. Mrs. The author desires to express his deep appreciation of the help given by Dean R. Rawson.C. D. Marshall. Dr. Dr. Washington. Smith. who. Needham. Zoological Survey of India.. D. Chicago. Iowa State College. Columbia University of New York. U.S. Associate in Aquatic Biolog}^.

. C.. External Appearance of the Animal B.. xvii A Chronological List of the Scientific Publications xxv xxv xxxv xxxv I Original Articles and OF Frank Collins Baker Books Biographical Contributions Book Notices and Reviews PART I.. Development of Helisoma scolare (Jay) B.CONTENTS Preface ix xi Acknowledgments A Memorial to Frank Collins Baker (1867-1942) . Classification and General ^Morphology General ^Morphology A.. General Internal Anatomy (a) 3 3 4 4 9 10 11 12 13 14 14 15 15 (b) (c) (d) (e) (f) (g) The Reproductive System The Digestive System (1) The Jaw (2) The Radiila The Renal System The Circulatory System The Respiratory System The Nervous System The [Muscular System Shell C. A.. 25 25 28 29 31 31 32 33 34 34 35 59931 . The General Ecology General Environmental Conditions Variations in Different Environments 17 17 18 19 19 19 20 21 Locomotion Food Food for Other Animals Length III. Nidification and Development in Other Species of Helisoma (a) Self-fertilization or Autofecundation (b) The Development (1) (2) (3) (4) (5) (6) of Helisoma Helisoma trivolvis lentum (Say) Helisoma trivolvis fallax (Haldeman) Helisoma duryi nor male Pilsbry Helisoma duryi eudiscus Pilsbry Helisoma tenue californiense F. II... Baker Other Records of Egg Laying Xlll . of Life The Planorbidae Worms XlDIFICATION AND EmBRYOLOGICAL DEVELOPMENT as Hosts for Parasitic ..

Distribution in Time and Space 37 37 38 39 41 . C. Pilsbry.) Charp. 1910 Genus Australorbis Pilsbry.. Geographical Distribution Dispersal of Species V. Genus Afroplanorbis Thiele. 1931 Genus Syrioplanorbis F. A.. 1934 Genus Planorbis Geoff roy. Historical Sketch of Classifications VI. 1837 Genus Odontogyrorbis Lorenthey. 1875 55 57 61 62 64 65 66 72 75 79 80 85 85 85 86 88 89 90 94 96 96 100 104 106 108 for Subfamily Segmentininae F.. 1840 Genus Taphius H. . 1905 Genus Armiger Hartmann.. 1817 Genus Hippeutis (Ag. 1855 Genus Trochorbis Benson. 1837 Subgenus Gyraulus SS Subgenus Torquis.) Charp.xiv The Molluscan Family Planorhidae IV. 1906 Genus Helicorbis Benson. 1939 Subgenus Lateorbis F. Adams.. New Subgenus . . New Subfamily Genus Segmentina Fleming.) Charp. 1855 . A New Classification of the Family Planorbidae General Discussion Anatomical Keys to Groups Systematic List of Genera and Subgenera Recognized This Work 45 45 46 48 in VII. 1767 Genus Anisus Studer. Genera. 1847) Subgenus Anisus SS Subgenus Costorbis Lindholm. Baker.. Systematic Account of the Subfamilies. Baker. and Subgenera Recent and Fossil — 50 50 51 Subfamily Planorbinae H. A. 1855 Genus Tropicorbis Pilsbry and Brown. 1906 Genus Gyraulus (Ag. 1837 Genus Polypylis Pilsbry. C. Baker. 1934 Genus Anisopsis Sandberger... 1820 (Gray. C. new name Heterodiscus West Genus Biomphalaria Preston. Geological History and Distribution B.. and A. 1914 Subgenus Tropicorbis SS Subgenus Obstructio Haas.Dall. 1926 Subgenus Bathyomphalus (Ag..

1840 Subgenus Helisoma SS Subgenus Seminolina Pilsbry. Adams. Baker. 1867 XI. 1934 Genus Paraplanorbis Hanna. Genera Wrongly Referred to Planorbidae Genus Nautilinus :\Iousson. C. 1853 192 194 196 . 1920 196 X. 1928 Genus Helisoma Swainson. 172 172 173 177 178 182 183 187 190 192 192 Groups of Uncertain Affinities Subfamily Choanomphalinae Germain. Systematic Account of the Subfamilies continued Subfamily Segmentininae F. Baker. 1924 Genus Acrorbis Odhner. 1922 Genus Pompholopsis Call. 1806 Subfamily Planorbulinae Pilsbry. 1842 Genus Carinifex ^Y.— Contents xv VII. C. New Subfamily continued Genus Pingiella F. New Genus 109 Genus Intha Annanclale. Bibliography 202 202 202 204 . . 1937 Subfamily Helisomatinae F. 1880 114 — Subgenus Drepanotrema SS Subgenus Fossulorbis Pilsbry. Binney. 1855 Subgenus ^lenetus SS Subgenus ]\Iicromenetus F. 1934 Subgenus Pierosoma Dall. Family Bulinidae Genus Indoplanorbis Annandale and Prashad. 1863 Genus Vorticifex :Meek. . C. 1935 Genus Menetus H. 1905 Genus Promenetus F. 1922 112 Genus Drepanotrema Fischer and Crosse. Baker. 1934 VIII. G. 1859 Genus Poecilospira Morcli. 1905 Subgenus Planorbella Haldeman. 1934 Genus Planorbula Haldeman. 1912 Genus Parapholyx Hanna. 1870 Genus Perrinilla Hannibal. C. Baker. 1922 Genus Platytaphius Pilsbry. 1923 Genus Choanomphalus Gerstfeldt. 1840 Subgenus Planorbula SS Subgenus Haldemanina Dall. Baker. C. and A. 118 118 119 120 121 123 123 124 129 134 150 154 159 160 161 165 166 . IX. 1888 Genus Planorbarius Froriep. 1872 Genus Palaeorbis Beneden and Coemans. New Subgenus Genus Planorbifex Pilsbry.

New Species New Variety New Variety anceps idahoense. . New Species 218 218 219 219 219 220 221 221 222 222 223 224 224 225 226 226 227 227 227 228 229 229 229 230 230 230 231 231 232 232 233 235 521 527 Illustrations of the Anatomy and of Shells Index to Text Index to Illustrations . . Foreword 215 XIII. . . New Variety trivolvis marshalli. New Species pilsbryi preblei. . Genus Planorbida Planorbula smithii. New Variety Menetus cooperi planospirus. New Variety campanulatum rideauense. New Species Menetus portlandensis. New Genus Helisoma Helisoma Helisoma Helisoma Helisoma Helisofna Helisoma Helisoma Helisoma Helisoma Helisoma Helisoma Helisoma Helisoma Helisoma Genus Parapholyx Parapholyx effusa dalli.. New Variety anceps politum. New Variety caribaeum cubense. New Variety columbiense.. . . 218 Tropicorbis shimeki. New Variety Parapholyx klamathensis sinitsini. New Variety Parapholyx pusilla.xvi The Molluscan Family Planorbidae PART II The Planorbidae Inhabiting North and South America AND THE West Indies XII. New Variety Menetus dalli. New Variety subcrenatum perdisjunctum. . New Variety Menetus labiatus. New Species kennicotti. . New Variety nnceps anticostianum. New Species Menetus dilatatus floridensis. New Species Genus Menetus Menetus color adoensis. New Species Menetus cooperi crassilabris. New Variety binneyi randolphi. New Variety campanulatum dalli. . Descriptions of Genus Troyicorhis New Species and Species New Varieties.. New Variety campanulatum pleistocenicum. anceps bartschi.

Baker's concept of the functions of a museum combined not alone the aspects of display and instruction but a third objective as well. 1942. he labored to establish and organize research collections. From that date onward until his death on May 7. Throughout his curatorship. and paleontology. Under his curatorship the Museum of Natural History became utilized by diverse student groups. His appointment as Curator of the L'niversity ]^Iuseum gave him the opportunity to cooperate in the development of a museum de- voted primarily to teaching. not as isolated representations of a dead past. he gave evidence of keen appreciation of the museum as a supplement to laboratory and classroom teaching. geology. For one whose career never placed him before classes for delivering formal instruction. archeology. and at all times offered hearty cooperation to the teaching faculty in creating habitat groups and other exhibits embodying both specific detail and broad principle. they were chiefly summer vacation employment. the dry framework of classification became clothed by artistic and balanced organization of his materials. namely. so likewise in his displays interest was always sought though it was never purchased by cheapening spectacular showmanship. In his synoptic display of the animal series. and geography students found the displays meaningful. and Urbana remained the focal point of his attention and activity. ^Ir. Naturally. 1918. students in agriculture came to study the well-organized habitat groups of pests of farm crops. the fields of his own interests were most emphasized and these included such diverse interests as the ]\Iollusca. Though he carried on extensive field studies during these years. that of research. and art students frequented the exhibition halls for sketching. Groups represented only by fossil remains were treated as integrated with modern expressions of life. xvii . Biology. As in all good teaching one of the first requisites is that of enlisting responsive interest by attractive presentation of fact or material.A MEMORIAL TO FRANK COLLINS BAKER (1867 to 1942) BAKER became FRANK COLLINS in the University Curator of the Museum of of IlUnois on January Natural History 1. To this work he brought a broad background of professional experience in every aspect of museum work and at the same time drew freely from his research experiences to create on the L'rbana campus a series of collections and exhibits the prime purpose of which was ever that of aiding in student instruction. his chief interests centered around his work and his associations on the L'rbana campus.

xviii The Molluscan Family Planorbidae In certain specialized areas of these three fields. had become one of the significant study collections in this field. for students of Mollusca regularly sent doubtful specimens . Baker by his own collecting and by personal exchange. This collection centered around his chief interest. To him who hath shall be given. New apparatus and improved optical equipment made him often express the regret that he could not go back and do again all the earlier studies which he carried on with less adequate laboratory equipment. the fresh-water pulmonate snails. Baker. whose services in cooperation with the State Geological Survey were recognized by his A appointment to the position of Consultant in Pleistocene Paleontology on its staff (1919). stimulated by a cooperative program of studies inaugurated by the State Geological Survey. at the time of Mr. in keeping with that period. Mr. began amassing a private collection of fresh-water Mollusca. Early in his career as a student of Mollusca. the University collections have assumed a significance greater than might be expected in a small museum. Thus. The operation of the old adage. the Pleistocene collections. and numerous collectors throughout the country sent samples for critical determination. Extensive field collecting was particularly stimulated by his association with the Natural History Survey of Wisconsin in the preparation of his exhaustive two-volume work on the Mollusca of Wisconsin (1928). which are now deposited in the State Geological Survey. Enormous series of Pleistocene material in quantitative samples were studied by Mr. His long association and collaboration with the Illinois State Natural History Survey gave him access to extensive collections. The Pleistocene collections under this joint arrangement between Museum and Survey assumed great importance and wide reputation as repository of critical materials. His earliest studies on Mollusca were. The collections of Mollusca are especially rich in representatives of the fresh-water and land Mollusca. and the esteem with which his results are held gives evidence of the effectiveness of his activities in scholarly pursuits. His early interest in the Pleistocene fauna was revived and. Baker's active research program. Baker's death. Baker's interest in research was deep and genuine. Mr.' here finds exemplification. revival of interest in researches on the archeology of the State by the University resulted in amassing within the Natural History Museum huge quantities of pottery and artifacts for the care of which Mr. Baker became responsible. especially in late years when he was engaged in an intensive field study of the land snails of Illinois which culminated in the Fieldbook of Illinois Land Snails published (1939) by the State Natural History Survey. the direct result of Mr. restricted largely to the study of the shells but he early became an advocate of the study of the soft parts of the living animal which produces the shell and the volume which this sketch accompanies is the culmination of his program of anatomical studies.

would not be complete without some narration of his early years and background. was born in Warren. These statements regarding Mr. at the fountainhead of molluscan studies. In fact. He left Brown to accept the Jessup Scholarship (1889-91) in the Philadelphia Academy of Sciences and there. and there the boy received his early education and for a time attended a business college. as indicated elsewhere in this volume. Ottawa Field Naturalist's Club. A. was presented to the United States National Museum. son of Francis Edwin and Anna Collins Baker. He was a fellow of the American Association for the Advancement of Science and held life membership in the Chicago Academy of Sciences and in the Illinois State of Science.Memorial and Bibliography xix to him for classification or verification. even when he faced some of greatest tragedies. which has always been an important aspect of the Academy program. gradually increasing deafness which tended to shut him off from an earlier appreciation of music and from easy communication with his friends gave him opportunity for uninterrupted study in which he could become so completely immersed as to ignore surrounding distrac- Academy With the life's tions entirely and to forget adversities. At the time of his death. Pilsbry. on December 14. In later years he often referred to the inspiration which he obtained from mere proximity to Joseph Leidy and other notables in the library and in the programs of the Academy. and there he learned the techniques of exactness and accuracy which he later maintained were of value in his scientific work. Rhode Island. His family later moved to Providence. Ecological Society of America. Other societies in which he held membership include: American Association of Museums. gripped him. At the time of his death he was president of the American Malacological Union. Frank Collins Baker. the die was cast for his scientific career. the two families Lymnaeidae and Planorbidae were particularly well represented in the Baker Collection. He was a fellow of the Geological Society of America and likewise of the Paleontological Society of America. he cherished particularly the election as Corresponding Alember of the Zoological Society of London. Even before this time he had developed a consuming interest in shells which he often remarked were the favorite childhood toys brought to him from distant oceans by a seafaring grandfather. and samples of those submitted were added to his cabinet. 1867. years. Museum Association of Great Britain. and the tradition for sound molluscan studies. and Sigma Xi. . his zeal for research never waned. A member of many scientific societies. he was for a time employed by the Gorham Silversmiths. as a man and a scientist. There likewise he became associated with H. He attended Brown University for one year (1888-89). which. Still struggling between the alternatives of business and science as a career. American Limnological Society. Baker.

After his return from Mexico. a description of the country. Baker. and T. Chamberlin. and ^Mexico afforded by this expedition had on him the same broadening influence which travel has brought to all the great naturalists and inspired him to consider natural faunal areas instead of political units in subsequent programs of comprehensive taxonomic studies. he became a frequent visitor in the home of John Hall. An intimate correspondence with Henry A. Few strictly educational institutions in the early history of this country have wielded the influence exerted by this private concern in the training of natural scientists.' which wove together a narrative of the trip. At the close of the Exposition. William Stimpson. w^here he succeeded and had association with men of the calibre of Robert Kennicott. especially those headed toward museum work as a profession. the Ward exhibit was purchased by Marshall Field as part of the nucleus for the beginning of the Field Columbian Museum. ended in an arrangement wherein Mr. On June 12. C. Mr. 1892. Baker was engaged as curator in the newly established museum (1894). Baker at the same time selected and organized an extensive synoptic display of museum specimens representing all the groups of the invertebrates. after about two decades with the Academy. and scientific observations. Mr. Upan completion of this task. Mr. Ward of Rochester. Air. Thus his monograph on the Lymnaeidae (1911) was not confined to the boundaries of the United States but encompassed a more natural zoogeographical scope of 'North and ^Middle America. was a member of an exploring party from the Philadelphia Academy into Mexico. a small popular work entitled 'A Naturalist in Mexico. long established Chicago Academy of Science. as zoologist. which was being prepared for exhibition at the Chicago World's Columbian Exposition in 1893. results of which are embodied in this volume. a change in administration placed . he married Lillian Alay Hall who was his constant companion until her death in 1934. Baker published his first separate book (1895).XX The Molluscan Family Planorhidae In 1890. The travels in Cuba. integrated with his professional work. It was here that he perfected the techniques of museum display and administration and for the first time had opportunity to develop an intensive program of research. Baker was to become associated (1891) as Invertebrate Zoologist with the influential Ward's Natural Science Establishment. But that same year he was chosen curator of the old. While his employment by the Ward Establishment was primarily for the identification and organization of the Ward collections of Mollusca. Baker accompanied the exhibit to Chicago where he supervised its installation. Unfortunately. Yucatan. was extended to include even broader scope in order to secure comprehensive treatment of a taxonomic unit without reference to political subdivisions.' and the program on the Planorbidae. and Mr. part of which has been available to the present writer. While residing in Rochester.

The work on this volume marked a new epoch in his research program. Chamberlin of the University of Chicago. While with the Chicago Academy. C. and it was then published by the University Press. Baker's close personal attention to the dredging as it proceeded. he exerted marked influence. Apparently endless material was exposed by the dredging operations. Three of his outstanding volumes were published during the twenty-one years of Mr. Stimulated by an intimate and cordial relation with Professor T. Graduate students in the University of Illinois always found him inspiring and helpful when they approached him with their problems. Some measure of his accomplishment during the Chicago period is available in the larger monographs which punctuated the continuous stream of smaller contributions from his desk. Thus. Baker had embarked upon. — — . thus taking the educational influence of the museum into the schools rather than depending wholly on the unilateral relation of expecting the schools to send their pupils to the museum. These are: two volumes on the Mollusca of the Chicago Area (1898 and 1902) and the monograph on the Lymnaeidae (1911). Baker came to the University of Illinois (1920). The material which served as the original basis for this study was made available in unusual quantity by construction of the Chicago Drainage Canal. However. for nights and holidays were free from dictatorial domination. Baker's direction.Memorial and Bibliography xxi the Academy in the hands of an individual who acrimoniously opposed the place of research as a museum function. Exposure which he experienced at that time led to infections which affected his hearing. In the museums under Mr. and a number of present-day scientists trace the awaking of their interest to his kindly encouragement and enthusiastic attitude toward the field of science. The interest in children. He began the construction of small portable displays of habitat groups for loan to public schools. Conditions there established became progressively worse until in later life his hearing was very seriously impaired. ]\Ir. Baker developed a service feature w^hich has become an important aspect of most public museums today and links them closely with the educational program. Baker's personal interest in research swung toward the study of faunas of the past and interpretation and reconstruction of environmental conditions under which they existed. Even this adverse atmosphere brought no reduction in the research program which Mr. and in education. Another undertaking of particular importance The Life of the Pleistocene was virtually completed during the years with the Chicago Academy. but much of this opportunity for research would have been lost had it not been for ]\Ir. publication of this volume was not afforded until after Air. Baker's association with the Chicago Academy. developed by these contacts always remained with him. Mr. though he never held formal appointment as instructor in any educational institution. the urchin was as welcome a visitor and was as thoughtfully treated as the scholar.

. Baker found deep who of his scientific work. bound. Forbes and Frank Smith. his colleagues helped him celebrate entrance upon status as Curator Emeritus. Baker to the queries of an unknown and apparently unpromising urchin frequenting the museum of the Chicago Academy. employed him as an investigator. For two years he carried on a comprehensive evaluation of the relations of bottom fauna to fish life in Oneida Lake. In 1936 Air. Baker reached the age established by the University for retirement from active service but his good health and vigor and the best interests of the University combined to bring about postponement of his retirement for three additional years. the Museum of Natural History had been in charge of members of the faculty whose foremost obligation had been that of teaching or administration. In 1939. Baker w^ho. in his seventy-second year. This program culminated in two volumes of distinctive contributions which were published by the New York State College of Forestry. and presented to him at a dinner in his honor. She explained that the son. In that same year. which permitted him to devote his entire time to completion of researches in progress.xxii The Molluscan Family Planorbidae satisfaction in following the careers of men attributed part of their early zeal for science to boyhood contacts with him and to inspiration growing out of his enthusiasm and kindly interest in their personal problems. One instance of an influence of which he was personally unaware stands out prominently. The newly established New York State College of Forestry. The esteem with which he was held as an individual and acknowdedgments of personal debts of gratitude were as sincely expressed as was the evaluation Mr. A mother called at his office in Urbana to present him with a copy of a scientific book written by her son. the Museum had prospered and was able to secure the services of very unusual collectors and competent preparators w^ho added materially to its development. 1918. The position was offered to Mr. Under wise guidance of men such as S. on the campus of Syracus University. Ever since the founding of the University of Illinois. at the height of his professional career. Letters of congratulation and appreciation from colleagues and from fellow scientists throughout the country were assembled. definitely attributed his aspiration in the field of science to the benevolent and gracious attention of Air. Mr. entered upon his duties in January. A. Toward the end of President James's administration. who had died while on archeological explorations in a foreign land. Baker resigned from his position with the Chicago Academy in 1915. he embarked on a new phase of his career in which his time was given over entirely to field study and research. a renewed interest in archeology and in natural resources together with rapid increase in University enrollment led to consideration of the problem of securing a full-time curator for the Museum of Natural History.

Baker has left an undying impression of honesty. Relatively early he began to shift his interest in Mollusca from purely conchological to malacological lines (about 1897) when he began to study the radulae and anatomy of the soft parts of the molluscs. Following the operation he never regained his health but became rapidly weaker and died on 7. Tryon. 1929. and familiarity with the history of the subject led to his being commissioned to prepare biographical sketches for five of the distinguished students of Mollusca in the Dictionary of American Biography (Gould. Faunal studies with considerable attention to ecological relations of the molluscs began early and found particular expression in his monograph on the Lymnaeidae (1911) and in his ecological studies on the fauna of Oneida Lake. In this aspect . integrity. illness impaired his opportunity for continuous work in the laboratory. but even under this physical handicap he continued to devote all the time that his strength would allow to the final studies for and preparation of the manuscript for his the monograph on The Molluscan Family Plannuu/nuifi opus orbidae. For more than half a century (1889-1942) Mr. mild in disposition. His long identification with the program of molluscan studies. Baker continued to publish the results of his researches and observations.Memorial and Bibliography xxiii Soon thereafter. Stimpson. These studies stand as background to the interest which found final expression in his contributions as associate editor of Xaturalist's Guide to the Americas (1926). The interest in Pleistocene paleontology first found expression in a paper published in 1898 and continued through the rest of his life with two posthumous papers on this subject. Kennicott. was ready for the editor and work was progressing on Volume II when he was taken critically ill and hospitalization for surgery was unavoid- — able. 1930. His publications in archeology were stimulated by his responsibility for the University collections and were concerned largely with analysis of shells and other remains of animal life encountered in excavations of Indian village sites in the Middle West. his early publications dealt largely with May distributional studies and problems of taxonomy of the MoUusca. and Sterki. Slight in stature. 1933. and unbiased in all of his personal and professional relations. 1942. charitable. Mr. Aside from purely incidental notes and compilations. which he continued to publish throughout his career. 1933). Volume 1. and genuineness in the memories of his many friends and associates. Interest in museum administration found expression in a series of papers issued during his years with the Chicago Academy and was revived when he became established in the museum at the University of Illinois. and kindly. kindly personal qualifications. 1932. unpretentious in attitude. based on exhaustive anatomical researches.

the Mollusca. Moorehead whose reports on studies conducted for the University of Illinois were edited by Mr. Baker wrote numerous reviews of current works on Mollusca for Ecology. is here published in full with but few slight editorial changes and corrections. Throughout his active career Mr.xxiv The Molluscan Family Planorbidae of his work. which he prepared. Mr. a copy of which was in his desk at the time of his death. From 1913 to 1915 he was assistant editor of Home and School Reference Work. In his total contributions. published in Chicago in seven volumes. in addition to serving as associate editor of Naturalist's Guide to the Americas. From 1921 to 1926. Harley Jones Van Cleave . K. more than 250 deal entirely or largely with his chosen field of research. minor notes. Baker edited the Bulletins and other publications of the Chicago Academy of Sciences. This list. exceeding 350 original articles. Baker kept a carefully compiled record of his publications. some quantitative evaluation of Air. In 1926. To supplement this analysis it should be recorded that in the period from 1900 to 1915. Baker. Earlier (1900) he prepared a one-page article on Mollusca for the United Editors Encyclopedia published in New York. Baker's productivity may be reflected in the fact that more than a dozen of these works enumerated are of such size as to merit classification as separate books. to which he personally contributed 125 articles on birds. he collaborated with various specialists but had closest relations with the late W. While many of these are but short. he prepared the references therein to molluscan distribution. Mr.

Bull. Proc. Chicago. Also m . 1890 7. Preliminary outline of a new classificatioii of the Muricidae. 17. 3:80-81. 2:19-40. 19(1) :45-48.. Remarks upon the round-tailed muskrat. Op. Nat. 1894:223-224. Oologists Exchange. 1:129-137. collection of shells from southern Mexico. Remarks on the Muricidae with descriptions of new species of shells. Sci. New York. with a consideration of the Genus Magilus of Montfort. Acad. 3(11). Notes on a collection of shells from the Mauritius. 9(3). 13. cit. 1891:45-55.. 18. Ward's Natural Science Establishment. 16 pages. 4. XXV . Descriptions of new species of Muricidae. Notes on Florida shells. 115-117. 1889:266-270. Philadelphia. Philadelphia. 2(9). Conchology department in 'The Old Curiosity Shop' writer was editor). 1891:56-61. Sci. A catalogue of conchological abbreviations. Sci. Proc. Remarks on Urosnlpinx perrugatus Conrad. 15. cit. Acad. Catalogue and synonymy of the recent species of the family Muricidae. Chicago. Proc.. For the year 1895. Philadelphia. Cahfornia. 1:153-172. On the modification of the apex of Murex. Rochester. Proc. Sci. Nat. Rochester Acad. 21. Proc. Cincinnati Soc. Nautilus. Acad.. Science (n. Oologists Exchange. gorgonians. 3:53-54. (of which department the 3. with remarks on the apices of certain forms. Florida. Sci. 4:89-91. 16. 2:179-183. Notes on the food of birds. Proc. 1896 : + 26. 1889: 271-273. 1896:680-682. 6. Nat. On 8. Acad. the Indian River. 25. Further notes on embryonic whorls of the Muricidae.s. 1895 viii 145 pages. Nat. A 24.A CHRONOLOGICAL LIST OF THE SCIENTIFIC PUBLICATIONS OF FRANK COLLINS BAKER ORIGINAL ARTICLES AND BOOKS 1889 1. Philadelphia. Rochester Acad.Y. Description of a new species of Ocinebra. Jour. Old Curiosity Shop. Catalogue of specimens of Echinodermata. On the correct position of the aperture in Plajiorbis.82 pages. 23. Acad. Philadelphia. Op. Hist. and Mexico. A naturalist in Mexico: being a visit to Cuba. Op. Philadelphia. 1890:66-72. Syllabvis of biological lectures. Chicago Acad. 1890:46-47. A few notes oi. northern Yucatan. Nautilus. 2. 27. 4:29-30. Nat. 10. 11. Sci. 9. 2:169-189. Nautilus. Neofibcr allcni True. Nautilus. 5. The Observer. Notes on the mental development and intelligence in animals. 2:2.. and sponges. corals. (Delivered before the North Side Science Section of the Chicago Institute of Education). 1891 12. iv -f. Sci. Nat. The geographic distribution of ttie Mollusca. Nat.i ornithological collecting. 14. Notes on a 1894 19. Directions for collecting alcoholic specimens of the Mollusca.. Thirty-eighth annual report of the Chicago Academy of Sciences. N.).. 1895 20. Acad. glance at the Chicago Academy of Sciences. Sci. Op.. The caves of Yucatan. Sci. 22. 8:109-111.. ^ Proc. cit. Proc. cit.

Op. 26 page-s. Acad. Description of a new species of Limnaea. 2(3) 191-211. 7:371-391. Op. Birds and Nature. Sci. and G. 34. 54. Chicago. 57. Trans. riyer. The digitations of the mantle in Physa. Nautilus. St. Snails of pond. ll(3):28-30. 68. 35:659-664.. Notes on the mollusks of Lilycash Creek. Snails of the ocean. Cincinnati Soc. Sports Afield. 7:155-157.. cit. Notes on the Mollusca of Owasco Lake.. St. Chicago Acad. Birds and Nature.. Further notes on animal intelligence. 43. Limnaea auricularia in America. Trans. On the modification of the apex in gastropod mollusks. Notes on mollusks from Acadia. collection of Pleistocene shells from Milwaukee. yariety Mighehi Binney. Nat. 38. Hist. Sci. cit. 56. Op. 72. Note on Natrix grahami B. St. Hist. 5:447. The gross anatomy of Limnnca cmarginata Say. 14:69-71. Nat.s. 29. 42. Cincinnati See. 32. (n. Study of fresh-water mussels. Nautilus. Jour. 20:215. Op. Louis. 33(5) :99-100. 44. Sci. 9:222-227.. Nautilus. 12(6) :63-65. The Mollusca 45. 34:187-188. 27 pages. 73. 1899 47.. Louis. Snails of the forest and field. Op. 70.. 9:176-181. Sci. 15(2):17-18. 1900 New York.. Collecting about Chicago. Cincinnati Soc. Nat. 14:93. Illinois. For the year 1897. 30.. 31. a collection of mollusks from 1898 39. Birds and Nature. cit. Nat. New yarieties of fresh-water shells. day on the Chicago drainage canal. A new museum tablet. The educational yalue of natural history museums. Shell collecting near Rochester. 3(1):1-130. 13(3) :30-31. 15(5) :59. Some interesting molluscan monstrosities. 8:179-181. Trans. 13:112-113. Fortieth annual report of the Chicago Academy of Sciences. Nat. 35. 13:57-59. Collecting fresh-water shells. Nautilus. cit. Sports Afield. 61. 20:314-316.. Nat. 19(2) 28. Nautilus.Part L The Pelecypoda. 49. Chicago Acad. On the pulsations of the molluscan heart. 2(4) :225-228. 19(3) :81-93. Nautilus. Wisconsin. Acad. Acad. 8(5) 41. Nautilus. 55. On a collection of fresh-water . For the year 1896. 64.). Suryey Bull. 7:191-192. 46.. 11:36. Birds and Nature. 34:283-284.shells from Rhode Island. Critical notes on the Muricidae. New York. Bull. cit. 9:685-704. Chicago Acad. 67. Sci. The prong-horned antelope. The Nautilus and other cephalopods. 19:276-278.. Science On the effect of music on caged animals. Popular Sci. Bull. St. The teeth of snails. 69. Jour. Sci. Op. 40. Notes on Radulae.. Op. The molluscan fauna 71-94. Nat. 73-78. Do cats remember? Sports Afield. oyster and its relatiyes. 9(1): 1-24. 63. 33. A A of the Chicago area. 19(2) :112-113. Amer.. The . (With F. cit. 51. Sci. Nautilus. A reyision of the Limnaeas of northern Illinois. cit. 9:74-79. Amer. 9:26-31. a : Notes on A 1901 60. 71. 36. 8:193-194.: xxvi The Molluscmi Family Planorhidae 1897 Thirty-ninth annual report of the Chicago Academy of Sciences. Trans. 65. Nautilus. 2(4) :229-230. Woodruff).. 53. 59. Louis. Reyiew of Education. 9:128-133.. Collecting in Florida. M. 11:143-146. of western New York. 62. 50. new Sphaerium. Mollusks in grass. Sci. Hist. 66. Nautilus. Louis. 48. The Obseryer. On Grand Tower. 14(2) :16-24. York New Acad. 12:65-66. The rock shells. 58. Jour. Acad.. Missouri. 52. Popular Sci. Hist. 1897:460-463.. Amer.. Popular Sci. Reason among animals. The molluscan fauna of the Genessee Riyer. 37.. 19:175-177. reyision of the Physae of northeastern Illinois. 33:194-197.. and brook. Ann..

98. Coral reefs.. 1903 81. Op. 17:50-51. 107. 14:179-181. 17:201-202. 18:125-127. 78. 18:141-142. The mollusks of Cedar Lake. 7:361. (Nora de plume.. 13:225-227.. cit.. 90. 18:50-51. 37(439) :481-488. cit. Corals. The stonv corals. Gastropoda. (Nom de plume. Acad. Collins Thurber) : 1905 102. The loggerhead shrike. (Nom de plume. Collins Thurber) The cerulian warbler.: Chronological List of Publications 1902 74. Collins Thurber) Notes on the genitalia of Lymnaea. Birds and Nature. Some snails of the ocean. The arrangement of the collections of Mollusca in the Chicago Academy of Sciences. (Nom de plume. The descriptive arrangement of museum collections. Indiana. Op. Collms Thurber) The on the smaller Lynmaeas. xvi 175 pages. Op. 14(2):99-105. Nat. Collins Thurber) The American flamingo. Op. 13:179-181... 88. familiar introduction to the study of the mollusks. Note on PUutorbis tnmcatus Miles. cit. The marsh hawk. 13:83-85. Birds and Nature. cit. 16:158-161. Op. Sci. Op. Birds and Nature. sea-fans. St. 85. 117. (Nom de plume. 100.. Trans. Op. 94. Collins Thurber) New varieties of American Limnaeas. Rib variation in Cardium. 3(2) :131-410. Hist. 18:23-24. Amer. cit. Nat. Nat. 18:10-12. New species of Lymrmea. A. New American Lynmaeas.. 18:119-121. corals. Collins 108. cit. (Nom de plume. and sea-whips. Birds and Nature. 5:50-55. Sea urchins and sand dollars. Nautilus. Nautilus. land and water.. 15:131-133. with notes on Lymnaea parva Lea. on museum articles. 18:62-63. (Nom de plume. cit. Spire variation in Pyramidula alternnta. Trans. Op. The bluebird. The Mollusca of the Chicago area. cit. New Jersey. 103. 75. Birds and Nature.. 99. Critical notes 105. Collins Thurber) of McGregor. 17:57. Mumford and Co. Shell collecting on the Mississippi. Nautilus. 38(453) :661-668. (Nom de plume. 95. (Nom de plume. 101. The cowries and shell money.. Collins Thurber) The flicker. Op. 19:51-53.. Some American grasshoppers.. 112. Nautilus. Collins Thurber) 106. n. 79. 18:167-168. Chicago. (Nom de plume... Nautilus. Chicago Acad. long-billed marsh wren.. Birds and Nature. 96. 89. 15(10) :120. 14(2) :107-110. II. 39:665-679. Starfishes. var. Collins Thurber) . The museum and the public school. Nautilus.. 83. Chicago. Bahamas. Sci.. Iowa. Amer. The molluscan fauna Thurber) 113. A + 1904 92. Pleistocene mollusks of White Pond. Museum Jour. 19:27-28. (Nom de plume. Review of Education. 110. Editorial 80.. 77. cii. 109. W. 17:38-39. Louis. 17:112-113. Nautilus. 12:35-37. Collins Thurber) The bald eagle. cit. 104. The hydroid 86. Op. The hermit 114. The belted kingfisher. 12:134-139. 76. 93. Nautilus. Op. (Nom de plume. (England). 91. Birds and Nature. Sci. The molluscan fauna of the Dells of Wisconsin. Amer.. Notes on Murex marcoensis Sowerby. Birds and Nature. 2:106-110. 16:102-105. Acad. 17:131-132. Op. A new species of Lymnaea from Ohio. 17:179-180. Birds and Nature. 16:232-236. thrush. A partial list of the marine mollusks of San Salvador. 13:131-133. 116. cit. The yellow-bellied sapsucker. Part II. 2 354-360. xxvii Fhnwrbis bicariitatiis striaius. Louis. Shells of 87. cit. 12:86-90. Nautilus. Nautilus. Museums Journal (England). Op. Museums Journal (England). Notes on a collection of shells from Bass Lake. 84. 115. 111. 97. 18:213-214. 82. Nat. cit. Indiana. cit. Survey Bull. Op. 17-88.. 15(3) 249-258. St.

23:19-21. 149. (Nom de plume. cit. The ecology of the Skokie Marsh area. 1910 155. Some observations on museum administration. 132. Sci.. 143. Op. Science. 20:125-127. 2:15-19. Nautilus. 1907 130. 138. Annual report of the Chicago Academy of Sciences.. 22:45. . Op. Nat. Michigan. The Chicago Academy of Sciences: its past history and present collections. cit. 19:146-147. Louis. 8:423-428. Science. cit. 150. 27:423-424. A catalogue of the Mollusca of Illinois. The relation of the State Academy of Science to the Natural A A of Sciences. Nautilus. Annual report of the librarian. 3:26-34. Op. Bull. Hist. sp. 136. The Natural History Survey of the Assoc. 157. Mus. 154. 2:1-7. Nautilus. Notes on a collection of mollusks from the \-icinitv of Alpena.. Photo Era. Nautilus.s Linn. 119.. 19:110-111.. Provincial museums.. 53-136. Amer. Collins Thurber) The blue-gray gnatcatcher. Bull. For the j'^ear Chicago Acad. 3:1-21. 19:213-214. 139. Application of De Vries's mutation theory to the molluscs. 22:20-23. 1909 145. The bob-white or quail. Proc. Amer. Nautilus. 23:41-42. 16(2):1-15. new species of Lymnaea. 3:26-45. 1908 134. Science. 26:666-669. sp. 7(6): 121. Bull. Hist. 19:23-24. A new Lijmnaea hinklcyi 124. Some instructive methods of bird installation. Acad. 8:437-499. 129. Op. 147. Nautilus. The Chicago Academy of Sciences. Suggestions for an educational exhibit of mollusks.. 23:91-94. Sci. 1:52-57. 20(2) :21. Amer. Mus. 3:128-130. 19:142-143. Op. new American Planorbis. 160. Op. cit. 125. The Chicago Academy of Sciences: Its past history and present 1908. 21:52-55. Collins Thurber) The least bittern. Proc. (England). 128. Bull. Op. 152. Suggestions for a natural classification of the familv 942-943. 1:68-69. 18:185-186. Chicago Acad. Science. Sci.. 19:120. Chicago Academy of Sciences. cit. 140. 159. 137. 31:221. A new species of Lymnaea. Illinois State Lab. The camera in science. Lvmnaeidae. Bull. Collins Thurber) The great blue heron. 19:71-72.. 158. 120. Nautilus. Two 133. Rei)ort Museums Jour. 146. collections. the Chicago Academy History Sur\'ey of Trans.xxviii TJie Molluscan Family Plauorbidae 1906 118. (Nom de plume. Op. 142. Sci. special publ. Assoc.. A new Sphaerium from Illinois. 126. 131. 28:534-535. Birds and Nature. Note on free public museums. Science. 122. Exhibition cases without shelves. Sci. 20:55. cit. 148. 40: 327-334. Assoc. Mus. 23:112-113.. 22:140-141. Proc.. Nat.. The dickcissel. A new variety of Lymnaea stagnalis. Note on Planorbis binneyi Tryon.. Nat. 127. with special reference to the Mollusca. cit. Op. 144. 123. cit. St. Chicago Acad. Nautilus. Illinois State Lab. Mollusks from Kansas and Oklahoma. (Nom de plume. small collection of shells from Texas. 135. Description of a new fossil Lymnaea. n. Science.. Nautilus. Annual report of the curator. 151. 28:684-685. 156. Collins Thurber) variety of Planorbis nautileu. on the educational work of 23:80 east American museums. 28:138-140. cit. 141. 125-126... Collins Thurber) Lyninaea daniehi n. 27: Chicago Acad. 153. Op. Amer. (Nom de plume. some 3:56-59. Range of Lymnaea umbiJicata. Descriptions of new species of Lyninaea. Op. Nautilus.. Trans. Note on Lymnaea desidiosa Say.. Preliminary note on the life of Glacial Lake Chicago. The Chicago Academy of Sciences... Sci.. Nautilus. (Xom de plume. cit. Illinois State Acad. 31:715-717. cit. 3:45-46.

Amer. New York. Proc. Lawrence Riyer. Further notes on the po. 3:xyi 539. St. Notes on post-glacial Mollusca. 1916 191. 163. 29:36.. The A . 170. Interglacial records in York. Assoc.. 189.dli. Notes on post-glacial Mollu.... A method of exhibiting large descriptiye labels on the outside of museum cases.. Cass and Van Buren Counties.. 4:21-23. County. 196. Michigan. Wisconsin. 6:108-111. Amer. Science. On the classification of the Lymnaeidae. 27:54. cit. Nautilus. Proc. Nature Study Rey.. 3:151-152. Op. 24:58-60. A Trans. 182. The molluscan fauna Arts. Nautilus. 186. Op. Op. 199... New Emmet Nautilus. Sci. 1913 176. cit. 36:523-524. areas. Op. 195. 26:115-116. 180.. 27:104-106. 174. 178. + 171. 27:68. Campeloma leunsii Walker in Illinois. Bull. relation of mollusks to fish in (Dneida Lake. Op. Sci. 5:143-145.. 3:137-150. Pseudogalba. recent and fossil. Amer. cit. 193. Nautilus. The fresh-water Mollusca of Oneida Lake. and of Tomahawk Lett. 9:394-397. 173. cit. Mollusks from Magician Lake. Nautilus. 3:106-108.. Sci. Science. A new Lymnaea from Montana. new name for Si))ipsonia. Illinois State Acad. 44:273-275. 25:66-67. Op. L"se of enlarged 89-91. Illinois State 5:108-116. 30:5-9. 172.. Nautilus. Trans. Chicago Acad. Chicago Special Publ. mollusk injurious to garden yegetables. new Flauorbis from Michigan. II. Mollusks of Unionville. Soc. Science.. of the Skokie Marsh area with sjiecial reference to the moUusks. Amer. Annual report of the librarian.Chronological List of Publications 161. Trans. cit. 194. Connecticut. Mus. 168. (Abstract).s from Oneida Lake with notes on the luteola group.. 26:120... 1914 c. 164. Northern Idaho shells. Trans.. Science. of North and Middle America. Wisconsin. Science./. 25:118-120. Lake. 185. 179. New York. New York State Coll. 29:87-88. 38:858-859. 162. 24:68-69. Op. 1916. Op. 44:568. Illinois State Acad. Assoc. Michigan. Recent additions to the catalog of Illinois Mollusca. Glacial Lake Chicago. 177. The auroral display of August 26. (England). Mus 187. 29:20-24. Description of a new Lymnaea. Op. I. cit.. Proc. Illinois State Acad. cit. Wisconsin Acad. 167. 166. cit. 43:136. 28:8-10.. Galba jerruginea in Oregon.. 27:24. 30:74-77. mollusks from Illinois. An eyening with the aquarium and snailery. Suggestions for XXIX an cducutiomil exhibit of niollusks. Sci. 10:315-320 1915 188.. The ecology 1911 165. The Lymnaeidae Acad. 198. Technical Publ. Mollusks of Wellesley Island and yicinity. Muscvniis Jour. Acad. photographs in museum groups. For. Michigan.sks of Oneida Lake to the basin of lower South Bay. Post-glacial life of Wilmette Bay. The relation of the Mollu. Pleistocene 192. 183. Nautilus. 7:74-78. 169. School loan collections as prepared by the Chicago Academy of Sciences. Mus. 27:7-8. Nautilus. Op. Trans. Assoc. Ecol. Sci. new yariety of Lamp. 175. 4:36-40. 190. Sci. A North American faunal 197. 17(1) :200-246. A method of exhibiting insect collections. Mollusca of Wyoming County. Mollusks from Berrien County. Interglacial mollusks from South Dakota. 1912 Sci. Waukesha County. 29:47.. Op. cit. 184. Annual report of the curator. cit... biota of Glacial Lake Chicago. Nautilus.

Op. The Museum of Natural Historv of the University of Illinois.. 205. Baker's 'Life of the Pleistocene' (reply to criticism by Dr. Studies in aquiculture or fresh-water farming.. Soc.. New York. Sci.^ from Illinois. 33:123-125. 19:478-488. Science. New York. Amer. Description of a 210. Urbana.476 smithii. New York. 10:376-380. and Math. Amer.. . 219. For.. 54:152-161. 216.' a statistical study of the administration of President Edmund J. 207. Circular. Post-glacial Mollusca from the marls of Central Illinois. : Illinois. 222.. 220. 33:11-13. James. 20:135-138. Trans. the molhisks of lower South Bay. 227. Illinois Bull.: XXX The Molluscan Family Planorbidae 1917 200. Mollusks infested with parasitic worms. The Museum of Natural History. 12:57-63. 228. 49:519-521. : new 1920 217. A new Planorbi. Physa smithiana. 212.. pages 119-121. (With Frank Smith). 214. Hist.. Nautilus. Op. 211.. 41(13) :527-539. 226. 34:61-66. New forms of Pleistocene mollusks from Illinois. Science. 225. Trans. Mus. Nautilus. 206. 223. Preparing collections of the Mollusca for exhibition and study. Nautilus. Animal life in loess deposits near Alton. 33:142. The importance of ecologv in the interpretation of fossil faunas. Illinois. Amer. Shufeldt). cit. Op. Nautilus... Micros. cit. Nautilus. Molluska of the Crocker-Land Expedition to northwest Greenland and Grinnell Land. New York State 1919 209. For. Jour. Trans. 47:391-392. Further notes 202. Illinois State Acad. 215. 232. 21:323-327. 9:247-248. 13:271-279. cit. Description of a new variety of fresh-water mussel from Oneida Lake. with special reference to mollusks. 208.. A new form of Amnicola from the Ohio Pleistocene deposits. Nautilus. 32(3) 94-97. 40(l):31-46. 26(7) The 659-671. Appendix to New York State Coll. 31:81-93. Geol. 230. species and variety of Planorbis from post-glacial deposits. 21:1-34. The ecology of North American Lymnaeidae. on the Mollusca of Oneida Lake. Op. mussel survey of the ujijier waters of the \'ermilion River with special reference to the Salt Fork. 28(5) :439-457. 32(3) :97-98. Op. York State. In 'Sixteen Years at the University of Illinois. University of Illinois. New York. The animal world in a handful of algae. and sewage in the Genessee River. cit.. Notes on a small collection of shells from Alaska. 231. 38:155-156. Op. A moHuscan garden pest. with notes on a Physa from the same formation... The productivity of invertebrate fish food on the bottom of the Oneida Lake. Coll. 1918 201. Animal life The life of the down by the Urbana. new name for Phy. cit. Effect of sewage and other pollution on animal 1921 life of rivers and streams. 17(41) iv -f. 218. 224. cit.. Op. Op. cit. Nat. Illinois State Acad. School Sci. Univ. Technical Publ. Hist. Nat. Pleistocene or glacial period. 221.. The Guide to Nature. A 12:129-131. 2: 277-280. Bull. relation of shellfish to fish in Oneida Lake. cit. 229. 33:125-127. Jour.) Binney. Ecology. The value of ecology in the interpretation of fossil faunas. 204. 213.. Pleistocene Mollusca from Indiana and Ohio. A new species of Physa from New. with descriptions of two new varieties of land shells from the same deposits. Geol. 34:67-68. Sci. Notes on Acella haldemani (Desh. Nat. 41(11) :479-517. The Auk. Fresh-water Mollusca from Colorado and Alberta. 32:19-23. 9:1-264. Notes on nidification in Gillia and Amnicola. Nautilus.. as recorded in the deposits laid great ice sheets.. 203. 35:22-24.

In 'President's Report. 38-41. A new Physa from California. Description of a new Lymiiaea from Yellowstone Park. cit. The naiad fauna of the Rock River Svstem.. 36:22-25. Trans.. In 'President's Report. The molluscan faima of the Big Vermilion River. 64:249. 248.. Nautilus. cit. A quantitative and qualitative survey with special reference to the Mollusca. 7:81-87. 41: 250. Pilsbry). A. Illinois State Acad. 30(1): 43-62. 38:15. Wisconsin.. 252.' 1924 place of the museum in university instruction. Parasitol. Sci. 55-57. 238. New forms of Planorbis and Lymnaca with notes on other forms. 7(2):1-127. A new Anodontoidcs from Wisconsin. 40(3) 85-87. 40:49-52. 15:408-420. Monogr. Pleistocene Molluska from northwestern and central Illinois. 245. 1924-1925:117-119. (Reprinted in School Life. 236. Trans. 19:103-112. In 'President's Report.: Chronological List of Publications 1922 xxxi 233. the Cahokia Museum Work. 22:651-655. Sci. cit. curator. Trans. New species and varieties of Mollusca from Lake Winnebago.' Uni- versity of Illinois. University of Illinois. Science. Op. 256. with notes on shells from the latter state. : New .. 258. New varieties of Stagnicola from Wisconsin and Wyoming. 259. 254. 1922-1923:159-162. Some problems of a university The use of molluscan shells bv Acad. 1923 Pleistocene Mollusca from the vicinity of Joliet. mound builders. varieties of Helisoma antrosa from Wisconsin and Minnesota. Ilhnois State Acad. Fluke infections and the destruction of the invertebrate host. 251.. a studv of the law of stream distribution. 240. versity of Illinois. Op. Nautilus. Nautilus. Nautilus. 27-29. Wisconsin Acad. and Lett. 242. 1926 nature. Op. 35:130-133.. The 243. 244. Arts.. 253. cit. and Math. pollution on animal life. 41 23-24. 58: 239. with new records for this state. Bowmanville low water stage of Glacial Lake Chicago. Museums and The effects of 23-24. The museum. Op. Science.. Illinois Biol.. 255. 40: 82-85. Trans. 260. Description of a new species of Somatogyrus from Wisconsin.. 10:33-35). Report of curator. Sci. 237. 246. The 1925 249. Museum of Natural Historv.' Uni- 1927 257. 234. Nautilus. 261. 21:109-146. 235.. Trans. 36:19-21. Museum Work. Naturalist's Guide to the Americas. Jour. Sci. Fresh-water Mollusca from White Lake. Nomenclatorial notes on American fresh-water Mollusca. 41:24-26. Nautilus. and Lett.. 8:145-147.. GeoL. Wisconsin Acad.. 241. 1925-1926:145-149. Report of curator. Michigan. 247. 5:92-94. The fauna of the Lake Winnebago region. Sci. Description of a new species of Lymnaea from British Columbia. cit. cit. Arts.. the Museum of Natural History. The educational value of a university natural history museum. Jour. Illinois State 16:328-334. the original exponent of visual education. (With H. School Sci. Oakland County. with special reference to its modification as the result of pollution by sewage and manufacturing wastes. Op. Museum of Natural Historv. Report of curator. Op. 36:123-125. Illinois. 22:192-205. Illinois. New Lymnaeas from Wisconsin and Minnesota. 36:125-126.

shells. Helisoma corpulenta Say. 1929-1930:164-169. Micros. A study of the Pleistocene Mollusca collected in 1927 from deposits in Fulton County. 41:49-51. Descriptions of new varieties of land and fresh-water mollusks from Pleistocene deposits in Illinois. Nautilus. 285a. Trans. 283. Soc. 274.. In Henderson.' University of Illinois. 285. Nautilus. Certain anatomical features of the fresh-water mollusc. Nautilus.. 42:48-60. and the description of new species and genus of Unionidae. In 'President's Report. 11(4) 788-789. Molluscan associations of White Lake. Trans. The molluscan fauna 44(l):21-24. . Baker and Junius Henderson. 290. Fossaria perplexa F. 1927-1928:124-127. Notes on Professor Shimek's paper on land : . Report of curator. The + versity of Illinois. 264. cit. cit. 273. 268. On genus and species making. Op. Pelecypoda. 8(3) 353-370. Museum of Natural History. Amer. The Molluscan Family Planorbidae a 263. Museum of Natural History. 276. A new variety of Helisoma campanulata from Michigan. 277.. Arts. In 'President's Report. 22:186-194. 'Some Fresh-water Mollusca from Washington and Oregon. Illinois State Acad. : 1930 new record of Castoroicles ohioensis from Illinois. Univ. Trans. 72:37-39. New York. C. the division of the Sphaeriidae into two subfamilies. Illinois. 48(1) :44-47. 10(7) :220-223.' Uni271. 1527: vi 495 pages. In 'President's Report. Mollusca from Vermilion and Pelican Lakes. : On 1928 267. Ecology. Illinois. with the description of a new variety of Helisoma corpulenta. The use of molluscan shells by the Cahokia mound builders. Museum of Natural Historv. Report of curator. 41:132-137. Op. Minnesota. Univ. Nat. 131-136. 1928-1929:140-146. 21 :288-312. Part II.. Wisconsin. 7(19):11-12. of the southern part of Lake Michigan and its relationship to old Glacial Lake Chicago.' University of Illinois. 20:269-292. Science. The American Bythinia not whollv an introduced species. 279. 287. 292. Stagnicola couleensis F. 269.. 41:118-120. 272. Museum of Natural History. Amer. 278. 270. cit. of Sci. 288. Bull. 26(4) 147-154. 265. Op. 280. Molluscan life of the loess deposits of Illinois. A review of our present knowledge concerning the character and distribution of the Pleistocene aquatic molluscan life of Illinois. 291. 42:95-97. 284. Op. 1929 275. 42:103-104. University museum exhibition cases. Illinois State Acad. Ecology. Descriptions of new forms of Pleistocene land mollusks from Illinois. Sci. Ecology.. 266. Report of curator. Gastropoda. 11(3) :469-480. The Cahokia Mounds. Trans. C. Wisconsin Acad. Part I. In 'President's Report. Illinois State Acad. 41:108.' University of Illinois. Midi. The fresh-water Mollusca of Wisconsin. Op. A 22:41-64. Bull..xxxii 262. with remarks on other species. 282. with descriptions of new varieties... fresh-water Mollusca of Wisconsin. snails as indicators of Ecological conditions. 20:56-63. Ecology. 40:114-120. Sci. cit. Report of curator. Nautilus. The Mollusca of Chautauqua Lake. and Lett. The use of animal life bv the mound-building Indians of Illinois. The variation of molluscan life during the Pleistocene and recent time. Nautilus. The European starling in Illinois.. Michigan a study of a small inland lake from an ecological and systematic viewpoint. Vermont 281. 1928 :i-xx -f 507 pages. 286. Sci. Baker. cit. 1926-1927:129-132. 9:271-283.' Nautilus.. 289. 71:389. Museum News. 22:411-434. Influence of the glacial period in changing the character of the molluscan fauna of North America. with descriptions of a new variety of Ptychohranchus and of Helkoma. Influence of a changed en\-ironment in the formation of new species and varieties. Nautilus. Nautilus. 69:521-522. Science.

Op. Jour. Moorehead's 'Exploration of the Etowah Site in Georgia. 319. 326. Illinois. 47(2) :59. Self fertilization and nidification in Physa halei. Amer. Op. of Lvmnaeidae from British America. Minnesota. Ecological relationship of the genus Pomatiop. 49:60. 15(2) 178-186. 296. 49:10-13. 46(2) :48-49. 78:190-191. Illinois State : New Acad. Midi. Illinois Acad. 44(4) :119-121. cit. Canadian Field-Nat. K. 1929. London. 33:231-235. recent and fossil.. 322. pp. 298.. Jour. Indiana. in W. Sci.ssils in loess deposits. Trans. 1. A new form New species of Polygyra tridentnta from Illinois.iin with special reference to Pomon tioits/s Inpidaria. Jour.<> iimhilicatellu^ with the description of a new cit. The molluscan fauna of the great river valleys of Illinois.. (With Stanlev T. Nautilus. Nautilus. 49(2) :46-48. Op. new name for Limnaea montana Elrod. Canadian Field-Nat. species and varieties of Helisoma and Gyraulus from Canada. Mollusca from Turkey Run State Park. 80:118. Studies on the bottom fauna of fresh-water lakes. 311.. 12(3) :389-396. group of Planorbidae. Op.Chronological List of Publications 1931 293. Cahn). : new 1932 301. 23(11) :520-523. 47:30-32. Illinois State Acad. Proc. Canada. 309. (Abstract). 318. Morphol. variety of Valvata Icwidi Cm-rier.. 302. 314..s. 317. 1935 48:135. Pulmonate Mollusca peculiar to the Pleistocene period. New Lymnaeidae from the United States and Canada: and other western states. Soc. Nautilus. 324. cit. 300. 308. 48(l):17-20. Additional notes xxxiii 295. 46(1) :6-9." 1934 313. 299. 320. Trans. 312. 323.. Sci.. Nautilus. Stognicola apicina and Stagnicola walkeriann. 321. Canadian Lymnaeas. 310. Massachusetts. The ecology New A of Say's Limnaeus elodes. cit. California. 303. A re-studv of the interglacial moUuscan fauna of Toronto.. A new mammoth record for Illinois. The classification of the large planorboid snails of Europe and America. Land and fresh-water molluscs from North Star Lake and vicinitv. Sci. (With A. Two new races and a new species of Helisoma from California. Andover. Description of a new species of Gyraulus. Itasca County. Nat. The genitalia of the Indian mollusk. Op. Trans. Paleontol. Midi. particularly the loess deposits. 305. 5(3) :270-292. 1933 306. Science. The variation and distribution. 25:189. Ecology. on animal life associated with the mound builders of Illinois. new race of Pohjgyra apprcssa from Illinois. 145-149. Sci. cit. ^ Geol. 48:69-70. 315.). 13:286-289. Molluscan shells from Etowah Mounds. 307. 47: 140-142. 47:35. Mus. 33: 358-366. A new species of Gyraulus from Canada. 48-27. Illinois State Acad. Fresh-water Mollusca from central Ontario. Natl.. species and races of Lvmnaeidae from Newfoundland. Washington Acad. Science (n.' Department of Archeology. Canada. Stagnicola elrodiana. Ann. Nautilus. Nat. Phillips Academy.. Proc.scan fo. Soc. (Abstract). The variation and distribution. Plei. cit.. generic po. : New The Brooks). Ecology. for 1934:372-373.. 1931 (2):575-592. 47:4-7. R. Chapter V.. in Illinois. 41-64.. 26:129. cit. in Illinois. Zool. (With Junius Henderson). of the snail Polygyra profunda. 316.. Rept. Op. Trans. Nautilus. new Stagnicola from Montana.. Amer. Oregon. Canadian Field-Nat. pp... Stratiographic sequence of mollu.sition of Planorbi. 24(2) 149-155. 297. Sci. Nautilus. Op.. 327. Pohjgijru dorfcuilliann and Bulhuoides dcalbatus in Illinois. Nautilus. Indoplanorbis exustus. 325. 45:139-140.stocene historv of the terrestrial Mollusca of Fulton Countv. 304. 16(3) :257-272. A 55(1):1-13. Description of a 294. of the snail Polygyra profunda. . Amer. 48(3) 105-106. recent and fossil.

29:243-246. 'The Kingston Site. Midi.. 194. The Auk. 329. M. 341. 10(l):72-76. Cit. Nautilus. 195. 183-187. Natl. 79:1-37.s. 50(4) 113-117. Philadelphia. 53:106. Illinois State Acad. 1937. Mollu. The relationship of Parapholyx. new Pleistocene race of Polygyra apprcssa. 54:388. Canadian Jour. 53:138-141. 356. In Cole and Deuel. 10:1-223.). A new A A species of Drcpanotrema and some preoccupied planorboid names. A new HcIiso)na from the Pliocene of Florida. A new anatomical classification of the Planorbidae. 210-212). Op. Canada Bull. Simpson. Sci. Minnesota. 52:106... 330. The case of Linmacu. 32(l):51-67. Soc. Trans.. Op. Land and fresh-water Mollusca of New Hampshire. An Illinois record for the little brown crane. Manual 2. Quantitative examination of molluscan fossils in two sections of Pleistocene loess in Illinois.' Logan Mus. 347. Nautilus. Abundance of the European starling in Illinois. cit.. 345. March.. 49(3) ". 331. Amer. 1937:67-74. Trans.104. Nautilus. : A 1938 land and fresh-water Mollusca from the Ui^per Pliocene of Kansas and a new species of Gyraulus from Early Pleistocene strata. Illinois. Op.B. 342. 1941 352. 50:30-31. Lugn's 'The Pleistocene Geology of Nebraska. 332. study in ethnology of the prehistoric Indians of Illinois. 5:185-225. Stagidcola bulimoides vancouverensis nov. II. 55:105-106. Nautilus. The generic position of Plonorbis obstructus Morelet. Amer. 17:87-102. Saskatchewan. Land and fresh-water Mollusca from western Ontario. Surv. New : 1937 335. Surv. In 'Symposium on Early Man. cit. The Mollusca of the shell heaps or escargotieres of northern Africa.. Bull. 55:16-17. Research. : New Bull. 1939 343. 354. 337.. J. cit. Peoria Acad. . 338.. 334. Remains of animal life from the Kingston kitchen midden site near Peoria. 350. Lymnaea . Illinois. 1942 355. Op. Philadelphia. cit. Michigan..' pages 269-270. Amer. Philos. Jour..' held at Academy of Natural Sciences. Lvmnaeidae from the United States and Canada. 336. 'Rediscovering Illinois. 333. Op. Hist. Fieldbook of Illinois land snails. 340..' Appendix VI.xxxiv The Molluscan Family Plauorbidae 1936 328. 1939:13-15.' Nautilus.' Nebraska Geol. Op. 85:564. Part II. 351. Rept. 346. Contributions to the archaeologv of the Illinois River vallev. Tenth Ann. The fresh-water mollusc Hclisuma corpulentum and its relatives in Canada. Paleontol. 353. Pleistocene land and fresh-water Mollusca as indicators of time and ecological conditions. Remains of animal life from the Kingston kitchen midden site near Peoria. Nat. 1940 348. cit. 51(l):22-23. Science. Sci. Nautilus. 'Explorations in and around Fulton County. 344.'< sordidus Ku. Illinois Nat. Original copies of Say's 'American conchology. 339. L. 27:74-85.. 349. 166 pages. 190. Paleontology. Union. (Identifications and comments on pages 122.'itagnalis and Lymnnca (Radix) nuricularia. Part III of 'Prehistoric habitation sites in the Sahara and North Africa. new race of Parapholyx effusa. from the Prince Albert National Park. (n.. 54:96-97. D. Malacol. Nautilus. South American species referred to Planorbula. Nautilus. 51(4) 126-131. In A. Mus. 54:17. and Montana. 123. var.' by A. 49(4) 127-140.ster. Identification of shells. Lippincott. 1940:4-5.

Ann. M.' bv Russel Suter and Emcollected meline Moore. The Champlain Sea. W. BOOK NOTICES AND REVIEWS (In Ecology) 1921 The fresh-water Mollusca of the Parish of Aldenham. Pilsbry and J. by R. The Mollusca of Oundle. XI. and mountain ranges between the San Pedro and Santa Cruz riv(>rs. Baker. 4:431. 1941:5-9. B. 5:220. 55:130-132. 4:319. by A. new Gi/raulii. Winslow. E. Stream pollution. by Shira. Mollu.^ca of the southwestern states. Malacol. by Mina L. 5:222. Allen. 3:344-345. The Mollusca of Dickinson County. by I. by H. 3:340-341. Augustus Addison Gould. 5:221. Coker. A re^"iew of 'Stream pollution studies. Victor Sterki. Tryon. Grier. by H. From the Tucson Range to Ajo. F. Wevmouth. 2:314. Studies of the biologv of fresh-water mussels. Mexico. Natural historv and propagation of fresh-water mussels. 1922 The Mollusca by the University of Michigan-Walker expedition in southern Vera Cruz. Robert Kennicott. Mollusca contained in the test pit deposits. Correlation of shape and station in fresh-water mussels. 4:319. Urbana. 1923 the effect of salinity on Teredo navalis. Carnegie Inst. 3:318. Danglade. Mollusca of North Dakota. The Molluscan Family Planorbidae. Y. Kendall. . Rei)t." 3:342. of Washington. 4:318. by E. 3:345. Feniss. Collect inji 358. C. by H. III. 2:230. On The Kentucky River and its 1924 mussel resources. Field. Blum. xxxv A and preserving fresh-water snails. bv H. mussels. 4:318. Distribution and movements of Winona Lake mussels. by N.^ from the Pleistocene of California and a new Parapholyx from a supposed Pliocene deposit in Oregon. 1943 359. A. Boycott. bv Winifred Golding. Publ. Nautilus. C. by A. Biology and economic value of the sea mussel. 1945 360. and Howard. E. Arizona. 3:343. 1932. The Mollusca collected bv the Universitv of Michigan-Williamson expedition in Venezuela. by W. Clark.Chronological List of Publications 357. Amcr. George Washington 1933. by Rev. B. BIOGRAPHICAL CONTRIBUTIONS (In Dictionary oj American Biography) 1929. Variations in the shell of Teredo navalis. Baker. 1930. 538:117-119. The edible clams. Ortmann. H. E. William Stimpson. Miller. Baker. 5:306. bv F. B. and scallops of California. The University of Illinois Press. R. A. 2:231.'^ rdulii^. Mytilu. Union. Final report on the study and appraisal of mussel resources in selected areas of the upper Missi-ssippi River. bv H. Philadelphia. 1932.

Richard- 7:229-230. E. 5:416-418. 1920 and 1922. Chillicothe to Grafton: Its valuation. by R. 1925 of the Dutch Leeward Islands. H. by R. River bottom fauna in 1923. Richardson. Illinois in the small bottom fauna of Peoria Lake. Richardson. B. Krecker. 7:230.xxxvi The MoUuscan Family Planorhidae The small bottom and shore fauna of the middle and lower Illinois River and its connecting lakes. E. 6:320. Land and fresh-water Mollusca 1926 Changes son. E. and its relation to the fishery. 6:193. by H. Baker. Conditions imder which Goniobasis livescene occurs in the island region of Lake Erie. . by F. its resources of food supply. bv R.



more or less heavy and distinct. which may take over the function of breathing when the animal is submerged and can not obtain free air. and usually tapering to a point behind. not granular as in some of the land snails. The head and foot protrude from the shell and bear several important organs. . The Planorbidae differ from the related family Lymnaeidae in having certain of the organs on the left side (sinistral). called the pseudobranch or false gill. many blood vessels entering it from the lower part of the lung cavity. This procedure seems more logical since it is the animal which secretes and builds up the shell or calcareous covering (see plate 70. and there are no grooves on the body. jirovides a tube-like conduit for free air to reach the lung. The tentacles of the Planorbidae. A siphon-like extension of the mantle. wide. and are placed at about the junction of the velum with the foot. a modified lung as in the family Lymnaeidae. and tentacles. as in Lyinnaca. fig. and bears the head. it is usually somewhat semicircular in shape. This organ is highly vascular. is supplemented by a prolongation of the anal region (auriform lobe). in which the tentacles are flattened and triangular. The jiseudobranch is on the left side of the body and the pneumostome is to the right of this organ nearer the median line of the neck. The tentacles are very long and filiform. It is comparatively a recent organ. The variations of the pseudobranch are shown on the plates illustrating the anatomy of each genus. broadly rounded in front. varying from one-half to the whole length of the foot. It always bears a portion of the rectum and the anus opens at the upper right side. There is usually a ridge. which may be greatly expanded. 1). the organs in Lymnaeidae being on the right side (dextral). The foot is long and variably narrow. Physidae. called the pncumostome or respiratory opening. the animals resembling the Lymnaeas in this respect. leaf-like. being post-natal in origin. The pseudobranch varies greatly in shape. not having been observed in embryological EXTERNAL APPEARANCE OF THE ANIMAL CONTRARY TO GENERAL CUSTOM. In this respect the Planorbidae resemble the families Physidae and Bulinidae. This velum is broad and short and is capable of great extension and retraction. the characteristics A.I. The eyes are placed on swellings at the inner base of the tentacles. The breathing organ of the Planorbidae. which begins on the upper part of the rectum and ends in the middle or at the base of the pseudobranch. and Bulinidae are also similar and differ from the Lymnaeidae and Ancylidae. GENERAL MORPHOLOGY of the gross anatomy will be considered before taking up the shell and its variations. The texture of the foot is smooth. or rounded in different genera. being long and narrow. animal and its development. The mantle collar does not extend beyond the edge of the aperture of the shell. There is a large velar area above and in front of the foot which is retained from the embryonic animal. eyes.

the male opening just behind the left tentacle and the female opening a short distance behind the male opening in the side of the neck. muscular ring preputium penial gland penial gland duct pilasters ^ Male organs. The shell may be carried perpendicularly. especially over the lung and kidney. External form. albumen gland ' sperm canal diaphragm or Penial complex. the Planorbidae are hermaphroditic with the organs of the two sexes mostly separated. That self-impregnation of the female system by sperm from the extended penial complex is possible is easily believed when the closeness of the two sex outlets is observed. the animal having been killed while in breeding condition. f ovotestis ] ] 1^ ovisperm duct or hermaphrodite duct position the penial complex lies over the vagina and uterus (plate 39. or ultradextral. The color of the animal differs in the various genera. plate 24. 12). The body above the fig. The difi^erent organs may be tabulated as follows: vagma uterus oviduct ' vergic sac verge Female organs <{ nidamental gland spermatheca spermathecal duct . 3). 9). In some species the color is yellowish. 6. fig. 10). foot may be flatly rounded or it may be rounded above and flattened on both sides of the foot (plate 70. or it may lean to the left sicle. INIuscular. A narrow tube. When pulled away from this position its form is seen to be more or less cylindrical and to consist of a large bulbous lower portion. is in many groups heavily blotched with black or brown. and Nervous. the color being lighter on the base of the foot and on the tentacles. This descends to Male Organs. 11. grayish. There are often flecks or spots of white on the foot and tentacles. the vas deferens (VD) is inserted at the summit of the vergic sac. 7. Circulatory. fig. fig. as in Gyraulus and JMenetus (plate B. In the natural . the vergic sac (VS. Mostly it is bluish-slate or blackish. Renal. Digestive. causing the shell to be pseuclodextral 70.4 The Molluscan Family Planorbidae The genital outlets are on the left side of the neck. as in Helisoma (plate 70. [a] The Reproductive System Like other Basommatophora. 11). preputium (PR) and a smaller pyriform portion. Good figures of the animal of Tropicorbis are shown on plate 78. or brownish. GENERAL INTERNAL ANATOMY The internal anatomy includes the following systems: Reproductive. 7) . penial complex vas deferens prostate prostate duct muscles seminal vesicle Common to both male and female. The mantle. Respiratory. . fig. The most important internal structure for classification purposes is the reproductive system and this will be first considered. figs. In many individuals the penial complex has been observed protruding from the male opening (plate 29.

fig. but it is distinctly glandular and different from the ovisperm duct in structure. In Helisoma (plate 24. p. fig. 7. 11. cross sections of Hclisoma ca)npamdatum) The section is seen to be distinctly fanshaped. fig. 11) or short. see also plate 27. (plate 14. Mcnetus). 7. 1. plate 1.General Morphology 5 near the male openinti. This organ varies greatly in the different genera. The penial complex assumes many shapes in the genera belonging to the subfamily Helisomatinae. It also occupies the position above the prostate. This may be very long. Taylor (1900. 4 and 12. fig. 7). plate 39. p. fig. the penial complex becomes turgid by blood pressure. It is recognized as the seminal vesicle by Simroth (1912. In another subfamily (Planorbinae) there is no ])enial gland duct except in two or three genera. fig. plate 34. fig. DC). There do not appear to be any protractor muscles. 9. About midway of the ovisperm duct is a glandular enlargement which has been called the seminal vesicle (SV). figs. 5-8). figs. fig. fig. DCO). which in turn empties into the vas deferens below the prostate . 2) and a number of muscles believed to be supporting muscles to retain the penial complex in position. 14. DC) which normally lies coiled at a point between the vergic sac and the preputium (plate 27. There are one or two retractor muscles of the penial complex (plate 24. Just above the spermatheca the prostate appears. wlicre it is concealed in the muscular tissue of the neck of the animal and appears again near the female opening (plate 24. At its distal end the sperm duct unites with the oviduct to form the ovisperm duct or hermaphrodite duct (plate 27. RM. 6). plate 34. fig. The preputium may be pushed upward and the vergic sac may occupy a position on the side of the prei)utium as in Helisoma campamdatum (plate 27. The supporting muscles may in a measure perform the office of additional retractor muscles in some genera. 15). fig. PRS) it is somewhat elongated and fan-shaped and the compound diverticula empty directly into the sperm duct along a very small portion of that tube. 501) and by Larambergue (1939. 9. In some other genera the prostate duct is separated from the sperm duct and vas deferens and the prostate is greatly lengthened. fig. fig. RM. 14. p. the sperm duct (plate 27. The retractor muscles enter the columellar muscle but the supporting muscles are attached to the muscular wall of the forepart of the body. The prostate is composed of many or few diverticula arranged in a single series. 15 1. fig. A cross section of the prostate in Helisoma trivolvis illustrates this condition (plate 24. figs. In one group of species (the Helisomatinae) there is usually a duct (pcnial gland duct. the prostate the vas deferens is continued as a somewhat larger tube. figs. 13. The vas deferens is continued near the female complex and borders the uterus and oviduct where it is a small tube (plate 24. fig. plate 24. 94). 3. In some species the vas deferens is enlarged as it enters the vergic sac. where it lies in other animals. 9. Helisoma trivolvis). 15. as in HelUoma trivolvu (i:)late 24. the enlargement being similar to the epipliallus of the land snails (plate 39. as in Hclisoma anceps (plate 23. each diverticulum being attached to and emptying into the separate prostate duct. 11. especially in helping to retract the prel)utium after eversion. plate 24. fig. 359) calls the small sac-like organ into which the duct of the albumen Bevond . SPD. 15. Many students of both freshwater and land pulmonates simply call this region an enlargement of the ovisperm duct.

The ovisperm duct is a small tube of variable length. Gyraulus deflectus obliquus) diverticula may be filled with both ova and sperm. 1. fig. 6. The vagina to Both Systems. This gland or bulb is the carrefour which receives the products of the albumen gland and from which they enter the oviduct (see plate 34. plate 15. figs. male system VG) usually a thick walled. at which time they are fertilized by the stored sperm. . plate 35. the two organs being more or less synchronous in this respect. . a valuable feature in classification. 6. plate 1. Carinifex jacksonensis. Gyraulus albus). 9. fig. but as this organ is an adjunct of the in all animals it could scarcely be located in an organ of the female system. 359) erroneously calls the carrefour the seminal vesicle. p. Carinifex jacksoriensis) Taylor (1900. In the Helisomatinae it is many-lobed. passes imperceptibly into the uterus (U) which is usually much enlarged. fig. Both ova and sperm are produced in the same diverticulum. A small duct extends from the albumen gland and enters a large bulbous swelling which is attached to the end of the oviduct before that organ combines with the sperm duct to form the ovisperm duct. . sometimes called egg gland) follows the uterus and is usually placed over the uterus and the oviduct. The albumen gland (AL) is a large. 1. enlarged near the middle to form the seminal vesicle. Planorbis planorbis. In its natural position it lies over the stomach and a loop of the intestine passes under it causing a channel to be formed on the under side of the albumen gland (plate 38. fig. the diverticula branching fan-wise from the ovisperm duct (plate 27. Helisoma campanulatum . Drepanotrema hofjmani). They are said to arise simultaneously from indifferent epithelial substance and may in the liver Organs Common and fills . Menetus opcrcularis. fig. The oviduct (OD) lies back of the gland and is much narrowed as it passes backward to join the ovisperm duct. 4. Carinifex ponsonbyi. plate 14. the seminal vesicle. more or less elongated organ. fig. 9. The seminal vesicle is variable in form in the different species and genera and provides another feature for classification purposes (see plate 35. Carinifex ponsonbyi). composed of many small alveoli. Female Organs. fig. 9. In the Planorbinae and Scgmentininae the diverticula are in pairs. plate 34. The spermatheca or receptaculum seminis (S) is a bulbous or pearshaped organ with a narrow duct of variable length which enters the vagina at its junction with the uterus. Its function is said to be that of receiving the sperm (in the form of a spermatophore) during copulation and to store it safely until the eggs descend from the ovotestis. fig. 7. The swelling near the albumen gland is the carrefour. the nidamental gland (NG. 5. Gyraulus hirsutus) The organic relations between the albumen gland and the oviduct are somewhat complicated.6 The MoUuscan Fcunily Planorhidae gland enters. the two diverticula are attached directly to the ovisi)erm duct (plate During the breeding season the 16. The ovotestis (OT) is embedded about one whorl of the shell. fig. ( . The method of junction of the oviduct and sperm duct with the ovisperm duct varies in different genera. 2. A large sacculated body. wide tube. fig. It varies in form much as does the prostate. the ova either ripe for expulsion or developing (plate 10. The vagina is usually very short but may be of considerable length in some species. OT. 7.

Some cytologists declare that fertilization occurs in the ovotestis diverticulum and that the fertilized egg descends and is deposited without the necessity for copulation. fig. 2. In section it is seen to be cup-shaped. Gyraulus deflectus obliquus). 6. to be lined on the sides with folds and to have many small folds in the bottom of the cup. The preputium always has several vertical ridges on the inside called pilasters (PL) which may serve to guide the verge or the penial gland during protrusion. ST. In other genera there may be a fleshy papilla or appendage at the end of the verge (PA) with the outlet of the canal in the middle at the base of the papilla (plate 2. 200. 7. or elongated (plate 23. . The male organ. 75). descend during copulation.. 8 and 14. and Asiatic jilanorbids. 1927. 10. The gland is attached to the inner wall of the preputium either midway between the male opening and the muscular ring (diaphragm). Helisoma trivolvis. Gyraulus deflectus obliquus. fig. 201) the writer called attention to a peculiar gland found in the preputium of the genus Helisoma. 5. plate 24. Menetus cooperi) or it may be a dependent papilla (plate 23. which may be short and fat (plate 1. (TryonI are shown on plate 52. fig. plate 24. In 1926 (Trans. It is thought by most malacologists that the spermatozoa ripen first. The verge may be modified at its extremity by the presence of a daggerlike object called a stylet (plate 16. 5. fig. Helisoma anceps. See Clapp 1927 for a discussion of this subject. fig. Subsequent study has shown that this organ is found in several other groups or genera of American. figs. species. Euroi)ean. This gland is of variable shape. Drepanotrema hoffmani). fig. V. and that fertilization occurs when the ova and sperm meet in the lower part of the uterus or vagina. which I have called the penial complex. plate 14. Moietus cooperi calioglyptus) Other slight modifications are found in some genera. a lower swollen or bulbous portion called the preputium (PR) and an upper portion called the vergic sac (VS). All grades of length and diameter between these extremes are to be found in the different genera. 6 Gyraulus albus) In these examples the outlet canal for the passage of the spermatozoa is at the side. Typical eupyrene spermatozoa and nucleated o^'lm^: of Gyraulus circumstriatus p. 14-16. 6. This may be a simple ring with a central perforation (plate 39. 15. fig. pp. figs. or somewhat nearer this ring in some . fig. Between the vergic sac and the preputium there is an organ which I have called the diaphragm (D). The latter contains the penis proper or verge (V). Planorbis planorbis) or very long and narrow (plate 10. . 6. 5. rounded. plate 16.General Morpliology 7 be found developing together in a single diverticulum (see Crabb. The whole organ is divisible into two parts. contains a number of features which are of great value in systematic diagnosis. just before the depositing of the egg capsule. 6. and are stored in the spermatheca of the female system until the ova ripen and descend through the oviduct. Acad. Segmcntina nitida) In still other genera the termination of the verge is simply rounded with the canal outlet in the center (plate 38. The Penial Complex. Helisoma trivolvis). 1 and fig. fig. Helisoma anceps. iiyriform. This is a muscular ring which separates the two parts of the penial complex. . Wis. figs. 22. fig.

7) the verge and penial gland protruded together from the male opening. The duct terminates at the muscular ring or diaphragm (D). 10) also shows the canal in the muscular ring. 3. Helisoma anceps shows this feature well (plate 23. Promenetus exacuous. It appears possible that one of the functions of the penial gland might be to expand the opening in the muscular ring or diaphragm which is normally nearly or quite closed. 7). 7. . The long duct begins at the bottom of the cup and terminates in the vergic space (DC) of the vergic sac (VS) just above the muscular ring or diaphragm (D). fig. figs. H. figs. The bottom of this cup is. 2. 7. figs. In Menetus and some other groups there is a small channel in the muscular ring (plate 39. fig. the walls of which are lined with vertical folds (GF). figs. Helisoma anceps) When coiled on the outside of the preputium it is bound to that organ by short muscles. as Helisoma duryi and its varieties. 1. but also the whole penial gland. simply lies on the surface of the preputium (plate 23. there is no external duct. Parapholyx effusa klamathensis (plate 36. 12. the duct begins in a small cavity or cup at the end of the gland (plate 33. 4) the gland shows a large cup-shaped cavity. and plate 27. 3. 581). fig. plate 41. fig. fig. 4). fig. In all species of Planorbidae the verge is extruded during copulation. each of which may have a different function. 10). trivolvis lentuni. fig. if long. 6). 11. the walls of which have vertical folds. 1. 1. But there is another function of the gland. Another group of Menetus has a short duct on the inside of the preputium (plate 39. and piercing the integument of the prea duct of variable length (DC) which. Helisoma trivolvis) or in a groove in the ring (plate 36. pierces the wall of the lower part of the vergic sac. In section this organ is seen to have two parts. figs. 8 The Molluscan Family Planorhidae From the base of the cup. plate 24. fig. This has been observed in many species of Helisoma (H. 9. 10. . Parapholyx effusa) The function of the penial gland is at present insufficiently known. The larger cup-shaped cavity has no connection with the smaller cavity or its duct. to the diaphragm to facilitate the passage of the verge.. fig. p. fig. figs. 2. is . plate 29. plate 33. and terminates just above the ring (plate 23. 1. 1931. notably in the genus Menetus. and prepare it for the protrusion of the verge. 9). secreted by the vertical folds in the gland cup. lined with folds laid crosswise. duryi normale. The penial gland duct possibly carries a fluid. fig. Helisoma anceps. 9. It was once thought by the writer to be connected with the formation of the spermatophore and was called the spermatophore gland (Baker. . for in the species possessing this gland there is no special apparatus for this purpose such as occurs in other species putium. Sections through the cup are shown on plate 23. 2. fig. In some species. 16. Helisoma campanulatum) or. 6. It has been observed that during copulation not only is the verge extruded from the male opening. 6. . if short. 9. is folded on the outside of the preputium (plate 27. fig. It is probable that the gland portion with the cup-like cavity lined with vertical folds is an excitatory organ or sarcobelum. In Helisoma trivolvis (plate 24. In some species. but this belief scarcely agrees with the structure of the gland. connection with the vergic space being effected by an open channel (plate 42. In a specimen of Drepanotrema lucidum examined (plate 11. D). plate 24.

and while the animals were seen in apparent coitus on several occasions. then the gizzard. There are retractor and protractor muscles and nerves received from the buccal ganglia. only the spermatheca and its duct vary in form and length to any degree. especially before and after breeding. and other genera). This condition occurs mostly in HcUsoma (plate 25. but includes also the ]:)rostate and seminal vesicle. As in the Lymnaeidae. The stomach is a large muscular body composed of three parts. a duct entering the buccal sac on each side near the oesophageal connection. figs. The male organs. the gland being pushed upward in the preputium causing the vergic sac to appear as though attached to the side of the preputium. the protrusion of the gland was not seen. first the crop. More than a dozen acjuaria were kept under observation by the writer for several years. which is an indistinctly bilobed body of large diameter. that this is not so easily accomplished as might be supposed. These organs are very uniform in the different species of the family (see plate 70. [b] The Digestive System digestive system of the Planorbidae is similar in many respects to that of the family Lymnaeidae. since the characters of the different groups are constant and uniform. When ojiencd it is found . 7 being the nonual form and figs. this modification occurs mostly in the male organs. Observations of the living snail in aquaria should help to solve some of the questions. are composed of The many small lobules. Anisus. figs. Another feature observed in connection with the penial gland is that the i)enial complex is often seen in a bilobed condition. The oesophagus is usually rather long (about one-fourth as long as the intestine). and 8 showing the gradual change to the bilobed condition. As was recorded in the study of the family Lymnaeidae (Baker. and the salivary glands. w'hich is an enlarged portion of the oesophagus. Much remains to be learned about the function of the penial gland. the second part the region of the stomach with the liver connections. plate 48). 1. the first consisting of the buccal sac. principally the penial complex. 1. The evolution of this feature was observed in specimens of Helisoma ccunpanulatum and is shown on phite 27. 12. therefore. Perhaps histological studies would help. 7. afford useful features upon which to base a natural classification of the family. usually cylindrical and elongated.General Morphologij 9 which have a horny stylet or fleshy papilla at the end of the verge {Gyraulus. 6. plate 24. It may be observed. fig. the oesophagus. and the third part the intestine and rectum (see fig. 6. there is great variation in the form of the genitalia of the genera of the family Planorbidae. beginning as a tube of small diameter and enlarging as it approaches the stomach region. 13). figs. 1. also. or opening into the intestine. The gizzard is not as distinctly bilobed as in Lymnaea but is of the same nature. The subject is worthy of a doctor's thesis. 1911). however. The two glands are joined behind and form a continuous loop or ring. 9. The salivary glands. 5. The buccal sac is usually pear-shaped. The female organs are uniform in position and shape. and third the pyloris. the oesophagus entering it at the upper middle portion. The system naturally divides into three parts. 2). The salivary glands are paired.

The striations of the superior jaw of the first group become separate vertical plates in the second group. The anal opening is always near this ridge on the pseudobranch. supports a central ridge or crest on the upi5cr side which extends to and over the pseudobranch. Another type is horseshoe-shaped and in this there is but one division which appears to represent the fusion of the lateral and superior jaws of the first type. the striations producing a saw-like condition on its lower. there is a finger-like process or blind sac which appears to be the stylotheca containing the crystalline style. producing a segmented jaw. [1] though in intestine sharply borders some species it extends of the Planorbidae is provided with two important organs which afford characteristics for classification. superior and lateral. These side jaws form a question-mark figure. The jaws of the Planorbidae fall into two principal types or variations. The jaws are moved by several distinct muscles. It has two unequal lobes which form the posterior part of the animal and in which the ovotestis is concealed. It is well supplied with blood vessels which form a plexus on and in the organ. fig. A large duct enters the pyloric portion of the stomach at the point of junction with the blind sac. In the upper part of the T lies the superior jaw. It then forms a single loop wdiich bends forward and extends to the fore part of the animal on the left side (the right side in the Lymnaeidae).10 The Molluscau Family Planorhidae taken in to contain small particles of siliceous matter. more or less digitiform. fig. brown. 8. with On the under side of the pyloris. 13). which is wide and low. The liver is composed of a great number of small lobules. Internally the sac contained a gelatinous substance or was empty. to the right and above this organ. striated vertically on its front part. The loop 1) of the the stomach in some species (fig. near the duct of the liver. two narrow. This blind sac is present in all species of the family examined. The liver or digestive gland covers the posterior part of the stomach region in its natural position. The most common is the group described above. The rectum. Helisoma anceps) The lateral jaws. This segmentation extends to the lower part . are attached to a cartilage which in turn is attached to the roof of the mouth. 1. as will be noted in the figures on plate 70. As in Lymnaea. the jaw and the radula. It bends around the stomach from the end of the pyloris and passes backward for a distance. The jaws. horny ribbons. 8). The intestine is very long. of a brownish color. Helisoma trivolvis) or elongated and narrow in other species (plate 48. cutting edge (plate 49. forward without a loop (fig. fig. evidently sand the food to serve for the breaking up of the ingested food. The Jaw The mouth . Bathyomphalus contortiis) . They are connected with the lateral margins of the superior jaw by a narrow projection from the upper part of the lateral jaws. emargin the lips or sides of the mouth. the greatly enlarged portion of the intestine. with large superior jaw and two smaller lateral jaws. short and rotund in some species (plate 48. fig. 1. The horny jaw lies in the upper part of the mouth at the fore end of the buccal sac (plate 70. Variation in the digestive system is noted only in the form of the stomach region. the mouth is somewhat T-shaped in repose.

and a smaller outer ectocone (see plate 57. There are protractor and retractor muscles which iiull the radula backward and forward. xiii). fig. The marginals of the radula also indicate a primitive form of radula. 12. fig. 16. There is a series of duplicating teeth on each side of the central tooth. Carinifex jacksonensis) Althcnigh belonging to the subfamily Helisomatinae the jaw is practically of the Planorbinae type. the radula is moved from behind forward. The lateral teeth are usually tricuspid and comprise an inner. not included in the usual segmented type of the family. as described by Sterki many years ago. as in the Lymnaeidae. showing that tlie hiteral jaws are inokidcd in the segmentation (see plate 50. over its cartilage. figs. The front end of the radula extends toward the mouth of the snail and the rear end is bent downward into a radular sac which forms a rounded protrusion lies or bump at the back of the buccal sac (see plate 70. The lingual ribbon of the Planorbidae carries many teeth in transverse rows as in the family Lymnaeidae. As stated by Pilsbry this law is as follows: All modifications of the teeth proceed from the median line of the radula outwards toward the edges. The mesocone usually persists in the earlier marginal teeth and may be recognized by its large size and central position. is that of Cariuife. as splitting of the ectocone or entocone into smaller cusps. during the act of cutting u\) food particles. p. In use. The intermediates are those teeth between the typical lateral and marginal teeth which show certain modifications.000 teeth in one ribbon. fig. showing a close relationshii) between these two . like the tongue of a cat lapping food from a plate. and marginal teeth. 18. a large median mesocone. [2] The Radula The radula of the Planorbidae is ribbon-like as in the Lymnaeidae. A peculiar jaw. The central tooth is always bicuspid and in some groups (as in Drepanotrema) there are one or more accessory cusps on either side of the central cusps. iacksonensis) This type of segmented jaw is suggestive of the polyplacognath group of land snails. Gijraidus parvus). Carinifex ponsonbyi.. The marginals are usually somewhat claw-like with a variable number of small cusps. 19. The radula grows forward from this sac as the functional teeth at the front end of the ribbon become worn by use. The law of mesometamorphosis applies to the Planorbidae as well as to the Lymnaeidae. into lateral. . The marginals of the Planorbidae are of the same ])rimitive type as are those of the Lymnaeidae. 13). intermediate. the outer marginal teeth being the last to be modified (Guide to Study of Helices. There may be as many as 200 rows with 85 teeth in a row or a total of 17. The cusps are usually dagger-like and may be wide or narrow. 1). This row is divisible. short entocone. Figure 17 on plate 49 shows the large size of the cartilage in this genus (C. fig. General Morphology 11 of the side of the horseshoe.T in which the jaw is b()w-shai)ed (or roundly horseshoeshaped) and is composed of many narrow plates fastened to a heavy cartilage (plate 49. It on a subquadrate cartilage near the rear end of the buccal sac. It is said to be a very ancient type of jaw which would seem to indicate that the Planorbinae group of the family is more ancient than the Helisomatinae group (except Carinifex which has the primitive form of jaw).

working on another group of snails. 4. is regularly marked by internal the lumen for a greater or less distance. pointed out the fact that the kidney may reflect specific characters. a small upper saccular portion with the pericardium at the right (SK) and an elongated lower portion (TK). the tubular portion septa which project into side by a large vein. and discharges into the mantle cavity. have long and narrow reflections reaching far below the base of attachment and with the distal end and the outer margin multiserrate (see plate 57). Viewed from the ventral side. the pulmonary vein on the renal vein on the left side (RA). [c] The Renal System or renal organ (nephridiumt of the family Planorhidae is a large and conspicuous part of the snail's anatomy. usually more or less upward. The figures on plates 44 to 47 show the various forms of ureter and the relationships of the different parts of the kidney. 1. perhaps. A series of sections through the kidney of Australorbis glabratus (plate 45. This is a small tube which turns to the left. U. the genitalia being between the kidney and the ureter (see plate 14. the most conspicuous part being a distinct ridge (RK) To the left of this ridge (in the section) there is another conspicuous ridge on the roof of the mantle. wide reflections high up on the base of attachment and only a few^ serrations or cusps on the lower margins (see plate 69). In fig. fig. These exceptions appear to be due to a splitting of the three cusps of the normal lateral teeth (entocone. The tubular portion is surrounded by thick walls of spongy tissue. ectocone) found in the majority of planorbid genera. an arrow indicates the place where each . As with the other organs. figs. The other group. At the lower end of the tubular portion is the short ureter (UR). The lateral teeth are very uniform in both groups. In 1940. have short. the cusps rounded or sharp and dagger-like. A section through the middle of this portion of the kidney (plate 44. In studying the teeth of the Planorhidae it must be remembered that only the unworn teeth show the true characteristics of the radula. the kidney consists of two parts. It lies to the left of the lung cavity. indicating. causing the teeth to be 6 or 7 cuspid. The central of the kidney. the radulae offer sound criteria for classification purposes. One group. plate 45. 7 to 11) show the internal form of the kidney at different points in its length. mesocone. NG). including the genus Planorhis and other divisions of the subfamily Planorbinae as well as some of the subfamily Segmentininae. including the genus Helisoma and other divisions of the subfamily Helisomatinae. these being mostly tricuspid. The Planorhidae are divisible into two groups by the form of the marginal teeth. Also the teeth must be in their natural position on the radula ribbon and not tilted or otherwise disturbed. In several groups (as Drepanotrema and Segmentina) the laterals are modified by the addition of several cusps. 9) shows a central tube The kidney or lumen flanked on each right side (AP) and the tube. . that the Planorhidae may be somewhat younger in development. Mattox. occupying a considerable portion of the body whorl of the animal. fig. The lateral teeth of the Planorhidae show a wider degree of variation than do the lateral teeth of the Lymnaeidae.12 The Molluscan Family Planorhidae families and also their common descent from the tectibranchiate stock of marine mollusks.

as shown in figs. since many species of the Planorbidae live in habitats abundantly . the kidney is flattened and broadened and the veins are smaller. as shown in the figures on the plates. carrying blood to the auricle. and other systems in the posterior part of the body. however. RA). In fig. carrying blood to the organs in the head. but no exit of this nature was found in any species examined. is open to question. 9 and 10 (C. and shows the lumen partly filled with folds extending from the walls of the kidney. All had a short ureter. In figs. The Planorbinae are mostly without this ridge. genital. In fig. the branches becoming smaller. etc. partly reflexed. These arteries branch frequently. The blood vessels (veins) lie in close proximity to the tubular jiart of the kidney and pick up the blood to be carried to the heart. and the cephalic artery which bends around and forward. as well as on other plates. B) the same features are shown. [d] The Circulatory System The circulatory system of the Planorbidae is similar to that of the Lymnaeidae and other Basommatophorous Pulmonata. The ureter of the Planorbidae is described as straight. The heart lies to the right of the kidney (instead of to the left as in Lymnaea) and consists of a ventricle and an auricle enclosed in an oval pericardium (plate 44. VT). entering the saccular portion (see plate 44. 8 (D) the section shows the tubular portion of the kidney with a few folds extending from the walls. The venous system consists of blood vessels which convey the blood back to the heart after circulation through the kidneys. near the lower part (A). This statement. and terminate in contact with the venous system either by small arterioles or through lacunae or blood sinuses which connect the arterial system with the venous system. In the Planorbidae the blood (haemolymph) contains a substance (haemoglobin) which gives it a pinkish or red color. foot. 2. 2. 16 and 17 on plate 45. also the large pulmonary vein at the right and the smaller renal vein at the left. AP. fig.General Morplioloyy 13 section is made. All figures show the variation in the form of the ridge in different portions of the kidney. A large branchial vein borders the kidney on the right side. The ridge on the kidney is mostly confined to members of the subfamily Helisomatinae and offers a valuable criterion for the separation of this planorbid group. AU. The aorta (AO) extends backward from the ventricle and divides into two branches or arteries. 11. the visceral artery which supplies the digestive. and other parts of the system. A large renal vein also borders the left side of the kidney. PC. The folds iu' the tubular portion of the kidney are metameric in their presence and one section may pass through a portion of the kidney where these are present. fig. Its presence in Planorbis is said to be associated with the poorly oxygenated habitat occui^ied by most species of this group in stagnant pools or lakes and ponds poorly supplied with oxygen. This is an active substance occurring abundantly in the respiratory system and differs from the bluish blood (liaemocyanin) which occurs in most mollusks. Venous sinuses are present and store the blood after circulation to be carried to the renal and respiratory systems for purification. lung. or it may pass between two sets of folds. Figure 7 (E) is a section through the sacculate part of the kidney below tlie pericardium.

Two pleural ganglia are connected with the visceral. 2. 14 The Molluscan Family Plauorbidae supplied with oxygen. The left cerebral ganglion sends nerves to the penial complex (plate 48. Helisoma anceps) which divide to send branches to the vergic sac. [e] The Respiratory System Respiration in the Planorbidae is effected as in the Lymnaeidae and other aquatic pulmonale mollusks. 3. Free air is admitted into a large mantle cavity abundantly lined with blood vessels forming a plexus on the dorsal wall of the mantle cavity. the pseudobranch. next to the kidney. 1911. The respiratory. . The rectum usually has a large crest or ridge on its upper surface which diminishes in size at the lower part where it extends over and to the edge of the pseudobranch (see plate 25. There are two cerebral ganglia above the oesophagus (CG). In the Planorbidae the presence of the secondary brancliium. P). It is abundantly supplied with a plexus of blood vessels which absorb oxygen from the water. A) usually above and about at the middle of the pseudobranch. 8. 4. . fig. 3. This lung cavity lies on the right side of the mollusk. but is always an outgrowth of the branchial cavity walls and lies to the left of the pulmonary siphon to which it is attached (plate 14. and is bordered on the left by the pulmonary vein (plate 44. as well as to other parts of this comj^lex. fig. two buccal ganglia (BG). Below the oesophagus there are two pedal ganglia (PG). figs. because the group is sinistral. fig. P). 17. enables the animal to respire without coming to the surface. a dextral group. is [f] The Nervous System The nervous system of the family Planorbidae differs little from that of the family Lymnaeidae (see Baker. 1911. Helisoma pilsbryi. Physa) but on the left side in dextral species (Lymnaea). In the Lymnaeidae. fig. 18). A nerve ring surrounds the oesophagus posterior to the buccal sac (plate 48. L). the nerves arise from the right ganglia (see Baker. and circulatory systems are intimately connected and should be considered together to understand the conditions governing the respiration and blood circulation of the animal. plate 5). plates 1 and 5). pedal. It may be a simple rounded fold or it may be flat with a distinct ridge down its center. plate 28. The lung cavity is placed on the right side of the body in sinistral species iPlanorbis. The pseudobranch is probably helpful in supplying oxygen to the blood. fig. 4. The rectum passes through the pseudobranch and its outlet (anus. in company with Lymnaeas and other fresh water piihnonates. fig. as it is highly vascular (plate 14. Hclisoma scalare.. plate 35. and retractor muscle. and a single abdominal ganglion. Carinifex jacksonensis) The breathing orifice or pulmonary siphon (pneumostome) is an outgrowth of the mantle which is capable of forming a tubular conduit or siphon through which free air may be admitted to the lung (see the plates mentioned above). In the Planorbidae these nerves are sent off from the left instead of the right ganglia. fig. 4. penial gland duct region. renal. two visceral ganglia. This organ is of various shapes. and abdominal ganglia.

11). There may be. 45).. and have been noted in the genera Helisoma and Carinifex. like a watch spring. But the shells are not all strikingly Sinistral. as Helisoma pilsbryi. As most species are planorboid in form there arc few modifications of the columella. THE SHELL The shells of the Planorbidae are mostly orb-shaped. and are apparently dextral and are usually described as pseudodextral or ultradextral. But the smaller species are tipped to one^ side. Cincinnati Soc. as are found in the familv Lvmnaeidae. and of few or many whorls. It may be armed with lamellae or entirely free from such obstructions. and Indoplanorbis are obviously sinistral. or it may be almost globose. The whorls may be rounded on the periphery or this region may be sharply carinated or squarely flattened. 19. usually exhibiting a distinct right and left side. It may be as thin as paper or thick and very solid. or disc-shaped. p.General Morphology 15 [g] The Muscular System The muscular system of the family Planorbidae does not differ from that of the family Lymnaeidae except in i)ositions due to the sinistrality of the animal. 1896. mm. Sutures may be deeply indented or scarcely visible. Scalariform individuals may also occur in any species and have been noted particularly in the smaller species of the genus GyrauliLS. or it may be deeply indented forming a distinct umbilicus. Planorbarius. or loosely coiled. while truly sinistral. Hist. Nat. ovate. The shells of the larger speLcies^_especially in_ Helisoma. to the large Helisoma trivolvis macrostomum. Such types sometimes occur in other species of the family as abnormal or pathologic forms.wh^ii ia loconiption. which attains a diameter of 30 m. It may be so fiat that one wonders how an animal can find enough room to function in so small a space. appear as dextral. Such large groups as Helisoma. In such species the upper side . The shell may be closely coiled. C. 2 In size the planorbids range from the tiny Armiger crista. rarely becoming a conical spiral as in Lymnaea and Physa. In most works on this family the shells have all been considered dextral and figured as such. wheel-shaped. to the left. fig. as Drepanotrema cultratum. also. Quite a literature has developed in the controversy as to whether the shells of Planorbis are dextral or sinistral (see Baker. fig. as the Helisoma scalare of Florida. The outer lip may be thin and sharp or it may be strengthened by a heavy ridge or callus. Several physoid types occur. 7) or it may lie'ahiiost flat as in Planorbida^nksi (shown on plate 70. The aperture is most frequently rounded. In all species of the family Planorbidae the ^nital organs are on the left side^ -and the animal is sinistral. The lower (umbilical) surface may be so flattened that no umbilical opening is indicated. These shells. are carried almost perpendicularly as in jHelisojnq trivolvisjnaa^o^tomum (shown on plate 70. (right) is umbilical and the lower side (left) is apical. less than in diameter. Jour. The most important muscles for classification purposes are those of the male generative system. m . more or less of a callus on the parietal wall. or half-moon shaped.

In some groups. Some shells are light milky in color. as in Menetus opercularis. In some groups the shell appears devoid of an epidermis. as Australorbis. A few species border on red. The epidermis or periostracum of the planorbids is usually rather thick and heavy. Malleated individuals are comparatively rare in the Planorbidae although they are common in the Lymnaeidae. but there are no species known comparable to the greenish shell and purplish aperture of Bulimnea magasoma. The remarks on sculpture in the Lymnaeidae Monograph (pp.16 The Molluscan Family Plauorbidae The sculpture of the planorbid shell consists mostly of growth lines be very fine and thread-like or coarse and rib-like. In some species there have been developed costae or ribs more or less evenly distributed. the color is rich brown or chestnut. 4) apply equally well to the which may Planorbidae. Spiral impressed lines are found in most of the species of the family. the handsomest of the fresh w^ater pulmonates. Zebra markings occur among the planorbids as among the lymnaeids. The color of the shells of the family Planorbidae is usually some variety of horn color. others are ashy. the surface being shining and waxy. . These are usually rather heavy and conspicuous but in some species they are very fine. The range of color is about like that in the family Lynmaeidae. In a few species the spiral lines are very faint or absent. light or dark. In a few species the epidermis forms paper-like crests or frills. 3. as in Helisoma anceps royalense.

that chemicals and oil. A few inhabit deeper water and in the deep lakes of Europe and Asia. as lakes Geneva. The family is on the whole a distinctly shallowwater group. however. especially if rapid. The maximum numbers occur in shallow areas not exceeding six feet in depth. 17 . deep dredgings have produced no members of the family Planorbidae. Helisoma pilsbriji. Helisoma corpulentum. It has been observed. altitude. such as Helisoma campamdatum. the usual limit of rooted vegetation in water bodies. Almost all of the members of the family Planorbidae are littoral animals and are not usually found in water deeper than fifteen feet. Tropicorbis. they may occur in abysmal depths of from forty to 350 meters. and some varieties of Helisoma anceps. reproducit now has such ECOLOGYsubjects that diverse tion. ]\Iany such in New England have been examined and a few in Idaho and ^Montana. etc. and Leman in Switzerland. development. are abundant in small pools which may become wholly or partially dry during diy seasons. from the largest lakes to the smallest pools. It is here made to include everything that affects the animal either from external or internal sources. as far as personal observations have indicated. GENERAL ENVIRONMENTAL CONDITIONS Representatives of the family Planorbidae may be found in almost any body of fresh water. and Lake Baikal in Siberia. and usually no other mollusks. GENERAL ECOLOGY a wide significance and includes so many is necessary to define what is embraced under this term in this chapter. relations in food chains. Streams of all sizes harbor one or more species of the family. are inimical to the Planorbidae and Lymnaeidae and none have been seen which could resist this deadly type of habitat. may be used by these animals successfully as habitats. parasites.II. In our own lakes. also wood wastes from factories. Lake Balaton in Hungary. Clear. Planorhula. The Planorbidae are able to withstand unfavorable features of their environment better than most groups of mollusks. and some Gyraulus. Helisoma trivolvis is common in small lakes or bays of larger lakes. Certain groups. members of the subgenus Seminolina live in limestone pools of limited size. In Florida. as Menetus. Constance. the animal busily eating such small organisms as may lie on the surface. ]\Iany of the large forms. climate. as well as water containing sewage. contain no planorbids. prefer larger bodies of water like the larger lakes. cold mountain streams. chemical conditions of the water body. including such subjects as the habitat. The Lymnaeas also share this ability to live under unfavorable conditions. the shell hanging downward. In such habitats the snails may be seen with the foot attached to the under side of the surface film. in shallow areas where the shore is bordered by Typha and sedge. Thus water of some alkalinity and salinity. owing to their ability to breathe free air. where food in the form of algae and other vegetation is abundant. pollution.

The Swiss lakes are at comparatively high altitudes as are also some of the lakes of Asia which contain moUusk faunas of some size. 18941900. Planorbula crassilabris may often be found on a lake shore fully exposed to the waves. 1909. the whorls. Quite a literature has accumulated on the general ecology and particular habitat relationship of the Planorbidae and other fresh-water mollusks. C. 1911. Dybowski. in Lake Eustis and Lake Griffin. varies from a typical planorboid form to a distinct physoid form by the elongation of the spire. Promenetus umbilicatellus is an inhabitant of ditches. VARIATIONS IN DIFFERENT ENVIRONMENTS snails do not exhibit as much variation due to differences environment as do the Lymnaeidae (see Baker. These examples might be greatly multiplied but enough have been presented to show that variation is the rule among individuals of the family Planorbidae and that this variation may not be due alone to conditions of the environment but rather to the inherent 'law to vary' which is found in all living things. 1918. and swampy places. in Peru. Tavlor. while Pronienetus exacuous is usually found in larger bodies of water. 1920.846 feet. pp.000 feet. 1874. 1916. Baker.000 to 10. which contains many mollusks. Great variation is often seen in specimens of the same species from one habitat. Taphius andecolus exhibits many variations in the form of the aperture.. 1924. The shell aperture often increases in size in large lake localities where the animal is subject to heavy wave action. In the Rocky Mountains they occur in lakes at altitudes of from 9. Florida. In Lake Titicaca. 1922. This may be observed in Helisoma anceps percarinatum and in Helisoma anceps sayi which are lake dwellers in contrast with typical Helisoma anceps which is usually a river or creek inhabitant.18 The Molluscan Family Plauorbidae Altitude does not seriously affect the distribution of the members of the family Planorbidae. In Carinifex the individual variations are numerous among specimens of one species from the same habitat. including a peculiar group of Planorbidae [Taphius) is at an altitude of 12. more expanded aperture. . Some species of this family show preferences for particular types of habitat. pools. The remarks on tlie lymnaeas in the 1911 monograph on that family may be consulted with advantage. 1875. Among some of the species of the genus Australorbis of South America and the West Indies there is often great variation in the axial height of the shell. 1909. 1869. These are listed in the bibliographv and may be consulted under the following names: Adams. and the in the The planorbid condition of the umbilical region. 1911. or in protected places in larger ponds and lakes. The family Planorbidae does not appear to offer as good a field for the distinguishing of the so-called habitat or ecological variations as does the familv Lvnmaeidae. 29-32). Forel. F.^ 1910. Thus Helisoma duryi seminole. Lindholm. in Peru. J. Helisoma canipanulatum davisi is a small pool or swamp dweller while typical Helisoma campanulatum is a lake inhabitant and usually has a larger. A number of special papers and books have been published which bear on this subject. Thus Planorbula armigera is usually found in ephemeral pools or small bodies of water. Lake Titicaca. W.

p. Fidalgo Island. Young and immature planorbids are usually very active. however.) Mouth Common ) . the animal using it from behind forward as described by Dr. as do the lymnaeids. Baker. FOOD FOR OTHER ANIMALS Together with other fresh-water mollusks. Potamogeton. concerning the food sujiply of this family. both the large floating leaf species and the submerged leaf species. 42 for notes on the food of lymnaeas). They have been observed eating pond-lily leaves (principally Castalia). Some planorbids are scavengers. Chief among the animal groups feeding on mollusks are certain species of fish (see F. the family Planorbidae furnishes food for a large number of other forms of animal life. p. vegetable fibers. have been found in the stomachs of the following food and game fish: Whitefi. C. The method most usually observed is that of gliding. though not to the extent observed among the lymnaeids. the latter usually being rather sluggish in motion.) Pumpkinseed (Lepomis gibbosus (Linn. and algae of various kinds.) ) Sucker {Catostomus commersonii (Lacepede) Yellow Bullhead (Ameiurus imtalus (LeSueur) ) Common Bullhead (Ameiurus nebidosus (LeSueur) ) Fre. by varying the amount of air in or on the body or shell. 1911. 32). Specimens of Helisoma duryi seminole from Florida had the stomach and gizzard filled with small grains of sand and nothing else. No evidences have come under observation indicating that the group is carnivorous (see Baker. eat- Locomotion ing everything in their i)ath. FOOD Like the lymnaeids the food of the planorbids is largely vegetal. The radula can be plainly (Coregotms clupeaformis (Mitchill) ) Small Buffalo (Ictiobus bubahis (Raf. Stomachs and crops that have been opened and examined have usually contained a quantity of fine sand (especially in the crop) which probably helps in grinding the food before it enters the intestine. The members of the family are usually very active. and a quantity of flocculent undigested animal matter have been noted in the stomach. including planorbids. Little is known. Both old and young snails will eat avidly of the algal scum which collects on the glass sides of an aquarium. much more so than fully mature individuals. Sterki many years ago. In locomotion the planorbids resemble the physas more than the lymnaeids. . 1916).) ) Bluegill (Lepomis macrochirus Raf.General Ecology 19 LOCOMOTION in the family Planorbidae is accomplished by the same means as in the family Lymnaeidae (see Baker. 1911. Tlie planorbids have been observed to rise suddenly and descend\vater Drum (Aplodinotus grunniens Raf. v. Washington) distomids. Ostracods (six in one specimen of Helisoma subcrenatum from Pass Lake. Fresh-water snails. crawling about on aquarium walls quite rapidly.

newts and salamanders. eat fresh. such as the Lesser Scaup (Nyroca affinis (Eyt. consume some mollusks Ducks. in diameter.water snails.. . are a valuable food source for the larger animals. These are: Gyraulus parvus (Say) Gyraulus deflectus obliquiis (DeKay) Promenetus exacuous (Say) Helisoma anceps (Menke) Helisoma campanulatum (Say) species of birds. It is obvious that the planorbids. The waders. It appears possible that these specimens would have attained full maturity in another year. frogs. four or five years it is probable. Specimens of Helisoma trivolvis lentum were hatched in an aquarium and lived sixteen months. The stomach contents of many fish and birds have been listed as simply snails or Planorbis without identification of the species involved. Helisoma duryi normale has been hatched in an aquarium and has lived for over a year. Many nymphs. leeches.) ) and others feeding in shallow water. An adult lentum has five whorls and is about 20 mm.) ). Helisoma particularly. in diameter and had four whorls. or two and a half years from time of hatching. At this time they were about 8 mm. It is quite probable that many other animals which frequent fresh-water snail habitats feed on the smaller species or the young of larger species. not of direct value as food or game fish. Mature individuals of Helisoma trivolvis have remained alive for nearly two years in a wellbalanced aquarium. especially water birds.) ) has been known to feed on a planorbid (probably a Gyraulus) Other groups of animals which include fresh-water snails in their diet are turtles. Such small fish known to eat snails are: Fresh Water Killy {Fmululus diaphanus (LeSueur) ) Killifish (Fundulus dispar (Agassiz) ) Viviparous Top Minnow {Gamhusia affinis (Baird Girard) ) Manitou Darter {Pcrcino caprodes zebra (Agassiz) ) & stomach contents of Five species of the planorbids have been specifically identified in the fish. feed on snails. and other snails of the fresh waters. are yet of great value as food for the larger food fish. crayfish. LENGTH OF LIFE The life span of the members of the family Planorbidae is not definitely known. Just how long any of the planorbids live in their natural habitat is not known. 20 The Molluscan Family Planorbidae Many smaller fish. would be of value. Experiments with the larger snails of the family Planorbidae. The lymnaeids are said to live for three or four years. An adult Planorbarius corneus (the red snail) has been kept alive with goldfish for two years. and dragonfly in their diet. Many of these include the planorbids and other fresh-water snails in their diet. The Red-winged Blackbird {Agclaius phoeniceus (Linn.) ) and the Killdeer (Oxyechus vociferus (Linn. such as the Upland Plover (Bartramia longicauda (Bechst.

Thus. penetrate the soft tissues of the snail. Schistosoma haematobium. Tropicorbis. and. In the West Indies a genus of large snails known as Australorbis is parasitized by the Schistosoma mansoni. is also found in northern South America and many of the West Indian islands whither it was brought from Africa by slaves during the days of the slave traffic. This species originally parasitized small planorbids belonging to the genus now known as Afroplanorbis.General Ecology 21 THE PLANORBIDAE AS HOSTS FOR PARASITIC WORMS The family Planorbidae furnishes some of the best examjiles of the role of snails in the life history of certain trematode worms known as flukes. and possibly southern France. from the penetration of a single miracidium there may be developed as many as ten thousand or more cercariae. on reaching maturity. sary for the embryonated eggs to be swallowed by the snail. the Sudan. bore their way through the wall. on coming into water. The third species. Schistosoma japonicum. either by direct or indirect routes. Corsica. they have metamorphosed into the simple sacculate sporocysts. produce infection in human or reservoir hosts (from Faust. a progeny of larvae known as cercariae. Then later. and the Philippine Islands. Formosa. they utilize entirely different species of mollusks for the l)arthenogenetic stages. The small snails of the genus Drepanotrema have also been found with cercariae. 350). which has become a serious pest in these regions. These parasites require a mollusk as the host of the parthenogenetic generations of their life cycle. Eventually the larvae in the eggs become clothed with a ciliated epithelium. related to the African group Afroplanorbis. pp. in the course of half an hour or an hour. These miracidia are adapted to specific groups of snails. whereupon they hatch in the lumen of the oesophagus. In this life cycle the Schistosoma in the human host lays eggs which are voided in the faeces or the urine. Palestine. the egg shell bursts and the miracidium escapes. In the meantime. 1932. 350-351). Schistosoma mansoni. the Congo Basin. it is neces. they erupt from the tissues of the snail. Thus. Sicily. with a distribution in northern Africa. ^Mesopotamia. occurring in the same localities in Africa as S. When in their swimming they reach the proximity of certain snails they show a specific response to that species of mollusk. haematobium. group of worms known as blood flukes {Schistosoma) cause serious disturbances in man. with a distribution in China. and. A genus of smaller snails. In certain cases of other trematodes iClonorchis and Metagonimiis) however. Japan. These first generation sporocysts give rise to a parthogenetic progeny (either second generation sporocysts or rediae) From this second generation there develops (also parthogenetically). Three species are known from different parts of the A world. In most cases they readily attack. . probably due to a mucous secretion of the animal. whereupon it begins to swim about rapidly (Faust. Although the three human species of blood flukes (Schistosoma) are closely related. 1932. uses members of the genus Oncomelania. parasitizes species of the genera Bulinus and Physopsis. is also used as intermediate host. Attempts to use the genus Helisoma as a host for Schistosoma mansoni . and. ancl gradually migrate through lymph channels into the lymph spaces bathing the hepatic glands. p.

where the mollusk Aiistralorbis glabratus (Say) (=Planorbis guadeloupensis Sowb. Jones. A. E. P. and Hoffman. Lymnaea. 1934. Faust (1918) has observed cercariae of four species in Helisoma trivolvis in Illinois and one species in Helisoma sidjcrenatum (not trivolvis.. Elaborate studies have been made by Dr. and '*S. including many of the Planorbidae. p. 365424). true Segmentina not being found in the East. and Stagnicola) as well as one Physa. typographical error) from Louisiana to be hosts of many American cercariae. nitidella' which have not yet been examined for their anatomy and hence their position is doubtful (probably Polypylis) The value of correct specific identifications of the intermediate hosts of these trematode worms is of. Schistosoma haematobium is shown to use a species of planorbid. olivaceus. cubensis. pfeifferi. uses only the small planorbids belonging to the old genus Segmentina which now includes the groups Pohjpylis. Planorbis parvus (Gyraulus) and Planorbis trivolvis (Helisomci) as well as several Physidae and Lymnaeidae which are known hosts. antiguensis. However. Fourteen species of cercariae infest Helisoma trivolvis and three species occur in Gyraidus parvus. 22 The MoUuscan Family Planorhidae were unsuccessful. Fossaria. C. 1933. and Drepanotrema cultratus. Helicorbis. Hoffman of the School of Tropical Medicine of the University of Puerto Rico. Fasciolopsis buski. In a list of the described cercariae of the United States 1919) Faust mentions two species of the family Planorbidae. New Orleans. Aguayo (1938. summarizes the cercarial stages of parasitic flatworms. Papers by parasitologists in which molluscan hosts are mentioned are numerous. 1934.) is the intermediate host. as stated) in ^Montana (Faust 1917). . on various phases of the Schistosomiasis mansoni problem in Puerto Rico. this text was compiled at the very beginning of studies on trematode life histories on this continent. Faust of the Department of Tropical JVIedicinc in Tulane University. More recently there have been numerous de( . William A. A. although species of this genus are parasitized by other forms of trematode worms. 210) has summarized the species of snails used as intermediate hosts by Schistosoma and other trematodes. and by the late Dr. A. For . 'Planorbis dufuorii' as well as Bulinus and Physopsis. and Planorbis are listed. The liver flukes of the genus Fasciola parasitize certain genera of Lymnaeidae [Radix. The Schistosoma mansoni intermediate hosts include Ajroplanorhis boissyi. these studies see Faust. The large fluke of the Orient. Tropicorbis centimetralis. Pseudosuccinea. O'Roke (1917) found Helisoma trivolvis (probably the race lentum) carrying cercariae in Kansas.the first importance as has been clearly indicated by Ingles (1933) in a study of the parasites of frogs. in Ward and Whipple's Fresh-water Biology (pp. and McCormick (1923) found Helisoma trivolvis and Gyraulus parvus to be infested in Ohio. Australorbis glabratus. A few are here indicated. A. Planorbis coenosus { Helicorbis) Segmentina hemaesphaenda {=Polypylis) and \Seg)nentina sch)naken = . Four species are listed as intermediate hosts. which is closely related to Fasciola. Physa. Miller (1936) found Helisoma trivolvis from Illinois and Helisoma trivolvis lentum (not lantum. and Pingiella. Hoffman and Faust. sudanicus. gives keys for the determination of the larval stages and indicates the species of mollusks which have served as intermediate hosts. and Faust. Ward (1918).

albumen gland. Puerto Rico. Fall Lake. contained cercariae. Helisoma trivolvis lentum. Montana. Collected by Dr. Four out of six specimens with cercariae in different parts of the body. Sent bj' Mr. Poland. were observed to be infested. Cercariae were abundant. A. Idaho. Subfamily Planorbinae No -spocimens of this subfamily were found with cercariae or rediae. Heavily infested with stylet cercariae. Louis Co. British Columbia. C. A. Jankowski. Many infested with cercariae. Two-thirds of the specimens examined (about a dozen) were infested with cercariae. Helisoma corpulentum from Miles Bay. Collected by F. One out of five specimens with cercariae. mostly the li\'er. Austmlorbi. fourteen miles west of Hartford. Ontario. Sent by Dr.. from near Urbana. Canada. Helisoma hornii. collected by Henderson. Wisconsin. During the anatomical investigations made for this work careful note was made of the presence of larvae of trematode worms. A. north of Rexburg. Colorado. A. One specimen in sixteen contained cercariae. one contained cercariae. Uni^•ersity of Saskatchewan. Peoria. Sent by Dr. Helisoma pilsbryi from Chetek F. Cercariae in eight Lake. mostly in the liver. Connecticut. in several examples almost consuming the liver. near Gypsum. principally cercariae and a few rediae. D. Cahn. from Lake Kahnipiminanikok. In several specimens the cercariae were most numerous about the stomach. from Unionville. Subfamily Planorbulinae Menelus cooperi callioglyptus from Quatana. Heli^soma eorpulcntum multicostatum. St. Illinois. Hoffman. R. All specimens examined were infested. Some additional titles are listed in the bibliographies accomj)anying the papers mentioned in this chapter. Sent by Junius Henderson. Orcas Island. and Knife Lake. and ovotestis. Canada. Henderson and Hugo Rodeck. Clemens. Subfamily Segmentininae Scgmentina nitida from marshes in a meadow in Czneriakow. The following species. and St. From the Biological Boaixi of Canada. Ml 111 tux cooperi from small lakes on Mt. C. Puget Sound. One out of six with cercariae. Connecticut. British Columbia.stitution. Helisoma plexatum. 23 Many other works might he quoted but a sufficient number have been indicated to show the role played by the fresh. Con. Cahn. Received from Dr. Ontario. Collected by specimens with the liver and ovote. Baker. Subfamily Helisomatinae Helisomn anceps. Helisoma subcrenatum from Cottonwood Pass. Puerto Rico. Collected by F. Dale Foster. Meuetas dilutatus from L'nionville.mainly in the ovotestis which was almost destroj^ed. Hoffman. About one-third were affected by cercariae. J. Canada. Baker. H(lisoma ivhiteavesi from Kashabowie Lake. from Teton River. C..General Ecology tailed studies on the stages in fresh-water snails. Vancouver Island. Over a thousand specimens of mollusks were examined but only a few contained parasitic worms. S. R. from Paul Lake. Several hundred were observed in some siJecimens. Canada. in the life cycle of the trematode worms.stly affected. Canada. one out of six infected. Various degrees of infestation. Of fifteen specimens examined. Ontario. Cercariae in liver and ovotestis. W. particularly the Planorbidae. One specimen with lung ca\ity filled with rediae and one specimen with cercariae in liver and genitalia. Collected by Dr. Lake Nipissing. Barron Co. One out of four infested with glahratus from near San Juan. Lake of the Woods. Collected by Dr. Drepanutremn hoffmani from pond near Isabela. Collected by Dr. arranged under subfamilies. Baker..stis mo.water snails. collected by J. Joseph. Some specimens from a sluggish creek one mile west of Devon.. Helisoma campanulatum wisconsinense from Pirate Island. Minnesota. a suburb of Warsaw. . Sent by Dr. Kamloops District. Washington. Cahn. Rawson. Ontario. William A.

P. H.. Unfortunately the two genera Tropicorbis and Australorbis have been confused by this author and many species lumped under Australorbis. ( tode infections. Faust 1920. In many other cases. the cercariae finally leave the snail which recovers from the damage inflicted by the parasite. Wesenberg-Lund (1934) and ]\Iiriam Rothschild (1936) have recorded instances of gigantism in snails apparently attributable to trema. It is obvious that specific identifications must be accurately made if any value is to be attached to these forms as intermediate hosts of parasitic worms. 60 plates made page 213 et seq. New York. collected by Dr. LaRocque. Many species of cercariae remain to be discovered from the tissues of planorbid and other fresh-water mollusks. Twelve out of fourteen specimens were infested with cercariae. in which case. and stomach are affected and in large measure destroyed. The work of Dr. E. and Ontario did not contain larval worms. Lutz is good and the species have been carefully distinguished. Fifty per cent of specimens examined were affected by cercariae. many about the stomach. the work of Dr. In one case (Menetus cooperi) almost the whole snail was so filled with cercariae (several thousand) that the normal organization could not be distinguished. Vianna Martins (1938) of the Laboratorio do Instituto Biologico Ezequiel Dias. will be discussed and the species figured.* *The author's death came before he could for Part II are printed as an Appendix to finish Part II. often. the modern genera Australorbis and Tropicorbis being the principal groups discussed. to which they have little relation anatomically.. The cercariae may infest only the liver. In Part II of this work the recognized species. Missouri. the cercariae being packed in the space occupied by the organs and in aggregation taking their form. Same species from small lake near Hutchins. Canadian National Museum. Lutz. The effect of fluke infestation on the anatomy of the snail is variable.24 The MoUuscan Family Planorhidae Menettis sampsoni from slough of Meramec River east of Stanton.E. Specimens have been examined in which the genitalia were completely destroyed or the genitalia and liver obliterated. this work (see However. Nearly all specimens examined were infested with cercariae. In several specimens nearly all of the internal organs had been destroyed. Collected by Leslie Hubricht. Promenetus exacuous from small lake in Wainwright Park. based in part on the earlier work of Dr. of species of both Australorbis and Tropicorbis. but mostly confined to the liver and genitalia. 79) has given a good account of the pathological changes in the gastropod liver due to fluke infestation. as well as of other groups. Lutz describes all species under the common generic name Planorbis. It is unfortunate that parasitological assistance was not available when these anatomical studies were in progress in order that the particular species of parasitic trematode worms involved might have been ascertained. Franklin Co. some in the fore part of the bod}'. Wisconsin. This was particularly true of the genitalia. Texas. and also the synonyms.). p. Dallas Co. Alberta. Ottawa. Martins is a study of the genus Australoi'bis of Pilsbry. Cheatum. Michigan. Received from A.C. Among these are the late Dr. Dr. Canada. In the main. the liver and genitalia being principally affected. Adolpho Lutz (1918) of the Instituto Oswaldo Cruz and Dr. Specimens of exacuous from Maine. genitalia. Several parasitologists have essayed the role of conchologists in an endeavor to understand the specific limitations of the groups of molluscan species acting as hosts for i^arasitic trematode worms. the liver. however. A. .

369-458). made several times a day.— H. . DEVELOPMENT OF Helisoma scalare (JAY) In 1932. 67-108) has described and figured the early stages of Lijmnaca stagnalis appressa =jugi(laris) and Lankester (1874) has given us the full development of the European Lymnaea stagnalis. a very significant experimental study. Roney. . pp.J. appeared in Ecology. on the embryonic development of the different species of the family Planorbidae and especially on their nidification. B. Development was observed to be sporadic in many of the eggs. E. when twenty-eight fertile eggs were counted. sent the writer a piece of pond lily leaf upon which was a large egg capsvde of Helisoma (Seminolina) scalare (Jay). under controlled conditions. 20 eggs in gastrula stage . 4.: III.* A dozen or more aciuaria were operated during the years 1930 to 1934 and careful data were gathered concerning the period of egg deposition. 25 . period of development. pp. on the development of Helisoma eggs. Andrews. 24 eg^cs in segmentation stages.v H. Little work has been done. b. . Observations began when the specimen was received on March 2 and the last specimen was hatched on March 17. v. 4 in early trochophore stage. and becoming marked from the early trochophore stage. NIDIFICATION AND EMBRYOLOGICAL DEVELOPMENT of cell growth and development in the phinorbids {Hclisonia trivolvis) have been well described and figured bv Holmes (1900. The observations. number of capsules laid. 5. of the Johns Hopkins University. and the time element in the growth of the voung animal in ( THE EARLY STAGES its shell. Crabb (1927. Florida. showed the following degrees of development March 2-3. *After this manuscript was in press. 24. A. V-C. 5 in trochophore stage 3 in veliger stage 2 postveliger. the rotation beginning in this species the latter j^art of the gastrula stage. More recently (1934) Lowrance has very fully described the early development of Stagjiicola kingi. Dr. 18 in gastrula stage . pp. containing twenty-eight eggs in various stages of development. A. From this capsule a fairly good idea of the development of this planorbid was obtained. The different stages are figured on plate 51 and are described in the explanation of that plate. 7. 4 trochophore 3 ^'eliger 3 embryos with shell. This egg capsule was observed as opi^iortunity afforded until all of the embryos were hatched or had died. however. 4 in trochophore stage 4 in preveliger stage. Same condition as on March 3. changes taking place within the egg capsule. Observations on the egg capsule began on March 2. . There were resting stages between periods of great activity of rotation about the interior of the egg. This rotation is caused by the vibration of the numerous cilia attached to the foot of the embryo (CL in figures). the second day of development. 18 in gastrula stage 8. . This capsule was deposited in an aquarium but the parent came from Lake Butler. . and growth of the young snails within the egg and after hatching. 218-243.

The radula can be seen working with its peculiar lapping motion. 9. 3 postvehger. 15. Several adult Helisoma scalare received from Dr. Trochoiihore stage. Both of the above embryos completed their development in fifteen days. 12. 7. Like fig. . 3. . Like fig. 13. Veliger stage. 12. The eyes were black and very conspicuous. Like fig. 12. Young snail. 14. 18 in 12. Like Veliger stage. Embrvo stage. Probably some early cell divisions took place the day before observations began. 9 on plate 51 the snail is very active. 7. . 3 embryos with shelL 18 in gastrula stage. 1 Segmentation and gastrula. fig. 3 vehger. 3 veliger. Like fig. 6. 14. The temperature of the room was 70-75 degrees Fahrenheit. 3 postvehger. Like Veliger stage. 13. 11. Embryo stage. In the stage shown in fig. 11. Posttrochophore stage. Young snail. 18 in gastrula stage 2 postvehger 3 embryos with shell 1 hatched. Embryo stage. The egg capsules were deposited on the glass sides of the aquarium and were numbered so that development could be easily and accurately observed. collected in Lake Butler. In the above table it will be observed that only ten embryos completed their development. which would increase the development period to sixteen days. Hatched from egg. Florida. Like fig. . were placed in an aquarium where they bred and laid eggs which successfully completed their development. Thirteen egg capsules were deposited early in Alarch: . 12. 16. 7. many of the gastrula stage filled with Protozoa. . 18 in gastrula stage. Eighteen embryos did not continue their growth after reaching the gastrula stage. Young snail. 6. 13. . Hatched. Like 7. Andrews. Like fig. 3 embryos with shelh 18 in gastrula stage. Like figs. The other eggs in this capsule showed approximately the same development. 18 in gastrula stage 11. Like fig. for what reason is not apparent. 10. 5. Embryo stage. 8. One young snail four days out of the egg was transferred to an aquarium. 13. for it can be seen to emit bubbles. Early trochophore stage. Like figs. 5. Veliger stage. 9. 2. 8. March Veliger stage. Like figs. Veliger stage. Like fig. crawling about the glass sides of the jar. 10. 6. 1). Like fig. 8. 3 embryos with shell 1 hatched. three days later. All hatched or dead. Like figs. 14. The embryo in egg number 5 migrated to egg number 6 at the late embryo stage and both embryos occupied the same chamber until they were hatched. Embryo stage. . Early segmentation and ga. Young snail. . Like figs. 9. 15. 14. figs. Young snail. 13. 2 embryos with shell. 5. Like fig. Like fig. 11. Like fig. moving about in the egg and grazing like the adult snail. Several of the embryos that completed their development showed the following history. 10. 4. 8 10. fig. . . 3 veliger.26 March The MoUuscan Family Planorhidae 10. Like fig. EGG NUMBER March 2-3. 4 postvehger. 10. Like fig. fig. 13. 12. 15. EGG NUMBER March 2-4. 9. March 10. Eggs were first observed on Alarch 13 (see plate 52. 3. 5. . Like fig. 12. 17. Po-sttrochophore stage. Young snail. This specimen was very active. 1 hatched. 4. 13.strula stages. and the pulmonary siphon is functioning. 18 in gastrula stage 2 veliger 2 postvehger 2 embryos with shell 2 hatched. 2. 16. . 16. the heart is beating regularly. 4. 4 trochophore 3 veliger 3 embryos with shelL gastrula stage 4 vehger 3 postvehger. .

Capsule X"o. 2. 12. 13 14 eggs.0 by 3. mm. There were three adult specimens measuring 8 to 10 mm. Trochophore and preveliger stages. from ]March 15 to ]\Iarch 31.0 mm. 10. cap. capsule 4. One capsule with nine eggs developed but twelve days April 2. 23. Gastrula stage. These.5by2. Veliger stage. 12. developed and hatched the young snails in thirteen days. 10. 4.: : Xidification Xo. 11. as follows.5 mm. figs. All other capsules occupied fifteen to eighteen days. capsule 4. 21. Freshly deposited eggs.0 by 0. the process occupying Post^•eliger stages. The period hatching. 28. 6. capsule 3.0 by 3. 8. Early segmentation.0 by 3.small shell. 17. Last embryo hatched.5. The shells were very wide with broad spire and decided umbilicus. shell almost 18. with ten eggs. capsule 3. Freshly deposited capsule. 11. capsule 5.5 by 2. Young snails all hatched. Gastrula and trochophore stages.0 by 3.0 mm.0 diameter 2.0 mm. 2 eggs. lOeggs. 7.5 mm. from ^Nlarch 15 to ^Nlarch 27.5 by 3. Four efnbrv'os hatched. 11. 1. Segmentation. 3.0 by 3. Embryo filling egg. March 22. 6 eggs.7 mm. About two weeks later (iVpril 2) six capsules were deposited measuring as follows (see plate 52. with 14 eggs. curiously enough.0 mm. The young snails in the aquarium were approximately seven and one-half weeks old.0. Capsule 5.5 by 3. was fifteen days. 2. 26.5 by 2. 2. capsule 3. with 10 eggs. 10 eggs.0 mm. and Emhryological Development Xo. 5. 4. 6. with 19 eggs. diameter 2.0.5 by 3. with five eggs. March 13.0 mm. 11 eggs. 8 eggs. April 8. 9. 8.0 by 3. had all embryos hatched in seventeen days.'*ule3. 9 eggs.0 mm. and earlj' embrj'O Segmentation and gastrula stages.5 by 3. 21/221/4- . capsule 5. 13 eggs. in length. There were ninety-six young snails of scalare only sixteen of which were living. were in shape more like Helisoma duryi seminole than typical scalare. Embryo with . with 7 eggs. mm. 13. Embryo with shell filling egg. Capsule Xo. 9. Embryo stage. Gastrula and trochophore stages.0 mm.5 mm.5 diameter 3. 20.5 mm. mm.8 diameter 2. 19. Embryo with filling egg. On ^May 22 the aquarium was examined for its entire contents. 7. capsule 4. 2^>. 27 2. Egg capsule No. Early embryo stage.5 by 3. capsule 4. 16.0 by 4. capsule 5. Preveliger and veliger stages. 24.0 mm. 10 eggs. A few of these were measured and the number of whorls was counted: Height Height Height Height 2. Embryo with whorl. 10 eggs. filling egg. 5 eggs.0 by 3.5 by 2. shell of about one filling Embryo egg.5 mm. capsule 5. with shell half Embryo with fourths shell three- Gastrula and trochophore stages. Capsule 5.5 mm. capsule 5. 25. 3) Capsule 4.0 by 4. The shell when the young are hatched measures about 1.0 mm.5 mm. 3 with eight eggs developed as follows: 14. with eleven eggs. Capsule 5. 3. with 9 eggs.5. of development 6. Whorls Whorls Whorls Whorls 2.5 mm. Embryo filling egg.8 mm. Posttrochojihore and preveliger stages. Capsule 6. 4 eggs. Embryos hatching.5 mm. Capsule 5. 5.0 by 3. with 9 eggs. occupied a longer period (18 daj's) in Capsule Xo. 27. 3. 15.

mm. 1932. On September 3.7 mm. diameter and have a trifle more than one whorl. leaving one adult.0. 9. differences in the form of the embryos as would be expected in two such diverse families. The young scalare. it will be noted. and 1933. Embryos in but two of the capsules completed their development. 10. 11.0. Between September 3 and September 17 these young snails formed eighteen in egg capsules containing the following number of eggs: 6. Wliorls 3^2. The young snails lived until January. four of the young scalare were living. 4. 12. All li\'ed from March 5 to May 21 when three died. Whorls 3.5 mm. The parent snails of this last brood. 11.0 6. 6. 2. after a hot summer. 9. Baker .0 4. Five of these measured as follows: in Height Height Height Height Height 4. These measurements indicate that the shells had almost trebled in size about twelve weeks. although but one more whorl was added. C. mm. B. mm. in diameter and had three and one-half whorls. in height and 4. 1933. The young shells measured 2 mm. 10. NIDIFICATION IN AND DEVELOPMENT OTHER SPECIES OF Helisoma Plates 51 and 52 Living material belonging to five additional species and races of Helisoma was studied in aquaria during the years 1931. during a hot period of weather.0. of course. On June 26.5 mm. 13.7. in height and 0. Whorls 3. were the offspring of an isolated adult scalar^e. seven specimens remained in the aquarium.0.1 5. The species included were: Helisoma trivolvis lentum (Say) Helisomn trivolvis fallax (Haldeman) Helisoma duryi normale Pilsbry Helisoma duryi eudiscus Pil. 7. Two empty egg capsules were observed on the side of the aquarium and twenty-one young snails were crawling about.sbry Helisoma tenue calijorniense F.28 The Molluscan Family Planorhidae When hatched the shells usually measure 1 mm. 1934. 5. 9. 5. On jMarch 5. Whorls 3%. 5. 10) but that with increased temperature (26 degrees centigrade) this period was cut to from nine to eleven days. 1933.0 mm. 6. There are. diameter diameter diameter diameter diameter 5. On August 15. On June 25 the last adult died but nine of the young remained alive. Liimnaea stagnalis may require as much as twenty days or more for development. Lowrance found that young snails of Stagnicola kingi hatched in from fourteen to sixteen days (p. two adult snails of this species died.1 5. in diameter and had two whorls. about twenty-three weeks old (160 days) measured 5. which lived over winter isolated in a small aciuarium. 7. several egg capsules were laid by this individual approximately nine months after its isolation. had more than doubled in size in less than eight weeks. 7. therefore. The development of HeUso)}ia scalare is similar in many respects to that described by Lankester for Lymnaea stagnalis (1874) and by Lowrance (1934) for Stagnicola kingi. The young snails at this time. Whorls 3%. 7. in height and 2.0 mm. 10. mm.

173. [a] Self-fertilization or Autofecundation or autofecundation is possible among the freshwater pulmonates has been suggested by several zoologists during the past hundred years or more. 1902. 1918. On pages 541-543 he ]iresented a list which includes all of the authentic exami)les of autofecundation. that reproduction by parthenogenesis does occur in some groups of snails. Notes were kept of developments in each capsule.Nidification and Eutbnjological Develojwient 29 Several species were from different localities and these as well as each race were kei)t in separate aciuaria. and several groups of Planorbidae {Gyraulus. Menetus) lay fertile eggs by this method when isolated. p. producing twenty-seven young snails. The reproductive method in Campeloma was observed by morphological and histological studies and there is no question concerning the parthenogenetic method of reproduction in the species examined. \). Boycott (1919) has asserted that Paludestrina jenkinsi develops by this method. Some years ago. Each jar was labeled and the egg capsules lettered or numbered as they were deposited by the animals. 1902. a small quantity of weed was introduced. the snail deposited some eggs which were hatched in early July. many of which were still thriving under these confined conditions on December 10. His conclusions were that there was no evidence of parthenogenesis in this snail. or until August. and many others along this line. Colton (1912. Observations were made several times each day at intervals of three or four hours. Crabb (1927) studied Lymnaea stagnalis appressa {=jugularis) to determine whether snails kept in strict isolation reproduced by self-fertilization or b}' parthenogenesis. The data which follow have been condensed from the original notes. 1938) have shown that the viviparoid snail Campeloma rufutn develops solely by parthenogenesis although other species of the genus have been found to be bisexual. In evaluating this set of observations.v. Physa. ]More recently Van Cleave 1937) and ]\Iattox (1937. 265) conducted some interesting exi)eriments on Planorbis vorte. and more recently Larambergue (1939) has issued an elaborate monograph particularly relating to Bidimis contortus. but until comparatively recent time few experiments have been carried on to test the reality of this method of reproduction. the rest of the jar was filled with pure oxygen and the vessel was perfectly sealed. aften sixteen months of complete isolation. 48) has shown that Lymnaea. it should be remarked that only those experiments in which individual eggs were isolated have any conclusive value in establishing the occurrence of self-fertilization. p. Crabb (1927) has presented elaborate data on this subject. however. Chadwick 1903. In June. 1901. On February 2. This list includes all families of Basommatophora as well as a few among the Stylommatophora. 1902. That self-fertilization ( ( . The information sought in these studies was to ascertain the time required for development and the rate of growth of the shells of the young snails during their life in the aquaria. It is to be noted. a common English planorbid. he isolated an adult specimen in a jar three-fourths filled with boiled water. It remained in this condition for eighteen months. This is especially true when forms are considered for which there are no observations as to the length of time the spermatozoa remain viable following copulation.

1933. With this general statement the writer can not fully concur. The writer has observed many species of Lynniaea. 1931. 1932. and the planorbids in isolated laboratory stock lays about e\-ery third day for a period of about twentydays. 58. On June 20. 1932. In early April. why should the male portion of the genitalia be so elaborately developed with prostate and the penial complex provided with such diverse accessories in its makeup? The spermatheca has been found to contain a spermatophore containing sperm which was obviously deposited during copulation. 175). laid twenty egg capsules aggregating 325 eggs. on the contrary. An isolated individual which lived over winter (nine months) without egg laying. No other eggs were laid by these specimens which lived until Se]it ember 17. In the family Planorbidae the intervals are much longer.30 The Molluscan Family Planorhidae Crabb states (1927. Thirteen isolated individuals had the following interval between hatching and egg laying: 32. 1933. ]i. the parent snail was removed to another aquarium and on June 22 laid two egg cap- ary . was isolated Janu16. It was about 325 days old. this individual. formed no capsules. not only of the specimens placed in isolation. Specimens of Helisoma trivolvis lentum were placed in aquaria in 1930. However. A young specimen from the parent lentum. 56. rests a few days. at least in those species having a penial complex of sufficient length to reach the female opening. 63. If the method of reproduction mentioned by Crabb is the chief or possibly the only method. Physa. after inhabiting the aquarium for 132 days. It is doubtless true in many cases of reproduction during isolation but that it is the chief method among free snails when in aggregation in their native environment or even in the laboratory is scarcely possible. 92. Helisoma scalare laid thirteen egg capsules in early March. 91) that self-fertilization is the normal method of rei^roduction in Lynmaea and other fresli-water pulmonates and that cross-fertilization seldom or never occurs. 50. . 50. 1932. five My Colton's experiments on Lynmaea {Pseudosuccinea) columella indicated that the interval between hatching and egg laying was widely variable (1912. Of fifty-four L. or a period of 943 days. Even the control culture. but for those in aggregation. deposited two fertile egg capsules on March 5. This is continued throughout the year. Twenty-six days was the shortest interval and ninety-two days the longest interval. and 74 days. then begins another cycle of laying. consisting of two snails. Between May 27 and June 9. was very prolific. 35. 49. Crabb states that he was unable to raise progeny from isolated Planorbis trivolvis iHelisoma) four individuals being carefully reared in isolation for 377 days without the laying of a single egg capsule. 73). 26.appressa isolated from the same egg mass. a capsule was laid. p. Self-copulation probably occurs when a snail is isolated. on February 22 four capsules were deposited and on March 7 two capsules were laid. Ten to fifteen small empty egg masses were deposited. p.s. seven laid their first eggs fifty-eight days after hatching. 74. six additional capsules were produced. they do not reach their maximum laving capacity until they are about one hundred days old (Crabb. Between September 3 and September 17 the young snails. Lymnaca stagnalis. 57. coitu. 1932. which is the case in many species of the Planorbidae. On February 16. 160 days old. laid eighteen egg capsules containing from six to thirteen eggs in each capsule.

Of the original 125 eggs laid in February and March. 1933. W. with twenty-four eggs. On October 23 and December 26 the six specimens were still alive. No. This city is also Say's original locality for this species. A few of the second generation remained alive for about two years but laid no laid fifteen egg capsules.Nidification siiles and E tnbryological Developinent 31 and on June 27 deposited two more capsules. the smallest being 5. [1] Helisoma trivolvis lentum (Say) SoNoral . with foiu' and one-fourth whorls. 1932. nineteen specimens were counted ranging in size from 5. It measured 7.5 mm. 1931.spcfimoiis of the Ic ntiim race were received in 1930 from Dr. in diameter with four whorls. two specimens were living. On June 25.J. Two-thirds of the eggs laid completed their development. 1931. Pap. 1932.5 by 6. one with fifteen eggs and the other with nineteen eggs. one 9 by 4. in diameter with three whorls and the largest 14 mm. only fourteen specimens remained in the aquarium. On November 28. an egg capsule was observed. with four whorls. ranging in diameter from 4. All eggs were fertile and contained embryos in the early stages of development.5 mm. with four and one-fourth whorls. The snails were placed in aquaria and kept under observation for several years. On February 16. the smallest being 6 mm. only 25 young snails survived until September. H. in diameter and the largest 10 mm. three were li^ing.5 mm. the nineteen specimens were again measured. This has been done by Holmes (1900) and others. one 4 by 3. the same number of specimens remained in the aquarium.0 mm. On September 17. Their size had not increased. with four and one-fourth whorls. These capsules were and twenty days after the large laying period in the first aquarium. New Orleans. apparently freshly laid. 1931. with four whorls. Zool. August 15. 1932. Uni. On March 19. in diameter and had three and one-foiu'th whorls and the largest was 12. 226). in diameter with four and one-foin-th whorls. fi-\"c ha\ing died. the largest 10. Faust of the Tulane Medical School.5 mm. and one 10 by 4. These two specimens had li^•ed o\er two years in the aquarium. On September 12. The smallest was 6. the largest being 11 mm. and onl.5 mm. E. Mus. *Rearing methods for securing uninfected snails to be used in life history studies on parasitic worms have been practiced in many parasitological laboratories. Gyraulus. and Tropicorbis would react to such investigations is not known. in diameter. with two and one-foui'th whorls to 16 mm.5 mm. On June 10. Krull (1931) has published the jjrocedure for successful rearing of Gyrauliis parvus and found that this species may be reared to full size in from four to five weeks under laboratory conditions (Occas. On January 16. with nineteen eggs. 1931. with nine eggs. C. 1931. They were collected from the canal at New Orleans. Just how the smaller species of the genera Menetus. twenty-five young snails were counted. in diameter with three whorls to 14 mm.. five specimens were living. with twenty-five eggs. .y two snails were alive two and one-half years later. the smallest 6. On February 22 four additional egg capsules were laid. two other egg capsules were observed. only six specimens were alive.5 mm. No attempt has been made to go into the finer details of embryological development. 1933.* It is probable that had Crabb kept his Helisoiita trivolvis longer they might ultimately have laid eggs. Mortality among the young snails appears to be heavy in their earlj' postembryonic life. On March 7. one 7 by 4 mm. Michigan. with fourteen eggs. [b] The Development of Helisoma The notes that follow describe some of the results of aquarium observations on a few of the species of the genus Helisoma. in diameter with three and one-fourth whorls and the largest 14. These examples seem to indicate that the large planorbids do not breed as often as do the lynmaeids and are not as available for studies in development as are members of the Lymnaeidae.— H.5 mm. 1932.5 mm.V-C. On March 5.5 mm.5 mm. 1933.

from normal habitats. and containing six eggs. Tlie eggs of both capsules passed through their development and were hatched by February 12. in diameter with 3M whorls. in diameter with 2^/4 whorls. but was eaten by Protozoa after three days of development.8 mm. in diameter with 2^2 whorls. 1 specimen 5 mm. the seventeen young fallnx were still alive.5 mm. 1932. in diameter and had three and one-half whorls.. in diameter with Sy^ whorls. 1 specimen 2. This small subspecies of HeUsoma trivolvis lived in the aquarium 495 days or about seventeen months. The li\ing young snails measured as follows: 2 specimens 2. 1 specimen 5. and increased from one and one-fourth to three and one-half whorls. Massachusetts. only four young jaUax remained living: 1 specimen 3. the young ranging in size from 1. On August 15 there were fourteen living young snails which measured as follows: 1 specimen 2 mm. one measuring 4.5 mm. The last remaining young was. An examination on May 22 revealed seventeen young snails living.5 mm. fifteen specimens of tliis race. in diameter with 2i/-> whorls. are only tentative and are possibly subject to considerable error. in diameter when hatched to 6 mm. Full maturity would i^robably have been attained the third year after hatching. 1 specimen 3. containing ten eggs.2 mm. Rehder. On June 26. and one 5. in diameter with 3 whorls. 1 specimen 3 mm. in diameter with 3 whorls. therefore. March 4 a capsule was laid measuring 5 by 6 mm. 1 specimen 5 mm. Adult fallax measures 15 mm. On February 28 another capsule was laid measuring 5. Harold A. Conclusive evidence of rate of growth in natvu-e could be determined by measuring representative samples. All of the adult snails were dead. to 3 mm. On March 1. 1932. 1931.5 mm. On June 25 one young snail remained which measured 6 mm. in diameter with 3 whorls. It grew from a size of 1. 1932. in diameter with 2 whoils. taken periodically as practiced by Van Cleave (1932) and his students in the interpretation of life history data. in diameter with 2% whorls. 2 specimens 5 mm. were recei\'ed from Dr. 3 specimens 3 mm. in diameter with 3 whorls. in diameter with 3 whorls. 1 specimen 5 mm. and on Maich 10 another capsule was laid.5 by 3.5 mm. in diameter with 3^/4 whorls.8 mm. these four snails were still living and measured as follows: 1 specimen 4 mm. in diameter with 3 whorls.5 mm. in diameter with 3 whorls. in diameter and has five full whorls. in diameter with 2i/{> whorls. 1 specimen 4. in diameter with 2 whorls. United States National Museum). 2 specimens 4 mm. only three living snails remained: 1 specimen 4. in diameter with 2 whorls. in diameter with 3 whorls. 4 specimens 3. twenty-two young and five adult snails were counted in the aquarium. 1 specimen 3. 1933. in diameter with 2^/^ whorls.5 nun..32 [2] The Molluscan Family Planorbidae Helisovna trivolvis fallax (Haldeman) October 2.5 mm.2 by 4 mm. in diameter with 31A whorls.5 mm. On October 23. in diameter with SMi whorls. then in Harvard University (now x\ssistant Curator of Mollusca. 3. 1 specimen 5. On March 13 a capsule containing nine eggs was observed. with twelve eggs.6 mm. with thirteen eggs. . somewhat more than one-third grown. in diameter with 2V2 whorls. in diameter with 3^4 whorls. On May 1. 2 specimens 4 mm. in diameter. 1 specimen 3. Very early experiments on snail development tended to show that the size of reared snails is influenced by the volume of the water and the container in which they grew.8 mm. 5 specimens 4. in which length of life span is estimated from growth rates in aquaria. in diameter with 3 whorls.6 by 7 mm.5 mm. containing eleven eggs. 1933. collectecl near Cambridge. when it died. 2 specimens 5 mm.9 nun.5 mm. Conclusions of this sort.5 mm. 1 specimen 5. 3 specimens 2. in diameter with 2^2 whorls. On March 19. 1 specimen 5. Egg capsules were observed February 6.2 mm.5 by 5.

the snails had left the egg capsule and were feeding on the side of the aquarium. By February 6 most of the veliger larvae had been hatched. forty miles west of Miami. one containing fifteen embryos in various stages of development.2 mm. They were placed in aciuaria in January. one gastrula stage and thirteen embryos with well-formed shells. Another capsule contained fifteen eggs with one veliger stage. in diameter with 2 whorls. in area) and one with four eggs (4. in diameter with 3 whorls. 1 specimen 5. the liver brownish. specimen 5 mm. one capsule carried out its full development. Normandy High School. two died and were eaten by Protozoa and one completed its development. F. On December 26. one with ten eggs. B. On May 22 there were seven living snails and thirty-three empty shells. On September 1. 12). 7 by 4. On September 21. one with seven eggs (6 by 3. in diameter with 3^4 whorls. On September 12. By August 15 one had died. the walls of the egg capsules were broken. in diameter with 3^/i> whorls. and the snails were crawling about in the broken interior. three capsules were observed on the side of the aquarium.Nidificatio)} (Did Eiiibnjological [3] Development 33 HeXisoma duryi noymalt Pilsbry Specimens of this planorbid. but four snails remained alive. The tentacles are long and mobile. The radula can be plainly seen at work and the jaw appears blackish through the translucent head. 1931. 1932.7 mm. 1 specimen 2. Of the first series of three capsules. 1 specimen 2.5 mm. From the aboA'e capsule and from several others hatched at about the same time. On June 26 the seven snails were still alive. The six snails gave the following measurements: 2 specimens 4 mm. Of the second set.5 mm. . Florida. five embryos with partly formed shells and fi\e embryos with fully formed shells. indicating that mortality had been heavy. collected on the Tamianii Trail. three in veliger stage and twelve with wellformed shells. and the eyes black and prominent.5 mm. in diameter with 2^-2 whorls.1 mm. and actively glided about the aquarium. Shay of the Department of Science. the veliger stage was reached on September 15 and the embryo with a well-formed shell on September 19.y February 2. in diameter with 3'!4 whorls. in diameter with 3 whorls. a recently laid capsule containing sixteen eggs was discovered and was obser\ed until all of the embryos had been hatched (see plate 42. two additional capsules were deposited. St. in diameter with 3 whorls.8 mm. In this capsule.8 mm.5 mm. and one with eight eggs. in diameter with 3Vi! whorls. 1 specimen 5. 1 specimen 4 mm. The body in the shell is pinkish. 1 specimen 3. 1 specimen 5. W. 1 specimen 5 mm.5 mm. i On March 12. 2 specimens 3.7 by 2. Observations were made on the shells several times in March and April. 1 specimen 6 mm. These latter hatched on February 4. It is probable that this capsule had been laid for a day or two before first observed because the interval of nine or ten days seems too short for comjilete development. In this cajisule fifteen days elapsed between early segmentation and exit from the egg. forty young snails were released in February. 1 s])eciinen 6.4 mm. 1932. These were hatched by February 5. These showed the measvu'ements below: i. one with nine eggs. in diameter with 2->4 whorls. Another capsule contained ten eggs.7 by 3. On January 31 several egg capsules were observed. the foot hyaline.). 1931. in diameter with ZYi whorls. 1 . principally during the month of May. the four snails were actively feeding in the egg capsule. 6 by 5 mm. On January 17.sured as follows: 1 specimen 4.7 mm. By the middle of February the young snails measiu'ed 1.1 by 1 mm. in diameter with 3% whorls. On September 20. The seven living snails had the following shell measurements: 1 specimen 1. in diameter with 3^/4 whorls. three snails were living which mea. m diameter with 3 whorls. Louis.2 mm. 5. 1933. vu o u 1 specimen 4 mm. 2 specimens 4. • • . slightly yellowish.2 mm. in diameter with 3Vi whorls. were sent to the writer by Mr. eleven snails had been hatched and were crawling on the glass sides of the aquarium. in area. in diameter with 3^/4 whorls.5 mm. in diameter with 3 whorls. fig.

Van Hyning. with nine eggs. in diameter with 3^-2 whoi-ls.5 mm. 16.5 mm. in diameter containing 18 eggs. Specimens of cali^oyr\. 5 by 8 mm. whorls. in diameter containing 15 eggs. 12) completed its development. Another was disintegrated and the space was filled by Protozoa. membrane. 6 by 3. Early embryo stages. If this growth is normal. 7. [5] HeXisotna tenue. 21. Segmentation begun. Marion County. San Bernardino Mts.5 mm. egg laying began again with the deposition of a capsule measuring 4 b. They were placed in an aquarium on February 12. [4} HeXisoma duryi ^udiscus Pilsbry Si>ccimens of this race of duryi were received from Mr. in area with five eggs. 1931. and on February 12 a capsule with seven eggs. 25. On February 10 a large capsule measuring 5 by 4 mm. 2. 4 by 7. 24. in diameter containing 21 eggs. All . in diameter with 3% whorls. from February 13 until February 28. 14. 22. stages. Embryos with shell forming. 10) developed to the gastrula stage and then ceased growth. 27. 1933. On February 6. Embryos with shell ^3 filling 15. One capsule (Feb.1 mm. 4 by 4 mm. but two snails remained alive. Embiyos with shell completely filling membrane. 8. in diameter containing 38 eggs. Feb. of the Florida State Museum. on the green algal deposit which formed on the glass sides of the aquaria. in diameter. From June 14 to June 26 the eggs remained in the same stage and at the last date the eggs appeared to disintegrate.8 mm. was deposited containing twelve eggs. Early embryo stages. in diameter containing 27 eggs. to 6. this race % % 19. 1933. in diameter with 3^/i> whorls. Segmentation ancl gastrula stages. 28.. T. collected at Silver Springs. Gastrula and trochophore stages. Embryos with shell filling 17. 4. S.5 mm. so that the young snails were about one-thiixl grown after a development period of one year and seven months. 4 by 4 mm. 9. Development was fairl.i&ns& F.5 mm. near Redlands. hatched. Embryos with shell forming. A typical development is indicated below: Feb. 4 by 5 nun. On August 6. This temperature reached 90 degrees or more in the hot . each egg 1 mm. Berry.5 mm.34 On June The Molluscan Family Plaiwrbidae 25. in diameter with S^/v. Embiyos with shell filling 18. membrane. C. 1. The matter of food is also important since only filamentous algae were jilaccd in the aquaria. 119 days after the laying of the last egg capsules. 1 specimen 6. 4. The tem])erature in which the young snails were reared was ordinary room heat between 70 and 75 degrees F. in diameter and has five whorls. in diameter containing 11 eggs.v 3. S. On December 13 an egg capsule was laid measuring 2 by 2. in diameter containing 26 eggs.v regular and occupied fifteen days. of 26. the snails feeding mostly. A single adult cudisciLS was left in the aquarium and on June 11. all of the embryos being hatched. 23.summer months. They were received at the laboratory on October 2. and laid nine egg capsules on the next day. it must require three or four years for this race to attain maturity. 1 specimen 6. 5. in diameter with 3^2 whorls. 4 by 7 mm. the 1 same three snails showed some growth: specimen 5. Trochophore and preveliger and y^ of membrane. collected from Prospect Park. California.8 mm. 5 by 3. 1 specimen 6.5 by 7 mm. another capsule was laid measuring 2 by 4 nmi. and containing two eggs. 1932. Baker were recei\ed from Dr. Veliger stages. 6. 3. These 3^oung snails li^•ed 570 to 580 days after hatching and increased in diameter from 1. A third capsule (Feb. and in number of whorls from one to three and a half. These measured: 1 specimen 6 mm. unless there is considerable acceleration in growth during the second or third years.5 mm. The nine capsules contained 199 eggs as noted below: No. On Jime 13 the eggs had reached the gastrula stage. 20. howe\er. in diameter containing 15 eggs. in diameter containing 28 eggs. Early veliger stages. 1932. An adult nor male measures about 20 mm.5 mm.

in diameter with 3 whorls. All of these eggs completed their development in from twelve to fourteen da.0 mm. On June 5. but two specimens remamed alive. On May 1.2 mm. 1933. 3 specimens 3. On . in diameter with 3 whorls. to 6. four additional snails died lea\ing eight young which gave the following measurements 1 specimen 3. At this ratio full maturity would be attained between the third and fourth years of life. 1 specimen 5. Between June 5 and June 26. 2 specimens 2. in size and containing eighteen and twenty eggs. 1933. [6] Other Records of Egg Laying HcUfiODUi pilshryi infracnrinatiun F.suring 6. As in the cases of the other species described in this chapter. 1933. 6 specimens 3. mea.1 mm. 1933. and 6 by 4. in diameter with 2V-> whorls.0 mm. 1932. There were ninety-six li\ing young snails and forty-fi\e empty shells. in diameter with 2V2 whorls. in diameter with 2 whorls.5 mm. 1 specimen 4. 1 specimen 5. in diameter with 2 whorls. 2 specimens 4. 1932. in diameter with 3^/4 whorls. of the Canadian National Mu.0 mm. in diameter and has five whorls which would indicate that the young specimen sixteen months old had made about one-third of its growth.0 mm. On June 25. in diameter with 3 whorls. On March 12. in diameter with 2 whorls. food.5 mm. Sjiecimens of this species were received from Mr.5 mm.0 mm. The living 3'oung snails gave the following measurements: 8 specimens 1. They were collected in the Rideau River near Ottawa.5 mm. They contained three. in diameter with 314 whorls. only fourteen young snails were alive. Canada. 3 specimens 4. and temperature maj^ have retarded the normal growth of the young of this race. 40 specimens 2. in diameter witli 2 whoi'ls. two young snails died leaving twelve active 3'oung.sules were deposited each measuring 6 by 4. and the whorls increased from one to three and one-half. 9 b\' 6. in diameter with 21^ whorls. Adult californiense measures 20 nmi.1 mm.0 mm. eleven.0 mm. 1 specimen 4. in diameter with 2y-2 whorls.snails. 3 specimens 2. and twenty eggs respectively. On February 18 a capsule 9 by 6 mm.sevmi. A. leaving fort3'-two active young and three adult . 20 specimens 3. They contained thirty-five. On June 20 two egg cap. 1932. On August 15. in diameter with 2^/l> whorls. On October 23. 18 specimens 2. mea. in diameter with 2 whorls.5 mm. in diameter with 2 whorls.: Nidification and Emhnjoloykal Development 35 On February 15 three cajjsules were laid by the adult snails. 1 specimen 4. On December 28.0 mm. in diameter with 2% whorls. in diameter with 3 whorls. in diameter with 21-) whorls. until June 25.5 mm. It had grown from a diameter of 0.1 mm. an interval of 482 days or sixteen months. one specimen only remained. two capsules were laid containing thirty-one and twenty-three eggs. 1 . only five young . C. LaRocque.specimen 4. This specimen had li\ed from February 28.0 mm. and on February 20 four cap. The measurements of these survivors are shown below: 1 specimen 2.0 mm. in diameter with 3 whorls.5 mm.3 mm.snails remained alive.5 mm.5 mm. in diameter with three and one-half whorls.ys. as noted below: 1 specimen 4. in diameter with 3 whorls.2 mm. fifty-four young snails died. in diameter with 1V> whorls. the aquarium was examined and the shells counted and measured. Baker. On March 1 and 2. 1 specimen 3. 6 by 5. 1 specimen 4. the great mortality being largely due to the very hot summer. and thirteen eggs respectively and development occupied thirteen to fifteen days. 1932. from February 28 until March 12.0 mm.8 mm. in diameter with 2i/-i w'horls. factors of volume of water. 6 specimens 3. thirty-three. twenty-eight.suring: 1 specimen 4. in diameter with 3 whorls. 2 sjjecimens 5.sules were deposited measuring 6 by 5. was laid containing forty eggs.9 mm. 141 specimens in all which had hatched from the capsules enumerated above.0 mm.

Plnnorbis albus Miiller {^ Gyraulus albus (Miiller)).) {^^ Anisus vortex (Linn. 13. 5. pp. 16. 17. containing 394 eggs.)). the capsules measuring 0. Development occupied ten to twelve days. Eight capsules containing 10. and 20 eggs. Egg capsules 9. 1 capsule containing 10 eggs.5 mm. and 21 eggs. and 8 eggs. 5.)).) {^ Planorbarius corneus (Linn. Hclisoma traskii (Lea). 4. Gyrorbis co7)iplanatus (Linn. France. 1 capsule containing 50 eggs. 2 capsules containing 35 and 85 eggs. California.) ). The capsules and eggs are noted below: May two A 1 1 1 shell shell shell shell shell shell 1 1 1 with with with with with with 2 capsules containing 16 and 24 eggs.25 to 1. in diameter. 3 capsules containing 35. near the type locality for traskii. Planorbis contortus Linn. and obser\ed the development of some of the embryos. 11. Stanley Siegfus of Bakersfield. received from Mr.) {^ Planorbis planorbis (Linn. preserved in alcohol.: 36 The Molluscan Family Planorhidae 7 a capsule was laid which measured 7 by 4. Capsules 3 mm. 2 capsules containing 45 and 42 eggs. 13. and rarely 6 eggs Mociuin-Tandon in each capsule. California. in diameter. shells. Planorbis corneus (Linn. Gyrorbis vortex (Linn. Capsules containGyrorbis carinatus (Beck in Moller) { ing ten to twenty eggs each. Development required fifteen to sixteen days. 6. in diameter containing twelve to forty eggs each. Hippeutis jontanus (Lightfoot). Development occupied ten to fifteen days.) ). 7. 425-439) collected egg capsules from water bodies in the environs of Toulouse. Capsules 3 to 4 mm. had eleven egg capsules on the capsule containing fifty eggs is figured (13) on plate 52. in diameter containing four to ten eggs. Nine egg capsules containing 3. in area and contained twentyeggs. 11. 12. Brooks (1880) has recorded the early stages in the development of the small snail Gyrauhis parvus (Say) {= Planorbis parvus Say). Development occupied ten to twelve daj^s. Some of his observations are recorded below Segrnentina nitida (Miiller). (1855. Development was not obser\ed. {= Bathyomphahis contort us (Linn. 22. ^ . The specimens were from the Buena Vista Canal. and 15 mm.5 mm. in diameter containing six to eight eggs. Ten to twelve eggs in a capsule which were 4 mm. Development required ten to twelve days. Plnnorbis cnrinatus). outlet of Kern Lake. Six shells of this species. Several capsules with 3. 4.

The Pleistocene planorbid fauna is THE GEOLOGICAL HISTORY AND DISTRIBUTION ANCESTRY of the Planorbidae. Belly River. Eocene. it is obvious that the family in some form must have existed in earlier strata. is known from the Liassic of England. practically the same as the recent fauna. and in the Oligocene more than eleven species are known. but earlier records are very few in number. then migrated to the land and finally again returned to aquatic habitats. and Pliocene formations have yielded many characteristic species. were originally aquatic. On the continent a large number of species and races of this family have been recorded from Nevada and fresh-water and land mollusks nearly all of the later geological periods. Planorbis mendipensis IVIoore. have not yet been reported from these older geological periods. and also in the several divisions known to belong to the Upper Cretaceous. A ties are known from various localifrom strata of Pennsylvanian age. In America (see Henderson. Oligocene. Perhaps this was accomplished through such groups as the Auriculidae {Carychium for example) which have the Basommatophorous organization but inhabit the land. In Europe this family is fairly common in rocks of IMesozoic and Cenozoic age. and others. It is cjuite probable that these large species are members of the genus Planorbarius and are related to the large Planorbarius corneus so common in the recent fauna. Of these two are of Jurassic or Comanchean age. In the Eocene a number of species of this family occur. as Judith River. During Tertiary time the family was well represented. shrouded in mystery. believed to be at the top of the Lower Cretaceous (Comanchean). especially some of the large forms which are listed under the genus Planorbina for it does not occur in Euroi^e. a species. As the few specimens of planorbid shells from Jurassic or Lower Cretaceous time are not notably different in general characteristics from those of later geological periods or from the modern fauna. 1935) sixty-three species of fossil planor. In the Upper Oolite another species appears. 37 . Triassic and Physa (prisca WolCarboniferous (Pennsylvanian and Mississippian) cott) and other fresh-water groups are reported from the Mississippian of . as of other groups of the Basommatophora. In America the family apparently first appears in the Morrison formation which is considered of Jurassic or Early Cretaceous age. or at the base of the Upper Cretaceous. bids have been described. The family is not uncommon in the Bear River formation. ]\Iiocene. A Triassic planorbid has already been mentioned. however. Planorbis fisheri Forbes. Planorbids. In Europe. The family is not uncommon in later periods. as of other members is of the puhnonate molkisks. probably Jurassic. Wenz (1923) has listed all of these but many of the species have obviously been placed in the wrong genera. It is said that the ancestors of the Planorbidae. DISTRIBUTION IN TIME AND SPACE A.IV.

Vorticijex. five from Upper Cretaceous. as in Planorbula and Tropicorbis. basing the decision on the similarity of the fossil species to some known living species of which the anatomy is known. Carinifex. Nearly all of the fossil planorbids of America and Europe have been described as With the greatest of care it is possible to place some of these species in their correct genera as understood at the present time. Perrinilla.38 The Molluscan Family Plaiwrbidae two from Comanchean.' Tropicorbis. where some species of the latter genus have internal lamellae in the aperture similar to species of the genus Planorbula. A small group of Helisoma (Seminolina) occurs only in the peninsula of Florida. the Planorbidae are world-wide in their geographic distribution. 2 species. Planorbifex. 1 species. Baker's death prevented him from completing Part II of this monograph. thirty from Pliocene. GEOGRAPHICAL DISTRIBUTION Like the Lymnaeidae. and five simply described as 'Tertiary. *As explained elsewhere. Pompholopsis. . 1 sjjecies. 6 species. 2 species. Fossil species of the family have been recorded from Europe. It cally all of the proposed as Planorbis may later be placed in more in Part II of this work to review criti- American fossil Planorbidae. Platytapkius. Australia. 2 species. Menetus is American and does not extend south of the United States. This genus is numerous in North America. Menetus. Helisonia. Africa. 1 species. however. Asia. Helisoma is wholly American but is known from both North and South America and from some of the West Indies. There is scarcely a known region in which some of the planorbid snails do not live. eight from Eocene. 1933). Gyraulus. is more or less limited.* The geographical distribution of the extinct species of the family Planorbidae is not very different from that of the recent species of the family. Among American fossils the following genera are known to be represented by several species and races: 'Planorhis. Planorbula is found only in eastern North America from Alabama northward. some of the islands of the Pacific Ocean. principally the northern part of the United States and Canada. A fossil species has been reported from northeast Siberia (Lindholm. five from Miocene. The two genera are far ai^art anatomically. Parapholyx.' It is obviously very difficult to place fossil mollusks in appropriate genera in a classification based on anatomical characteristics. Typical Planorbis and Planorbarius are mostly confined to Europe and western Asia. 1 species. wdth possibly some forms of circum-Mediterranean distribution. 2 species. Paraplanorbis. Many West of the species now known is appropriate genera. however. can only be tentatively i^laced. Mr. Some forms. including several from the Indies and Central America. The distribution by genera. six from Oligocene. North and South America. Even among modern species tliis method is fraught with danger because of similar resemblances of the shells of several genera. B. and to endeavor to place each species in its correct genus. 8 species. 8 species. and the islands of the West Indies. 5 species. Carinifex and Parapholyx are west American in distribution.

Some species of Tropicorbis have the for example. Grenada. and in the southern part of the United States bordering the Gulf of Mexico. How. as Polypylis. Segmentina is typically European. a closely related group. isolated by large and deep areas of salt water? Examining the distribution of Aiistralorbis.s is known from South America and the West Indies but not found north of the Isthmus of Panama. we are struck by the similarity of the geographic routes of the birds with the geographic distribution of the snails. Taphius occurs only in Central America and in northern South America. is found in northern and central Africa and resembles Tropicorbis so closely. The presence occurring in eastern and northern South America the Lesser Antilles. Vincent. Hence our knowledge of distribution has become more definite and better understood than was possible when shell characters were given chief taxonomic value. Dominica. Marie . It is typically a northern group (palaearctic) but a few species extend southward into the tropical regions. Tropicorbis occurs in South and Central America. etc. Europe. to eastern Asia Dispersal of Species of species of mollusks in isolated places has been somewhat of a puzzle to many zoologists. Gyraulus is the most widely distributed of any of the genera of Planorbidae. Martinique. South America. the Greater Antilles. The shell alone is not sufficient to distinguish groups because there are parallels in growth tendency which render shells of widely separated genera confusingly similar in appearance. Several groups related to Segmentina. in Mexico. and north African. Puerto Rico. Helicorbis. through the Caribbean and West Indian islands as follows: Trinidad. St. Lucia. One route is from Venezuela. in both shell and anatomy. Australorbis. Asia. It is of interest to note that in considering the distribution of the genera. west Asian. Natural dispersal by migration is out of the question. the West Indies. Aiistralorbis and Helisonia. especially the wading birds and ducks. Instances of this are found in the species of Tropicorbis and Planorbida. Segmentina and Polypylis. details of the anatomy have been the only means by which the different groupings could be made with certainty.Distribution in Time and Space 39 Ai(. and Intha.str(dorbi. known as Afroplanorbis. or Drepanotrema found their way to so many of the islands of the West Indies. A former connection with South America occurred too remotely to answer satisfactorily the question. for example. are confined and some of the Pacific islands. The present study of the distribution of the members of the family Planorbidae shows conclusively the value of the anatomy in the classification of the species. populated by one or more species of planorbids or other fresh-water species? In what manner have the same species of Tropicorbis. that it is a question whether the two groups are not identical is generically. Pingiella. Driftwood might answer for land snails (at least to a limited degree) but not for fresh.water forms. being known from North America. Africa. Curiously enough. and Jamaica. When we examine the migration routes of the migrating birds. and some of the islands of the Pacific as w-ell as from the West Indies. in we find it and also many of the groups of islands — same distribution. are ponds without inlet or outlet and more or less widely separated. St. Anisus is mostly European and Asiatic.

especially beetles. pay no attention to what may be attached to the feet of their victims. especially in populating closely associated ponds or streanis. Tropicorbis. In the distribution and dispersal of the Mollusca. By this means in a few years a body of water would be a prolific habitat for the species. Some migrant birds pass up the west coast of Central America and ]\Iexico. and Cuba. Thomas. to be liberated when the birds make other stops. In the case of the Basommatophorous mollusks a single snail introduced into a pond would be sufficient to insure population since selffertilization is easily accomplished. Natural migration doubtless takes place in connected streams and other bodies of water and will account for the dispersal of related species within the drainage of a water system. Christopher. Jamaica. Antigua. St. Guatemala. close to Costa Rica. The genus Tropicorbis has this distribution and many of its species are widely distributed. would perform the same service. Floods often carry species of snails over natural divides. Young snails have been found attached to the wings (elytra) of beetles. eastern Mexico to the gulf coast of the United States. Honduras. The scarcity of records of snails on the bodies of migrating birds is primarily due to the fact that those who kill the birds. Another route is from Venezuela northward up the eastern coast of Panama. Haiti. the snails become attached to the feet or other parts of the bodies of the birds. do not carefully examine the bodies of the specimens and the snails fall off or are brushed off during the preparation of the skin for the cabinet. They stop at these islands to feed and rest. Puerto Rico. Aquatic insects. sportsmen and hunters. Ornithologists. and Drepanotrema inhabit most of the islands mentioned. Careful examination of the bodies of birds would doubtless provide many valuable evidences for avian dispersal of fresh-water and other mollusks. St. too. all agencies must be taken into account.40 The MoUuscan Family Planorbidae Galente. It is the writer's belief that the populating of many if not all of the islands of the West Indies with planorbid and other fresh-water mollusks has been brought about through the agency of migrating birds. Young snails have been found attached to the feet of many birds. The three genera Australorbis. Nicaragua. .

AV.— V. and Ancylinae. Planorbidae. Lymnaeidae. Ancylus was made a separate family. and Physidae. Bryant "Walker 1918) in his 'Synopsis' recognized four families. F.). within which he recognized Ancylinae. Planorbinae. P. HISTORICAL SKETCH OF CLASSIFICATIONS VOLU:\IES on the IMollusca. Planorbinae. George W. and allied genera in the family Limnaeidae. Pompholiginae. Li 1870. Physa he relegated to a separate family. Planorbis. C. and Ancylus are all placed in the family Limnaeidae. Limnaeinae. In 1872. and Ancylidae (Germain. W. as well other genera. Pompholiginae. Limnophysidae. separated Physidae as a family but places the subfamilies Limnaeinae. Tryon. in the third volume of his Structural and Systematic Conchology. nearly all of the genera of fresh-water pulmonates Limnaea. Bulinus. In 1902. Dybowski (1903) proposed a classification of tlie Basommatophora in which six families were recognized. placed all groups under the family Physidae. and Ancylidae. G. and Ancylus appear in the family Limnaeidae. Physa. Planorbinae. Paul Fischer. Amphipeplidae. placed all of the fresh-water pulmonates under the family Limnaeidae with subfamilies IXas THE EARLY — ( (including Physa. in 1895. and Ancylinae as subfamilies. Binney. in S. Baker in 1902 and Dall in 1905 separated Physa and Aplexa to form the family Physidae but placed Limnaea [Lymnaea) Planorbis. Physa being included in the subfamily Linmaeinae. and Planorbinae as subfamilies. Westerlund placed all of the Basommatophorous mollusks in the family Limnaeidae with four subfamilies. most of the important monographs and lists have accepted the four-family division of the Basommatophora. From 1920 onward. Jr. in which the fresh-water pulmonates are all placed in the family Limnaeidae with Limnaeinae. Physidae. Planorbis. by Baker. Woodward's Manual of the Mollusca. Chilina. and Ancylinae in the Limnaeidae. Limnaeinae. a year later (1884). Kennard and ( 41 . Planorbinae. but separated the principal groups as subfamilies: Limnaeinae. Ancylidae. Dall proposed a classification in which four subfamilies were recognized: Limnaeinae. Physinae. 1921-23. Physa. etc. and An1842-1844) who cylinae. Tryon. published a work on The Land and Fresh-Water Shells of North America.. Cooke. Camptocercinae. H. Limnaeidae. Ancylidae. Thus in 18511856. Planorbinae. in 1865. In this grouping he was following Haldeman |)laced Planorbis and other groups in the family Physidae. Planorbidae. in a Monograph of the Freshicater Univalve Mollusca of the United States. Planorbinae. by Forbes and Hanley. in the Manuel de Conchyliologic (1883). Lymnaeidae. were largely classified by the shells and were placed in families with wide limits as to the genera involved. the planorbids. Aplexa. Planorbidae. Physidae. ^legasystrophinae. in 1853 (volume iv) the fresh-water pulmonates of the genera Limnaea. and Pompholiginae. and Ancylinae. listed most of the fresh-water pulmonates under the family Limnaeidae. A. In A History of British Mollusca. H. evidently following Paul Fischer. Limnaeinae .

of which there are many. ]\Iost of the early papers or works on anatomy simply gave as illustration a familiar example. indicating a tendency to recognize smaller group differences and also indicating advance in knowledge concerning these groups. such as Clessin 1886) and Sowerby (Reeve. F. Preston. for example. Pilsbrv and bv H. Tropidiscus. Lindholm. and Segmentina. 1872). were diagnosed from shell characters for the most part. Many other works might be listed. Many generic and subgeneric names have been proposed during the past ninety or one hundred years but these were founded almost wholly on shell characteristics. Woodward. It is to be observed. the groups of which were thought to constitute a family. thiele. Haas. as Lindholm. mostly fossil groups. Soos. 1927. Anisus. recognizing Segmentina as a distinct genus. Baudelot (1863) published good figures of the genitalia and other organs of Planorbis corneus {= Planorbarius) In 1867. Menetus. including Planorbis corneus. Pallery. A. C. Armiger. 1929. Pompholyx. leucostoma. and Jontanus. especially by European writers. Baker recognized the groups Helisoma. Gyraulus. Studies of the anatomy of the Planorbidae began in Europe many years ago but were confined to only a few species. as. Tryon listed most of the group names as subgenera of Planorbis. and contortus. spirorbis. Ficinus. No such analytical studies as those on the land mollusks by H. listing all groups as sections under the genera Planorbis and Anisus. From one family in 1870 (Limnaeidae or Physidae) this grouping has grown to five at the present time. that all of these genera or subgenera as treated by early authors. published an article on the penis of Planorbidae in which he divided the European species of Planorbis into two groups. (1) with a penial stylet. Carinifex. The large monographs. the four previously alluded to and Bulinidae.42 The MoUuscan Family Planorbidae Pilsbry and Bequaert. and (2) without a stylet. Germain (1931). followed Dall. 1928. Mori (1938) in his classification of Japanese Planorbidae followed Thiele. Dybowski. an almost forgotten WTiter. Baker. and Gyraulus as being of generic rank and in 1928 fully described these groups. . In 1884. Odhner. Among European works. and others. In the matter of generic distinctions among the planorbids there is little of note in the earlier works. Planorbis corneus and Limnaea stagnalis figured by MoquinTandon in 1855. Baker were made until after the year 1920. B. do not help much in family distinctions and the lesser faunal lists of both continents. Kennard and Woodward (1926) did likewise. 1926. simply copy the classification as published in some of the works mentioned above. giving anatomical reasons for their distinction. In a review of the classifications of the Basommatophorous grouji the change in number of families and genera recognized is noteworthy. a great step in advancement over previous works as regards classification. in his Mollusks of France. including Planorbis vortex. F. Westerlund. Hippeutis. 1921. albus. In 1926. He erected the genus Appendicularia for Planorbis ( . The most important of these are listed in the synonymy under the different genera. 1926. C. and others). however. Thiele in his Handbuch (1931) was too conservative. In 1905. Lindholm in 1926 recognized as of generic rank Planorbis. recognized five families. Spiralina. Dall considered all groups but Segmenti7Ui as subgenera of Planorbis. Germain. and Choanomphalus are listed as genera with some subgenera. nitidus. Planorbula. Clessin. Bathyomphalus. 1931.

vortex. were given. Gyrorbis spirorbis. The difference between the prostate diverticula of Planorbis corneus and the smaller species. F.s and F. In 1917. and other anatomical features. I. No attempt was made.. In 1891. pp. Soos published a paper on the anatomy of the Hungarian Planorbidae. appeared F. to base a classification on the anatomical data. This was the most modern treatment which the family had received and the key is the first attempt to separate the genera by the use of anatomical sperm outlet in the center. one new and one following Annandale. adding greatly to our knowledge and clearing up many doubtful jioints in the classification based on the anatomy. and radulae of the species of Planorbidae inhabiting the Middle West. There was a good account of the excretory organ (kidney). several notable papers appeared on the anatomy of the family. Rao published an exhaustive treatise on the anatomy of Annandale's new genus Indoplanorbis. and Gyraulus albus. radulae. Also in 1931. Connolly (1925) figured the genitalia and radula of Planorbis pfeiffen which indicated that it was related to the West Indian genus Tropicorbis. C. of Wis. The subfamily Helisominae (Helisomatinae) was proposed as new on page 309 of that work. In 1923. 306-387) presented descriptions and figures of the genitalia. On page 22 a key appeared in which the groups were briefly diagnosed from the anatomical standi)oint. several critical papers appeared on the anatomy of the Planorbidae. rotundatus. Germain issued his work on the land and fresh-water mollusks of France in which outline figures of the genitalia. carinatus.. vortex. In 1929. Simroth's account (1912) of the genitalia of the Planorbidae was based on Buchner's work and added nothing to our knowledge of the antomy of the group. and some of the radulae. Odhner published an account of the Alollusca of the Tiikerns in which the genera were separated by characteristics of the genitalia. Baker's paper . Baker. published notes on the genitalia. and albus did not distinguish the peculiar gland of corneus from the penis. This is the best work on the anatomy of the Planorbidae which had appeared up to this time.Historical SkelcJ) oj Classifications 43 7iiti(lu. During the decade 1931 to 1940. Buchner pubHshed the first comprehensive account of the genitalia of the family Planorbidae. Tropidiscus planorbis. L. The radulae and the digestive system were also considered. the first attempt to chissify the Planorbidae by reference to details of the anatomy. in 1926. Bathyomphalus contortus. giving figures of the genitalia of Planorbis corneus. (2) nitidus and cotnplanatus with flagella and sperm outlet at the side. etc. (3) marginatus and carinatus with and (4) contortus. jaws. Annandale (1922) and Annandale and Prashad (1919-1921) added to our knowledge of the anatomy of the Indian Planorbidae. jaws. as far as known to the writer. and radulae of certain American Planorbidae and in 1928 (AIolI. Segmentina nitida. however. During the decade from 1921 to 1930. C. He divided the male organ into four types. nitidus. In 1931. fontdnus because of the presence of the flagella on the penis. with a stylet. w'as especially mentioned but no reference was made of the presence of a separate prostate duct. (1) corneus with a small penis. The classification of the genitalia is based on that of Buchner with the addition of two types. This is. He differences.

Planorbulinae. their genitalia were figured. in 1933. in 1935. The same author. Soos published a paper on the Planorbidae of Hungary which included good figures of the genitalia of the species inhabiting that country. Menetus. From the foregoing summary it is plainly obvious that a detailed study of the classification of the family Planorbidae. The classification of other groups. comparable to the methods used by Pilsbry on the land mollusks. Later.44 The Molluscan Family Planorbidae on the classification of the larger planorbes of Europe and America.discussed and a new genus Australorbis was proposed for the large snails inhabiting South America and the West Indies. In 1934. Pilsbry discussed the planorbid fauna of Florida. . It is not overstating to say that this paper is the most comprehensive account of the family Planorbidae from a modern standpoint that has yet appeared. based on anatomical features. Carinifex. was established. is still to be prepared. The peculiar penial gland of the genus Helisoma was discussed at length. Planorbis wmbilicatellus. It is believed that a foundation has been laid upon which to build a natural classification of the fanfily Planorbidae.inolina) was proposed for the group which is not found outside of the Florida peninsula. An attempt to supply this deficiency is the purpose of the present work. the same author published short papers placing certain disputed species in their proper generic position. A. In 1935. A new subfamily. . in which Planorbis corneus and the American Helisoma species were discussed and the genitalia and radulae figured. Tropicorbis. and a new name {Sem. based on anatomical information. The European genus Anisus was ably discussed. based on their anatomy. long thought to be a Gyraulus. was shown to be a member of the genus Menetus and a new subgenus was proposed for Menetus exacuous (Promenetus) In 1936. and Parapholyx was . Characteristic keys to the genera of American Planorbidae were given. a paper from the pen of H. made a study of the peculiar genitalia of Indoplanorbis exustus of India adding some features to the previous paper by Rao. A. Planorbis obstructus was shown to belong in the genus Tropicorbis and the true relationship of Parapholyx was indicated. Thus.

the form of the radula teeth. that have given criteria for generic and higher distinctions. No such comparable work has been done on the fresh-water pulmonates (Basommatophora). great stress is laid on the differences in the male genitalia. in addition to several new genera. flagella present or absent on the vergic sac. as the shape of the kidney. Henry A. A NEW CLASSIFICATION OF THE FAMILY PLANORBIDAE GENERAL DISCUSSION THE PAST FIFTY YEARS. the DURIXG(Stylommatophora) under the able leadership shells . While the shell has helped in many cases it has been the internal features. In the new classification here presented. chiefiy the genitalia. of the Academy of Natural Sciences of Philadelphia.VI. that only by the careful study of the internal organs of the animal can a natural classification be secured. This has been accomplished by the study of anatomical details of the genera and species. whether it is smooth or has a ridge. a penial gland in the preputium and whether this has an internal or an external duct or none at all. Pilsbry. The family Planorbidae offers better characteristics for purposes of classification than does the Lymnaeidae. whether the diverticula are in a double series or are multiple. has been brought to a high degree of accuracy and completeness. although the shell sometimes presents characteristics of generic importance. The results obtained by these studies have been accepted by nearly all malacologists and conchologists. there being a greater diversity of structural features among the different genera. were adequate to represent the different anatomical features involved. 45 . These represent a majority of the generic groups so far published. The pseudobranch also in the family. the shell offers less assistance than the organs of the animal. Other anatomical features also aid in classification. As in the study of the land mollusks. from different parts of the world. Few new groups were necessary because those already published. and the few attempts to apply anatomical criteria for generic distinctions. of which there were many. the shape of the stomach and the turns of the intestine from stomach to anus. the characteristics of the prostate diverticula and the prostate duct. the pseudobranchs of the two groups being markedly different. were dissected. whether simple or fragmented. For the data in the present work a large number of species of the family Planorbidae. whether at the end or on the side. as used in the Pulmonata. This was attempted by the writer for the Lymnaeidae in 1911 and some gratif^'ing results were obtained. For generic purposes. It offers some criteria although this organ is fairly uniform does effectively separate the Planorbidae from the Bulinidae. the opening of the sperm canal in the verge. It is obvious. the size of the preputium and vergic sac. the presence or absence of a stylet on the verge. classification of the land of Dr. bringing together groups which are allied by features of the natural organization of the animal. however. the genitalia have been of the greatest assistance. and the nature of the ovotestis. the condition of the jaws. the number of retractor muscles of the penial complex. have met with more or less criticism.

b. or less in diameter. end Verge with . 4a.stylet. ANATOMICAL KEYS TO GROUPS* Key la. or more in diameter. A few groups. . the contrasting conditions being arranged as 'a' and 'b' under the same numeral. Prostate diverticula not on separate duct. penial gland duct inside preputium. Platytaphius. ovotestis with paired diverticula. out 3a. 4 5 57). H. simple. — . 85) *In using this type of alternative key the options are always arranged in couplets. but it is believed that when examined they will be found to fit into this classification. This form of continuous and consecutive key is less confusing than the type in which descriptions to be compared are widely separated from each other and does not offer the technical diflScuIties of composition inherent in the indented or step form of key. b. If the description under la fits the specimen or concept under consideration. depression 5a. 2a. branched at 2 i). 46).. Vergic sac without flagella 3a. Vergic sac about as long as preputium.^'-C. to Genera of Planorl)inac+ Prostate diverticula on separate duct. (see p. It is confidently believed that the classification herein presented will form a reliable foundation on which to build future knowledge which may be obtained concerning this interesting family.J. . Vergic sac le. as Taphius. the numeral at the end of that line indicates the next couplet to be tried (except in those instances where la leads directly to an identification). Shell with b. Prostate a single row of diverticula usually placed on a separate prostate duct. and all of the genera. Many species of the family still remain of wliich the animal characteristics are unknown and it is possible that new genera or subgenera will need to be made for their inclusion. Species inhabiting 8a. .2 3 2a. Anniger (see p. and Trochorbis. Vergic sac with flagella b. Left side of shell with whorls depressed as though reamed (Subgenus) Torquis (see p. b.Planorbulinae (see p. readily fit into this subfamily classification. 47) Planorbinae (see p. tSome subgenera which have distinctive morphological characters are included in this key and in all succeeding keys to genera. 66).. outlet of sperm canal Aiifitralorhift (see 90).Gi/m»/i<. shell 3 than one-fourth length of preputium b. The reader always starts with la. 8 not costate b. Prostate forming a fan-shaped pattern in cross section. (see p. 51).s. Segmentininae (see p. Biomphalaria. 72) 6a. side at end 3 6 many closely coiled whorls not increasing greatly in diameter few loosely coiled whorls rapidly increasing in diameter Anisus s. fanwise Prostate forming a finger-shaped pattern with few diverticula. shell 5 mm. to Subfamilies . and so on through tile key until a group name is reached at the end of one of the options. . 62). unbranched at end b. 7 7a. 47).statement under la does not fit. Shell with . as far as they have been examined. (see p.s s. . Left side of shell flat without central depression b. outlet of sperm canal at Verge without stylet. 86) 9 Tropicorbis s. Prostate with multiple diverticula usually placed on the sperm duct . 46 The Molluscan Family Planorbidae It has been possible to recognize four subfamilies of the family Planorbidae based on fundamental anatomical differences. 48). Left side of shell with rounded whorls on a plane surface. Vergic sac longer than preputium Planorhis (see p. penial gland duct on outside of preputium. b. the alternative under lb should. Key la.s. ovotestis consisting of several diverticula arranged Helisomatinae (see p. Species inhabiting Africa America Afroplanorbis (see p. 75) usually costate b. "\>rgic sac much longer than preputium. If the . Left side of shell with deejily excavated (Subgenus) Bathijomphnlus (see p.s. are still unknown anatomically.

penial gland a simple cup attached 'to wall of preputium. with penial gland in preputium Genitalia without penial gland in preputium 2a. penial gland an elongated cup with wide a]ierture Carinifex (see p. the preputium much wider than the vergic sac. which is elongate. (Subgenus) Scminolina (seep. 161). Flagellum wide and short. often bifid at the end 7a. Polypylis (see p.s. thick. penial gland bifurcated.. pear-shaped. . (Subgenus) Fo. in diameter Acrorbis (see p. 6a. verge slender with vergic appendage at end Segmentina (see p. complex with vergic sac and preputium equal in length. in a coil between vergic sac and preputium 5 Penial complex with vergic sac shorter than the preputium. Preputium with external penial gland duct 2 3 6 . sac-like. . 96). 100). with a long "stem" attached to a short "bowl" Parapholijx (see p. Shell with campanulate or bell-shaped aperture (Subgenus) Planorbella (seep. external penial gland duct on preputitmi Without penial gland duct on preputium External duct very short. Key la. slender. 109).s. Classification of the Family Planorbidae 47 b. 85).s s. Penial duct twice as long as in 6a. not campanulate. PreiHitiinn narrow. (Subgenus) Picrostnna (see p. verge wide at upper part. . Preputium wide. 4a. Penial . 5a. complex with one retractor muscle complex with two retractor muscles Penial complex with but little constriction between vergic sac and preputium.Hclisoma s. . to Genera of Helisomatinae+ . Ijenial gland duct about half as long as preinitium. . 4a.. vergic sac about twice as long as preputium. wide. 104) Flagellum shorter than penial complex. . b. flagella more than twice as long as vergic .PlnnorbariJi^ (. Shell with b. jjenial gland duct several times as long as preputium.<>!^ulorbis (see !>. 118). Flagellum long and narrow. (see p. without penial appendage Hippeutis (see p.s. . Flagellum as long as penial complex. flagella consisting of small cylindrical sacs attached long ducts Pingiella 4 b. s.see p. b. swollen or club-. in diameter 6 b. not longer than preputiimi 6 Shell with simple aperture. to very (see p. b. Drepanotrema (see Drepanotrema p. penial gland pijie-shaped.4 2a. . (see Aperture with internal lamellae during a i)art of the shells growth (Subgenus) Ohstrurtio (see p. orbicular 10 with loosely coiled whorls rapidly increasing in diameter (Subgenus) Latcorhis (see p. 124) Penial complex with notable constriction between vergic sac and preputium. Prejiutium with i)enial gland but without penial duct b. Ajierture without internal lamellae at b. is *One subgenus included among the groups differentiated in tliis key. vergic sac longer than preputium. placed between vergic sac and preputium. A Xew Oil. 154). penial gland duct as long as the jn-eputium. (see p. any stage of growth Tropicurhi. not swollen at summit. 6a. Penial 3a. vergic sac and preputium of about the same length shell more than 5 mm. extending from upper part of ju'eputium to vergic sac. 150).. . club-shaped.5 mm. 134).. shell very small. (see p. b. Shell Shell with closely coiled whorls slowly increasing in diameter.sic. vergic sac shorter than preputium..?haped at sununit b. An 3a. 85) 85). 114) 7 Key la. b. Shell with rounded pcrijihery angulated or carinated i>erii)hery. 129). p. 118). . penial gland short and thick. 112). 10a. b. 5a. attached to short ducts Intha External duct about four times as long as preinitium. 1. 121). Genitalia to Genera of Segnientininae* 2 5 3 b. Penial b. sausage-shaped. half as long as preputium. 166). Penial duct on outside of preputium very short. tSeveral subgenera are also differentiated in this key.

Planorbis strauchianus Clessin Subgenus Costorbis Lindholm. along a large part of the ovisperm duct collar or SYSTEMATIC LIST OF GENERA AND SUBGENERA RECOGNIZED IN THIS WORK In the following list the genera and subgenera are arranged in the sequence which appears most in accord with their structural features. 1840 Planorbis andecolus Orb.t 1875 *One subgenus is included in this key. 1931 Genus Syrioplanorbis F. la. Planorbis janeirensis Clessin Planorbis pallidus C. tThe anatomy is not known for any member of JA group represented by fossils only. the surface Penial gland connected with muscular collar by an open slit in the muscular wall of the preputium. 187).s. Penial gland sausage-shaped with round. 178). Adams. and A. with open. Two groups of uncertain affinities. ovate. 1837. seminal vesicle consisting of many short diverticula stretched Frotnenetus (see p. Penial gland flattened. Family PLANORBIDAE H. it is obvious. Penial duct very short leading from gland direct to muscular diaphragm b. 1837 Planorbis albiis Miiller Subgenus Gyraulus SS Planorbis parvus Say Subgenus Torquis Dall.) Charp. . liebmanni T>nn\s. Planorbis krambergeri Halavdts Genus Odontogyrorbis L6renthey. C. The fossil groups are marked by a dagger.. . f 1910 Genus Australorbis Pilsbry. channel-like duct 2a.t 1906 Genus Gyraulus (Ag. (Subgenus) Micromenetus (see p. 1820 (Gray.sec \\. Baker (New) Genus Afroplanorbis Thiele. diaphragm gland connected with muscular collar by an open duct as long as the gland. 48 The Molluscan Family Planorhidae Key to Genera of Planorbulinae* 2 3 collar or (see s. Genus Taphius H. The fossil species. Adams Planorbis sudanicus Martens for Planorbis libanicus West.s. 1939 Subgenus Lateorbis F. They are placed in the subfamilies and near the genera with which they appear to be affiliated. The type species is indicated following the name of each genus or subgenus. p. Baker. the two pilasters sending branches from both sides to this slit. 1767 Genus Anisus Studer. Menet us 183) Penial duct long.. Genus Armiger Hartmann. B. seminal vesicle composed of many diverticula forming a mulberry appearance on Flanorbuln s. Penial b. Adams. Several groups are unknown anatomically and these are marked by an asterisk... (.) Charp. 173). Subfamily Planorbinae 1934 Genus Planorbis Geoffrov. Helix spirorbis Linn. preoccupied Genus Biomphalaria Preston. C. 1855 Pilsln-y. 1914 P. 1934 Genus Anisopsis Sandberger. lying above a single diagonal projection of the left pilaster. one recent and one fossil. extending up the side of the preputium to the muscular 3a. closed duct b. can never be known anatomically and these are placed near those genera which they most nearly resemble.Helix contorta Linn.Qr = orbiculus Subgenus Tropicorbis" SS Morelet Subgenus Obstructio Haas.t 1926 Subgenus Bathyomphalus (Ag. 1855 Genus Tropicorbis Pilsbry and Brown. are placed at the end of the family sequence. and A. 1847) Subgenus Anisus SS Helix planorbis Linn. f new name Heterodfscus West. Biomphalaria smithi Preston Planorbis glabratus Say Planorbis calculus Sandberger this group. 1905 Nautilus crista Linn.

PLANORBULINAE Pilsbry. 1922 Intha capitis Annandale Planorbis anatinum Orb. G. J 1934 Groups of Planorbis armigerus Say Planorbis icheatleyi Lea Planorbis exacuous Say .. 1855 Subgenus Menetus SS Subgenus Mieromenetus F. 1863 Genus Vorticifex Meek. Planorbis condoni Hanna Planorbis hcteroplearus P. 1922 Genus Pompholopsis Call. 1840 Subgenus Helisoma SS Subgenus Seminolina Pilsl^ry. J 1912 Geiuis Parapholyx Hanna. 1840 Subgenus Planorbula SS Sul:)genus Haldemanina Dall. Phil.! 1853 tThe anatomy is Choanomphalus maacki Gerstfeldt Valvata multiformis Zeiten. Planorbis umbilicalis Benson Trochorbis trochoid iics Benson Baker (New) Pyramidula pi'ipinoi-^lx Ping and Yen Intha Annandale.! 1855 Pingiella F. . C. Baker. 1842 Genus Carinifcx W. 1935 Genus Menetus H. 1934 Genus Planorbula Haldeman. Planorbis nitidus Mtiller Planorbis complanatus Drap.t 1859 Genus Poecilospira Morch. C. A group represented by fossils . 1817 Hippeutis (Ag. 1905 Genus Promenetus F.! 1922 Genus Platytaphius Pilsl)ry. f 1855 Trochorbis Benson. {=yzabalensis C. and A.) Physa scalaris Jay Planorbis Planorbis Planorbis Carinifex trivolvis = bicarinatus campanulatus Say newberryi Lea tryoni Meek Helisoma cordillerana Hannibal Pomplinhjx effiisa Lea Pontjiholap-sis irhitei Call Hdix contra Linn.) Charj). 1934 Subgenus Pierosoma Dall. J 1870 Genus Perrinilla Hannibal. C. and V. . C. Baker (New) Genus Planorbifex Pilsbry. only. 49 Siil)family SEGMENTININAE Baker (New) Segmontina Fleming. 1837 Polypylis Pilsi)rv.! 1888 Genus Planorbarius Froriep. C. Acrorbis petricola Odhner F. 1906 Helicorbis Benson. Planorbis opercularis Gould Planorbis dilatatus Gould Planorbis vanvlecki Arnold Uncertain Affinities Genus Choanomphalus Gerstfeldt. Adams. Baker.t 1924 Genus Acrorl^is Odhner. Subgenus Fossulorbis 1934 Genus l'arai)l:in()rl)is Ilanna. 1905 Subgenus Planorbella Haldeman. not known for any member of this group. 1928 Drepanotrema P^ischer and Crosse. Planorhis largillurli. 1880 Subgenus Drepanotrema SS Pilsl)rv. and F.). Jiinney.A New Genus Genus Genus Genus Genus Genus Genus Genus Classification of the Family Planorbidae F. 1937 Subfamily HELISOMATINAE Genus Helisoma Swainson. Planorbis cnltratus Orb. 1806 Subfamily Planorbis anceps { Menke Say Sowb.

Physopsis. The chief feature separating the Planorbidae from the other members of the Limnophila is the presence of an accessory breathing organ. principally in the penial complex. Adams Tvpe genus Planorbis Geoff roy. The shell is carried almost vertically in the typically sinistral groups but leans toward the left in other groups. Anisms. and other related groups should form a separate family Bulinidae. PLANORBIDAE. another distinctive feature of the family. 'planorboid' shell is. This has been recognized by Germain in his mollusks of France (1931) and by Larambergue in a recent publication (1939). The shell of Planorbidae is typically sinistral in some groups (Helisoma) and pseudodextral or iiltradextral in many groups. In these groups the spire is beneath and the umbilicus is above. having a ridged kidney as in Helisoma. and the presence of a penial stylet. Subfamily PLANORBINAE is H. The groui)s of genera related to Bulinm and Physopsis have in the past been included in the family Planorbidae because of the similarity of the radulae and the presence of a pseudobranch. A. being in the form of folds or ridges while in the Planorbidae it is smooth and leaf-like. There are also differences in the genitalia. The ovotestis has the diverticula arranged in a double row.VII. the pseudobranch. Armiger. In the Planorbinae the prostate diverticula are all simple sacs placed in a single row along the whole length of a long prostate duct which is separated from the sperm duct into which it enters. Bathyomphahis. The Indian genus Indoplanorbis has been shown by the latter author to be related to Bulinus and its place appears to be with the Bulinidae. Gyraulus. 1934 equivalent to the family Planorbidae as now recognized by systematists. of The subfamily H. are animals with spiral shells. Here again Australorbis is an exception. principally in the jaws. AND SUBGENERA RECENT AND FOSSIL together with the Physidae and Bulinidae. There is considerable variation among the different genera grouped in Planorbinae. of course. Tropicorbis. principally the smaller forms. These differences may be tabulated as follows: Radula with serrated marginals placed hijih up on the reflection: Planorbis. Pilsbry. and A. the radulae. The kidney is usually rounded and without a superposed ridge. Radula with claw-like marginals on a long reflection: Australorbis. The flattened. 50 . GENERA. the genitalia and the respiratory and excretory organs are on the left side. These differences suggest that Bulinus. An apparent exception is found in tiie genus Australorbis where the single row of glands is placed directly on the sperm duct and the ends of the prostate diverticula are branched. But this organ in Bulinus is different from the pseudobranch in Planorbidae. these being respectively the right and left sides of the shell. In Physa it is multicuspid. which takes over the office of a gill when the animal is submerged and can not obtain free air. The central tooth of the radula is always bicuspid. SYSTEMATIC ACCOUNT OF THE SUBFAMILIES.

Omalodiscus Bexsox. Syn. Gen. by tautonomj' Helix planorbis Linn. (of Cat. Heft 6. Gyraulus. a superior and two lateral jaws: Ajroplanorbis. 11. Alaska Moll. The fossil group supposed to belong to this subfamily.ard and Woodw. Jaw fragmented with many plates: Planorbis. Several groups are unknown anatomically and hence their position in the system can not be definitely known until the animal has been examined. (Xon Dejean. 1855. As subgenus of Planorbis. p. Genotype 6. p. Tropidiscus Clessix. 1774. Bathyomphalus. in three pieces. Xo type 1902.\rtm. [Fitzixger] Herrm. Ind. 76. Anisus.. Tropicorbis. p. As genus. Jour. Hi. ]k 111 type designation. 1924.. p. 1886. As subgenus. Twpicorbis. Xo type designation.\ll.\nx. Bathyomphalus. As genus. Syrioplanorbi)^.-Cab. Tropicorbis. 138. et Fluv.. Moll. Schn. 120. p.\xxsex. Bengal. (part) Hist. As synonym of Planorbis.xrd. p. p. Archiv. X'autilus but cites no type. As subgenus. Ajroplanorbis. pp. Hungarici. p. Genus 1767. Type Planorbis marginatus Drap. Tropidiscus Westerluxd. 1847. Had. Heft Ideogenotype Helix planorbis Linn. 1898. Gen. 428.Subfamilies. and Subgenera — Recent and Fossil 51 Radiila with interstitial cusps on laterals or marginals: Australorbis. 1883. Jugoslav. Syst. Man. 1767 Genotj'pe by tautonomy Helix planorbis Linn. nomenclatorial position 1917. 1905. 7. Planorbis PLAXORBIS Cleoffroy. Conch. Anisus 1855. Centr. Ind. 1921. u. Ill.\RX. Tropidiscus Fischer. France. p. 1888. Taphius. p. Biol. Muller 1774) Wexz. 58 year. 58 year. 9.. Traite Coq. II. 1906. Genera. II. p. 85. (vide Lindholm.\ll. As genus. 40.\p. Terr. Herrmannsen. 12. 1523. ^'erge with stylet Verge without stylet: PUmorbis. Ind. 66. V. Amer. Placorbis DR. p. Rec. Planorbis complanatus Stein... Auslralorbis. Part. Oester. Mai. Paris.. 509. As subgenus. 252). XVII.. Comments on XV.artexs. (part) Beitr.v Gray. As fixed b. Planorbis planorbis (Linn. Muscheln Berlins. Spiralina H. Such are Costorbis. — 1852. : Anisus. British Xon-Marine !Moll.-T\xd. Tropidiscus (Linn... Fauna Palaearc. XXIV. :Mo11. V. and Biomphalaria.. X'at.. Asiatic Soc.). Musei X'at. Tropidiscus (of Stein) Lixdholm. vox M. 64. Conch. Archiv. p.. p. Genotype Helix planorbis Linn. Planorbis 1926. 1923. Planorbis Muller.. Xot Anisus Studer. XXI. Anisus Fitzi. Tropidiscus Sees. fur Mollusk. Geoffroy. X"o type cited. As subgenus. Tj'pe . fiir Mollu. 1885..). As genus. As section. p. France.n-ger.. p. p.. (part) Verm. Tj'pe designated by Clessin. 141. Tjqie: Planorbis umbilicatus Miiller. Ann. Reg. 22. 1886 Planorbis marginatus Drap. Planorbis Kexx. 34. Armiger. No 1833. Otnalodiscus D. Coleoptera). Omalodiscus marginatus Drap. Auslralorbis. Akad. No tj'pe Mai. Planorbis (Geoffroy) cited. 105. p. 1821. Moll. Lande. 1905. 1850. 151. Type Planorbis umbilicatus Muller = XIX. Terr. GyrorbU Moq. 1847. Gyraulus. Tropidiscus Steix. Mus.286. Hist. Suppl.). As genus. can only be placed by shell resemblances and this points to the Planorbinae. Type Planorbis p.\rD. Xat. 152. Jaw Armig( r. No type cited. T3-pe designated by Kennard and Woodward. As genus. Tropidiscus Westerluxd. 380. Tropidiscus D. Fossil. Type Tropidiscus planorbis Germaix'. 1927. planorbis (Linn. 252.

especially at the summit where the vas deferens enters. Oc. et Fluv.) cited. 379.). The prostate diverticula are cylindrical in form and a trifle more than 0. elongate-cylindrical. Mus. Allat. (Linn. Planorbis carinatus Miiller Planorbis planorbis (Linn. p. Mus. eyes sessile at the inner base of the tentacles.) by absolute tautonomy. p. pp. Germain. Seminal vesicle (SV) 3. France. p. 7). Abh. Cien. Both sperm duct and prostate duct enter the smaller vas deferens near the vagina (fig. rounded before and pointed behind. Sperm duct (SPD) very long (14 mm.. Univ. 1936. Tj'pe evidently intended to be Planorbis phniorbis (Linn. As genus. Internally (fig. 29. As subgenus under genus Anisus. B. The verge (V) is short and wide in repose and nearly fills the vergic sac. Type Plaiiorbis planorbis 1935. Nat. As genus. upper and lower (left and right) sides flattened. and p. the preputium (PR) is long (3 mm.) adherent to the oviduct. XIII. a yellowish liver. Mich. Handbuch.) and somewhat less in diameter than the preputimn. figs.). XXXII. rounded. Male Organs (fig. No type desig- 481. Planorbis (of Miiller 1774) Haas. The long. Teil 2. aperture small. 1. p. Tropidiscm Thiele. A cross section through the prostate and oviduct shows the relationship between the sperm duct and the prostate duct (fig. long and consists of forty to forty-five diverticula placed in a single row on a long prostate duct (PD) which is separated from the sperm duct. absolute tautonomy Helix planorbis Linn. The vas deferens is a long (12 mm. 431. A group of five or more small muscles near the base of the preputium and a longer. a carina encircles the body whorl on the left side. The prostate (PRS) is about 5 mm.y H. Baker.) narrow tube (VD). nation. Planorbis b. YD. tentacles long and filiform. Trab. or oval. The walls of the preputium are thin but the walls of the vergic sac are thickened. As genus and subgenus. ANATOMICAL CHARACTERISTICS PLATE 1 GENITALIA. regularly increasing in diameter. 4) the preputium has two vertical pilasters and a large number of longitudinal muscular ridges (PL). There is one long and narrow retractor muscle (R^M) attached to the vergic sac near the junction of the latter with the preputium. II..7 mm. 1930. Planorbis 1931. Planorbis planorbis (Linn. narrow muscle attached to the upper part of the preputium have been called supporting muscles (SM) They are not attached to the columella muscle but to the fore part of the head.). Natur. Type 1931. 42. 521. the right side in advance of the left side. Planorbis (of Miiller 1774) R\as. wide. cylindrical. IV. Zool. . Gesell. Tropidiscus Soos. Papers.5 mm. 1). External appearance. narrow muscle may play a part in retracting the preputium after copulation.52 The Molluscan Family Planorbidae 1929. Kozlem. Barcelona. As genus. the foot short. The penial complex is elongated (fig. Shell (plate 77. 26-28) discoidal with few whorls. PD).5 mm. 210. and the vergic sac (VS) is short (0. Senckn. in length. covered for the most part by small pustule-like glands. long and 0. Animal. 2).7 mm. when removed from the shell the animal of Planorbis planorbis has a blackish or lead-colored mantle. and a pinkish ovotestis. As subgenus. A section through the verge and a . Moll. Terr. No. 25.. 516. The body is small. No.

Germain's figure (1931. A smaller tube. p. showing for the first time the separate prostate duct. The figure of the related species carinatus JMiiller (p. 5).Subfamilies. long. The ovisperm duct (SO) is divided into two external parts separated by the seminal vesicle. The genitalia of Planorbis planorbis have been figured by several European conchologists. 6). and Subgenera — Recent and Fossil 53 part of the vas deferens shows the sperm canal to be located in the center and to have a central exit (fig. The albumen'gland is large (3. The respiratory tube or pncumostome (PS) forms a . The prostate shows relatively the same number of diverticula. The duct of the prostate is shown as very short while in the Poland material the free portion of this duct is much longer before entering the vas deferens. through which it passes. 523. The seminal vesicle shows an enlargement. wide. Hermaphrodite Organs. extends back of the sperm duct at its junction with the oviduct and gradually enlarges as it enters the seminal vesicle. not rounded as was observed in all specimens personally examined. p. The pseudobranch (P. Beneath the albumen gland the sperm duct and the oviduct are joined to form the ovisperm duct (fig.) and wide. A small duct enters the carrefour (CF) which in turn enters the oviduct in front of the junction of that organ with the sperm duct. The absence of a distinct muscular ring or diajihragm is noteworthy.) and about as narrow as the sperm duct. fig. Genera. narrow duct about 1 mm. The spermatheca is shown on a long duct and the spermatheca is pear-shaped. 1). and attached to the uterus by a short.5 about mm. Soos's figure (1917. One portion. The spermatheca. long. The free oviduct (OD) is short (about 2 mm. Soos's figure (1935. 8). There appears to be some variation in this organ perhaps due to seasonal or sex condition. The penial complex in Soos's figure is not shown in sufficient detail. A. long and 2 mm. Germain's figure is like Soos's in this respect. The spermatheca is shown in a more elongated form. passes from the seminal vesicle to the ovotestis. about 3 mm. fig. but the position of the retractor muscle indicates a short vergic sac. 533) is in general like the figures in this work. Female Organs. L. but there is lack of detail. The sperraatheca 1. The nidamental gland (NG) is as long as the uterus and surrounds a large part of the oviduct (see fig. The rectum (R) pierces it after bordering the columella muscle and the anus has its exit at the lower edge (A). is shown of a pyriform shape and on a long duct. 9).5 mm. The uterus (U) is as wide as the vagina and is about 5 mm. about 2. 25) appears more nearly correct. The ovotestis (OT) consists of many clubshaped diverticula placed in a double row (fig. in length. Respiratory and Renal Systems. The ovotestis is buried in the cell-mass of the liver (fig.5 by 2 mm. The vagina (VG) is very short (less than 0. flattened.5 mm. 41) agrees in general with that shown in this work. It is greatly lobulated. (S) in the specimens examined is long and 1 mm. p. The large seminal vesicle is indicated by a swelling in the ovisperm duct. 24) given by Soos is more like the Poland specimens of planorbis personally examined. wide. but this organ is subject to more or less variation. globular or pear-shaped. long (SD).) and is a mass of small sacs (AL). The lower portion of the female complex is shown very much longer in Soos's figure than was observed in the Poland specimens. about 5 mm. however. 3) is leaf- shaped.

the lower or cutting edges V-shaped. in rows 1 to 156 the upper part of the central tooth measured 6 microns and the lower part of the base of attachment measured 12 microns in width. The jaw many is Geographical Distribution. Species Considered as Valid.54 The Molluscan Family Planorbidae is short V-shaped lobe to the right of the pseudobranch. In one membrane some of the marginal teeth (20) had the entocone split into two sharp cusps. There is no ridge on the lower side. 25-1-25. 25 km.5 mm. In rows 157 to 176 these measurements were 5 and 12 microns respectively. The ureter is short (1 mm. distributed throughout Europe from Sweden southward to the Mediterranean and from Great Britain eastward to western Russia. The pericardium is long-oval in shape. 6. the mesocone largest and longest. the fore part near the mouth somewhat tubular. the glandular portion occupying only about one-eighth of the length. 1 The kidney mm. large and can (plate 46. wide). The central teeth varied somewhat in width in the same membrane. 205. Asia Minor. the width being about one-fourth of the heiglit and there are four short. fig. subeciual cusps. 21) is horseshoe-shaped and is composed of small plates. The radula sac is distinct and rounded. (plate 50. The central tooth is bicuspid with a wide base of attachment. Digestive System. narrowly tubular and turns sharply to the left into the mantle cavity. fig. 7) formula is 24-1-24. 9) is very long and narrow (10 mm. the cusps large and spade-shaped. and 209. The outer marginals are very narrow. 10). For the above anatomical data eight specimens were examined. long). the blind sac about as long as the stomach. The stomach (plate 48. lateral teeth (1-10) tricuspid. tlie cusps long and spade-shaped. The stomach does not show distinct divisions into crop and gizzard. with four cusps which become subequal toward the outer margin of the radula membrane. It is also found in northern Asia (Siberia). Poland. Intermediate teeth (11-13) with the reflection reaching only about half way to the lower edge of the base of attachment. with four cusps. the ectocone split into two cusps. Marginal teeth (14-20) very narrow with the reflection placed high up on the tooth. and northern Africa. 2. No parasitic worms (cercariae) were observed in any of the specimens examined. cylindrical. This was rare and occurred only in atyjiical rows. For example. long and 1. The genus Planorbis. as herein understood.2 mm. southeast of Warsaw. Although many species now assigned to other genera were originally placed in the genus Planorbis there are now . the reflection reaching nearly to the lower edge of the base of attachment. long. A. 176. A cross section of the kidney at about midway of its length shows the lumen to be flattened and widened with a vein at each end (fig. and joined behind. In most membranes the central teeth were very uniform in width. The radula (plate 67. fig. wide. The salivary glands are much elongated. figs. 7) is elongated and pyriform in sliape. The intestine runs directly forward and does not make a backward loop as in Helisoma and some other groups. or 26-1-26 and the number of rows of teeth varied as follows: 175. about thirty on the upper or horizontal part. collected by Mr. It be greatly expanded. The buccal sac is elongated and shortly pyriform in shape. Jankowski from a pond in Swidry-Mate.

The genus Planorbis is peculiar anatomically in the form of the prostate with many single diverticula and in the comparatively small size of the vergic sac as compared with the preputium. the selection of which is governed by the rules of tautonomy (see H. 1800 (Vermes). as advocated by Kennard and Woodward (1924). Spirorbis Daudin. Soos 1935. The Helix corneus Linn.. Malac. The name Tropidiscus Stein (1850) is in use bv several European authors (Germain 1921.mxson. however. Dall 1870. Planorbis antiochianus Locard Planorbis aniotddi Germain Planorbis marmoratus Michaud Geological Distribution. Xaturwiss. 337. A.) Pln7iorbis carinntus Mi'iller now restrieted. Natursvis. l^lmiorbis sieversi Mouss.. p. selected b}' Gra}. Another group of authors have cited Geoffroy (Traite Coq. as genoty])e (Kennard and Woodward. have presented very convincing arguments in favor of retaining the Planorbis of this atithor. After reviewing the evidence very carefully there seems to be no valid reason why Planorbis Geoffroy. 12. Anzeiger Allg. Geoffroy 1767 is post-Linnean but this author was not in all cases binomial and is rejected by some students for this reason. 1820 (Gray. Planarbis atticus Bgt. Gesammt. giving ^Vlontfort 1810 (Conch. Oligocene to Recent time. Pilsbry and Bequaert 19271. is the genotype of Planorbariiis Froriep. Fischer and Crosse 1879) quote Ciuettard as authority ( 1847 Studer. is very small. should not be used. Treat. the author for this genus and the species for its type have been in dispute.. according to Wenz. Guettard 1756 is pre-Linnean and also non-binomial and would appear to be ruled out on these accounts. Tyjic Planorbis vulgaris Swain. Pilsbry and Bequaert 1927. 1767. II. 1924. Helix planorbis Linn.. Binney 1865. p. Some authors (Tryon 1872. Sci. 1884. Soos 1917. However. A fewhave used Guettard 1756 and ^liiller 1774 in combination with Helix corneus as genotype (Germain 1921. the genotype for both is Helix planorbis Linn. 1840. Baker 1902. 12.'Thiele 1931) but this name is a synonym of Planorbis. The verge. B. Fischer 1883. 9). The Planorbis of ]\Killer (1774) is preferred by some late writers (Dall 1905. Not . Schweiz.. p. Genus 1820. For many years. Other authors have cited the genus as of Miiller (Verm. Paris. 3te Jahrg. 152. Acad. Anisus ANISUS Studer. Baker 1931). also. p. having the same genotype. Hist. short. 1774) with Helix cornens Linn. Syst. 1767) with Helix planorbis Linn. who cite the Helix corneus of Linnaeus as genotype. Spirorbis Sw.. L. whether Geoffroy or Miiller is selected. It is probable that some of the sj^ecies included in this genus by paleontologists really belong to other grouj)s. p. Kennard and Woodward (1924. Ges. 1931). Baker. p. 91. 1756). and wide. Remarks. differing markedly in this respect (as far as known) from any other group of planorbes in Europe.SubjafNilies. p. Genera. and Subgenera— Recent and Fossil 55 The hut relatively few sjjecies which belong in this genus as best recognized of these are: Planorbis planorbis (Linn. 151. Xr. 1847) Type Helix spirorbis Linn. 270) as the first author to select a genotype. The nature of the genitalia and the form of the shell set this group apart as a distinct genus of the family Planorbidae. 1930).. as type (Dall 1905. 1926). 1806.

Y. op.. 1917. As synonym of subgenus Anisus.. Tr. 1926.^rtens. Cat. p.Y. 1883. by Lindholm in 1922 and Kennard and Woodin 1924. Fauna Flora Fennica.. pars. or Hesse 1909. Cent. Wenz.. Paraspira Wenz. 1926. Mollusk. Anisus Fitz. As genus. Not Diplodiscus Dies. 252..). Spiralina The Molluscan Family Planorhidae Hartmaxn. 64. Honigmann. 143.. As subgenus. As genus. 1905. Amer. Cien. Soc. Planorbis rotundatus Foiv. Russia.. Type evidently Helix . Musei Nat. Haldeman. ward 1902. 428. of Drap. p. op. 381.). p. Non Dies. Sj'st. p. Moll. Diplodiscus Westerlund. Paraspira Dall.). Uebersicht.. Archiv. p. 68. As subgenus. vortex Linn.). 1820) H\as. 1899. Zeitsch.. As subgenus. Type Planorbis 1921.. XIII. non-marine Moll. II. No type cited. Spiralina (von Martens. Monogenotype Spirorbis vulgarise^ Planorbis vortex var. 7. Type designated bv Lindholm. Anisus Studer. Gyrostoma Haas. 22. Not Plmiorbina Dall. Diplodiscus Westerlund. Type Planorbis spirorbis 1870. Playiorbis rotundatus Poir. Mus. Type Planorbis rotundatus Poir. Type designated by Lindholm. Schneck. As genus and subgenus. Lindholm. Spiralina Hartmann. As section of Planorbis. 58 year. Hungarici. p. Knietniewski 1897. 1929.. p. '= Planorbis leucostoma Millet. Trab. Type Anisus vortex (Linn. \). Fluv. Planorbis leucostoma Millet. Type Planorbis rotundatus Westerlund. Moll. 1530. Gray. As subgenus. Stein (part). p. f.. Type evidently Planorbis spirorbis (Linn. As subgenus. und Musch. 82.Vf first species (Anatomy). As genus. Anisus (Studer. 1921. 7.. 1899.-Ga.. Musee Zool. f. f. An. jip. op. 253. Genotype Spiralina vortex (Linn. Paraspira 1922. Nat. Trematodes. Ann. 1836. IX. Type designated by Pilsbrv 1934.. Ter. Dall. Gyrorbis L. Gyrorbis spirorbis 1909. Anisiis (S. Diplodiscus p. LV. Reg.). Rec. Spiralina von Martens 1899. p.. cit. "Not Gyrorbis Fitz.sterop. Syn. Lebend. 1897. II. 1850. Poir. = 1929. p. p. 76. 1885. Spiralina (Hartmann MS) von Martens. Spiralina 1923.. pp. 281. 81. XV. Akad. As genus. Sci. Germain. 45. 1529. cit. Mon. Soos. Type designated Planorbis vortex (Linn. 120. As sub- genus of Pla7iorbis. cit. 1926. Type Helix vortex Linn. As section of Planorbis. XVII. 1926. T^-pe by original designation Planorbis rotundatus Poir.. 1843. 1886. von M. cit. p. Lye. First species Planorbis vortex (Linn. p. Heft 6. Freshwater Univalve Moll. Hist. 507. Haas. Nat. Proc. pp. 1926. 423. 253. de Conch.). Kennard and Woodward. Germain. Ideogenotype Planorbis spirorbis (Linn.. Physidae. Biol. p. 296.). Berlins. Spiralina (Hartmann MS) von Martens. 14. Spirorbis Herrmannsen. p. Rad. op. 1899) vortex Linn. 1926. 1847. 180. (Linn. Planorbis spirorbis Linn. Barcelona. Tome XXIII.s'^. Molluskund..56 1840. Type Planorbis rotundatus Poir. 1820) Lindholm. 151. p. As genus. p. vortex (Linn. 43.. Man. 115.. (Hartmann) von Martens.). Gijrorbis Tj-pe Helix spirorbis Linn. pp.. Fossil. Gyrorbis Fischer. France. Jugoslav. 1833. 'Syst. 320. Str.=^ Planorbis leucostoma Millet. p. 352. Type designated Gyrostoma rotundatum Poir. As subgenus of Planorbis. Helix 1905. Not Gyrostoma Kirby and Spence 1828. Alaska Moll. 1923. Erd-und Siissw. 1836. Ann. 1923. XIII. Planorbina 1847. 49. Ind. XXI. Anisxis (Studer. Tropidiscus 1855. Archiv. cit. Zool. Brit. 34. Cab. Act. op.. As subgenus. Mai. Ideogenotype Helix vortex Linn. As genus. Soc. First species. Type designated by Lindholm Planorbis vortex (Linn. L'Acad. Ind. p. Mus. Gen. Gyrorbis Westerlund. 86. As genus. Moquix-Tandon. Lindholm.) Lindholm.' Xude name. 492.. p. Conch.).. Genotype Paraspira rotundata Poiret. p.. Gyrorbis Clessin. 395. Fauna Palaearc. Naturw. As section of Planorbis.). N.

spirorhis. but only about 0. long in spirorhis and leucostoinus. p. figs. Spiralina (Hartmann) von Martens. in septemgyratus fifty-seven. The prostate is long in vortex and septemgyratus and short in spirorhis and leucostomus. 12. fig. genus of A)iisus.. p. LXXXVI. plate 7. Anisus vortex (Linn. 2. convoluted. leucostomus. 480. narrow retractor nniscle and apparently no supporting muscles. which may be nearly twice as long in vortex as in spirorhis (figs. Type p. Ter. 1931. The preputium is always shorter than the vergic sac. As subgenus of genus A^iisus. in vortex). figs.). As sub- (Hartmann) Thiele. 1931.. narrowing greatly in diameter where it joins the preputium. Studer. Acad. Genera. 1820. The vergic sac is elongated. o\i. iv teil. IVpe Planorhis Icucostoma Millet. Anisus (Studer. and. cit. The preputium is cylindrical. Male Organs (plate 6. Fluv. II. .. figs. The vas deferens is shorter than the sperm duct by about half its length. von Type probably intended cit. 3 vortex. fig. SS. 1931. cit. Sci. 3 mm. is a narrow tube. (Linn. Cross sections through the oviduct and prostate show the separation of the two ducts (plate 6. cylindrical. op. As genus.). The prostate duct varies in length in the different species. and in leucostomus over twenty. and Subgenera — Recent and Fossil 57 (Hartiuann. As genus. fig.Subfa/inlies. 29. usually swollen at the upper part. in leucostomus. 12. 6. Pilsbry. It is about 3. 1935. Spifdliiia 1>. XXXII. plate 7. 1931. 20-22.. in spirorhis. ANATOMICAL CHARACTERISTICS PLATES 6-7 fig. The diverticula of the prostate vary in length and diameter as shown in the figures (plate 6. aperture small. plate 7. 6). As genus. and are placed on a duct separated from the sperm duct which joins the latter some distance above the vagina to form the vas deferens. as usual. The diverticula are sac-like. Spiralinn 1934. Anisus (SS) Thiele. p. septemgyratus.) ) discoidal with many tightly coiled whorls very slowly increasing in diameter. Proc. . op. in vortex being about sixty. septemgijratus) Seminal vesicle (SV) long. about three times the diameter of the ovisperm duct. in spirorhis about twenty. long in vortex. GENITALIA. Moll. designated by Gray in 1847. Subgenus Ty]:)e ANISUS Helix spirorhis Linn. cylindrical. modified in carinate species by the peripheral carina. 62. As subgenus of Anisus. Kozleni. As subgenus. 1820) Thh^lk. fig. long in spirorhis. Soos.). Nat. Gekmaix. Anisus spirorhis (Linn. 1899. 7 mm. 526. France.2 mm. Animis Studer. 16-18. 1 and 13 on plate 6). It is about 1 mm. The number of diverticula vary correspondingly. rounded or ovate. Externally resembling that of Planorhis. 1935. to be Planorhis vorlex (Linn. pyriform or club-shaped. 2 mm. 29. 8 spirorhis). Animal. Type Planorhis vorlex p. and 4 mm. 481.. in septemgyratus. A7iisus leucostomus (Millet). and leucostomus. fig. The penial complex is greatly elongated in all species of the genus Anisus. fig. 19. Anisus Handbuch. Martens). whorls rounded or with a sharp carina at the periphery. Shell (plate 77. long in vortex. AUat. There is one long. Soos. Sperm duct (SPD) very long and slender (5 mm. p.5 mm. Helix planorhis Linn. 481. 2. all of the whorls visible on both sides of the shell. 1820.

horny stylet. but rather short and wide in leucostomus. In several specimens of septemgyratus the spermatheca was cylindrical and acutely pointed at the summit. long in spirorbis. 4. plate 7. 2 and 6. bulbous at the upper part where it is attached to the verge. the spermatheca is short and bulbous with a short. to be small or large. fig. however. spirorbis. It is relatively thin and is capable of considerable contraction and expansion. fig. Between the preputium and the vergic sac there is a thick muscular ring which shows as a swelling on the exterior surface of the upper part of the preputium. 6. and septemgyratus but large and swollen in leucostomus. The spermatheca in vortex. fig. 6). The spermatheca and its duct are over 1 mm. The albumen gland is elongated. The stylet is sharp and needle-like. The ovotestis is made up of many club-shaped diverticula arranged in a double row on the ovisperm duct (i)late 6. AL). 5. figs. figs. The sperm canal turns to the side just above the stylet where it has its exit on the side of the verge (plate 6. varying in this respect from Buchner's figure. narrow. thick spermathecal duct (plate 7. and spirorbis. 2. fig. at the end of which is a small.58 The Molluscan Family Plauorhidae Internally the penial complex has certain distinctive features. Several diverticula of vortex had ripe ova. Possibly changes due to breeding condition may be the reason for in fig. fig. 12). 5). fig. 4. fig. which causes the hole or foramen in the middle. long). The verge of vortex is much narrower than that of any other species examined. . plate 7. spirorbis (3 mm. OT. The nidamental gland is long and cylindrical in vortex. The oviduct shows the same differences. vortex. long) and of larger diameter in leucostomus. 10 . as shown on plate 6 ispirorbis) The preputium has two vertical ridges or pilasters. cylindrically pyriform in shape above and tapering rapidly below the middle of its length to form a slender tube. There is a fine groove extending along one face of the stylet (plate 6. 2). 10. In those specimens of the four species examined the ring has been widely open and the papilla has been extended below the area of the ring (plate 6. 1 on plate 5. and somewhat arched (plate 6. and septemgyratus. The oviduct is always of greater diameter than the sperm duct (see figs. but it is there shown in section as almost closed and is placed below the thick papilla. The vagina (V) is very short (less than 0. is much elongated and cylindrical and septemgijrntus with a spermathecal duct about as long as the spermatheca (plate 6.3 mm. 2. figs. plate 6). The uterus (U) is long and cylindrical in vortex. 5). this difference. as viewed from above. A large papilla (PP) is suspended between the walls of the muscular ring. Female Organs. fig. 12. being long and cylindrical in vortex. fig. spirorbis) The papilla is larger and somewhat differently shaped in the figures on plate 6. 12. The ovisperm duct is divided into two parts separated by the . but is short (about 2 mm. One specimen contained a large spermatophore (plate 7. Buchner shows this ring (Ringwulst) in his fig. septemgyratus.) in all species. Hermaphrodite Organs. In leucostomus. 7. the walls of which are very thick. The verge is long. 2. With the exception of the 'Ringwulst' the penial complex of the four species of the genus Anisus agrees with the description and figure of Buchner. 9. fig. Two diverticula of leucostomus are shown on plate 6. spirorbis. fig. plate 7. Below the large muscular swelling there is a second smaller ring which is here called the diaphragm (D). spirorbis and leucostomus). but less than a millimeter in leucostomus. plate 7.

the glandular portion occupying only a small part of the entire length. The portion of the ovisperm duct between the seminal vesicle and the oviduct is a long. The pericardium is very short as compared with total length of the kidney. Central tooth with two spade-shaped cusps reaching nearly to the lower edge of the base of attachment. 539). In vortex (fig. 527. about as wide as high at first but becoming narrower toward the marginal teeth. but in vortex. The ureter is very short and is sharply bent upward into the mantle cavity. Lateral teeth (1-10) tricus]')id. wide. with the reflection short and wide. In the first two species the enlargement is of the same diameter as the seminal vesicle. tafcl 4. fig. and spirorbis there are small pustule-like bulgings which ai)i)ear to be glandular extensions of the seminal vesicle. fig. The side jaws are also fragmented. In these species the kidney is somewhat triangular in shape to fit the sharp angle caused by the peripheral carina of the shell. which is copied by Simroth. the reflections very short and wide. Germain's figure of rotundatus (=leucostonius) is similar but lacks detail (1931. Cross sections show no evidence of a ridge. In leucostomus (fig. SubjaniiUes. The kidney 11 spirorbis. INIarginal teeth (12-15) narrow. The genitalia of the four species here recorded are in the main similar Buclmer's fig. wide but in vortex it is 12 mm.3 mm. Respiratory and Renal Systems. The pneumostome is cylindrical and in the specimens examined is rolled into a cylindrical tube with a large cleft extending its whole length. 11. 25) is similar to that of Planorbis with about twenty plates on the upper part. septemgyratus. 2. (plate 47) is very long 1 leucostonius) and narrow. p. and plate 7. figs. buccal sac. 13) it is 5 nun. four-cuspid. Genera. 6. long and over 0. 6 to vortex. 6) but is of small diameter in spirorbis and vortex. The jaw (plate 50. In a cross section of spirorbis (fig. narrow. fig. 16) the kidney is seen to be very wide and much flattened with the veins near the middle of the lumen. and flattened in all species examined. 14) the kidney is as thick as wide and the veins are placed at the base of the rounded lumen. four-cuspid. The numbers of rows of teeth vary from 185 to 195. slender. The pseudobranch is large. In spirorbis (fig.. 15) the kidney is wider than high with the veins lower on the lumen. fig. The kidney of Anisus does not differ in any essential way from that of Phinorbis. and general digestive system are similar to these organs in Planorbis. It varies in length in the different species (plate 6. The radula (plate 67. Digestive System. pp. as is also the figure of vortex (p. Soos's figure (1917. There is but one intermediate tooth (11). long and about 0. spirorbis) formula is 15-1-15 or 16-1-16.3 mm. 50) is like those in this work. and Subgenera — Recent and Fossil 59 yeiuinal vesicle. smooth tube in leucostoinus. The stomach. The formulae for the other species examined are as follows: 16-1-16 with 130 rows sejptemgyratus 16-1-16 with 142 rows vorlex 16-1-16 with 130 rows leucostomus . It is pierced by the rectum which is long and narrow. as in the ii'enus Planorbis. fig. The prostate is shown too long as compared with the Poland specimens personally examined. The fourth cusp appears as a splitting of the ectocone. The portion between the seminal vesicle and the ovotcstis is enlarged in septemgyratus and Icucostomus (plate 7. 53). of which it may be a part.

In Planorbis the vergic sac is only one-fourth as long as the preputium while in Anisus this organ is always longer than the preputium and the verge is long and narrow and bears a stylet at the end. The last two species are now members of the genus Drepanotrema. Soos. Poland: vortex. 68). 286) listed Spiralina Hartmann in the synonymy of typical Planorbis without comment or question. Species Considered as Valid. septemgyratus. used by Germain. Poland. In 1899.) centrogyratus (West.) compressus (Michaud) Anisus Anisus Anisus Anisus Anisus Anisus Anisus johanseni (Mozley) perezi (Dupuy) (Germain) jragilis (Millet) paravortex (Ping and Yen) millcti charteus (Held. Anisus Anisus Anisus Anisus Anisus Anisus Anisus Ariisus spirorbis (Linn. Kennard and Woodward. of Warsaw. the formula of which is 25-1-25. spirorbis. Mai. Three examples from turfy meadow in Drewnica. The following species are understood to conform to the concept of Anisus. which is absent in Planorbis. thus. One species is reported from China {paravortex). Gen. Thus the distribution is about the same as that of Planorbis. plate 6. a nude name published by Hartmann in 1840. The shell is always flattened and many-whorled but from the shell alone the two groups could scarcely be separated generically. Poland.) septemgyratus (Ziegler) leucostoynus (Millet) vorticulus (Troschel) villai (Adami) ressmannianus (West. 320). Poland. But in 1847 Herrmannsen (Ind. as . northeast of Warsaw. The proper name for the group of European planorbids typified by Planorbis spirorbis. The presence of the muscular ring and papilla in Anisus is also another striking difference. The seminal vesicle also differs markedly in the two genera. 4. Anisus). 12 km. Jankowski. The genus Anisus is distributed throughout Europe from Great Britain to Russia and Siberia and southward to Italy and Algeria. leucostomus. and others. plate 1. but the teeth are similar in form and number of cusps. and leucostomus has been in considerable doubt. and by Kennard and Woodward in 1924 (p. p. Poland. Planorbis. von Martens used the name for the European Planorbis vortex and the American Planorbis cultratus and sumichrasti. A. II. The name most commonly found in the literature. Anisus has fewer teeth in a row than Planorbis.) Geological Distribution. northeast of Warsaw. This distribution may be enlarged when other si)ecies arc examined anatomically. 10. Eight examples from stream in meadow at Jab-Tonna.) vortex (Linn. Warsaw. Thus the type of Spiralina becomes Planorbis vortex and this species was formally designated the type by Lindholm in 1922 (p.60 The Molluscan Family Plauorbidae The teeth are all very uniform in form. Remarks. Three sj^ecimens from small ponds in Bielany Park. with fig. Geographical Distribution. northeast of Warsaw. This genus begins about the middle of the Miocene period. all received from Mr. Lindholm. is Spiralina. Eight specimens from a trench in Struga. For the anatomical information herein presented the following material has been available. vortex. 4 km. 18 km. The group of European planorbids under the name Anisus differs from the genus Planorbis notably in the form of the penial complex (compare fig.

Gyrorbis. pp. 62). killing the p. As subgenus of Anisus. The group is anatomically unknown but the shape of the shell would indicate that it should be grouped with Anisus. spirorbis' is valid for Anisus only if the identity of spirorbis Studer and spirorbis Linn. The writer agrees with Pilsbry that. j). versely costate. Russia. Heft 257.) strauchianus Shell (plate 76. fig. genot. But this name is H. the ciuadrate. As subgenus of Anisus. B.Subfamilies. Clessin's figure (copied from Conch. It is a question whether this group should be separated from Ani.' Thus no subsequent use preoccupied by Diesing in Trematodes. the aperture Animal. as noted in the list of synonyms of Anisus. A more serious question relating to Anisus is whether a generic name is valid when admitted by its author to be for a group formed by the union of two prior genera. Other names have been proposed for this European group and would be available but for previous use in other groups. 2. Teil Clessin. 48L Type A. Costorbis COSTORBIS Lindholm. and Subgenera — Recent and Fossil 61 49) 'effectually clothing and simultaneously of this name is permissible. Archiv. 1820 and gave some very good reasons for accepting this name for the group tyjufied by Planorbis vortex and spirorbis. 5) is reproduced on plate 76. since it was proi:)erly clothed as a genus by Gray in 1847. Sci. differing principally in the costate sculpture. 1931. V..s is similar in form to that of Anisus septongyratus. fig. In 1934 (Proc. Unfortunately Opinion 9 of the International Commission on Nomenclature is not explicit on this point. Geographical Distribution. Acad. Subgenus Type by 1926. 1926 original designation Planorbis strauchianus Clessin Lindholm. 3. Wuestia. in (C. Anisus is used by Thiele in his Handbuch. . and containing nothing else. The original description of the shell appears on page 204 of Clessin's work. The best interest of nomenclature is served by the use of Anisus rather than by coining a new name. 3). and Gyrostoma are thus affected. 253. as type. name Spiralina. XVII. Mono- 193L Costorbis Thiele. Pilsbry revived the early name Anisus Studer. I see no sufficient reason for doubting it. Species Considered as Valid. Cab. this being the first species listed. Genera. Handbuch. 58 year. The shell of strauchiami. plate 31. Diplodiscus. Spirorbis Swainson. Differing from Anisus being heavily trans- body whorl angulated above and below. p.vpe Planorbis strauchianus Clessin. fig. Remarks. Phil. fiir Mollusk. In this connection he remarks Gray's type designation of 'H. The only species definitely referable to this subgenus is A7nsus (Costorbis) strauchianus (Clessin). accepting Gray's subsec^uent designation of Helix spirorbis Linn. Baker remarks (1930.. since the name Anisus has had wide use in the literature it had better be retained for this group.. is admitted. but as Anisus has been widely used it had better be retained. Dall (1905) used the name Diplodisciis Westerlund and cited Planorbis vortex as type.

Bathyomphalus Clessix. 58 year. AUg. France. Not difi^ering in general characteristics from Anisus. Polygyrus Gr. Moll. Zool. As genus. Nat. Museo Cien.) and composed of forty-eight to fifty relatively short and wide GENITALIA. Teil 2.arpentier. Reg. 86. 1929.. Mai. p. Mus. p.ard and Woodward. Kozlem. Hist. 65. Genotype Bathyomphalus contortus (Linn. (Agassiz MS) Charpentier. coJitortus (Linn. Cab. Akad.^in. 23-25).. Type Planorbis Linn.. 82. Type Bathyomphalus contortus (Linn. Archiv.). 83. Pars 22. p. 45. Bathyomphalus Lixdholm.5 mm. As subgenus. Bathyomphalus Westerluxd. Lye.. p. Man. 1633. As subgenus of Planorbis. 181. Proc.). 507. Rad. 120. V. Musei Nat. Hist. X.\y. No tyjie cited. II. Planorbis contortus (Linn. p. Small. Conch. Cat. 1902.. 1926. p. 481. Bathyomphalus Dall. j.). Gen. p. 10) Seminal vesicle (SV) very long of larger diameter than the ovisperm duct with many small tuberculous glands protruding from the surface.). 1931.). II. 352. Suppl. Bathyomphalus Germ. Bathyomphalus Kexx. p. XV.) cited. 1886. 253. British Non-marine Moll. Fauna Palaearc. 2. 1917. Type evidently Planorbis contortus (Linn.). Syn. Bathyomphalus H. As genus. Linn.. Type evidently Planorbis contortus (Linn. Handbuch. Hungarici. As contortus (Linn. 34. 68. Bathyomphalus Fischer. Ind. As genus of Planorbis. Male Organs (3. 1837).). Barcelona. Animal. (fig. or Beck. Bathyomphalus Westerluxd. Alaska Moll. 1931. section of Planorbis. France.) (not Berlins. Type Helix contorta Linn. p. 520. 151. 1870. As subgenus of Planorbis. Schneck.. pp. p.).62 Subgenus 1837.). As genus. Ind. Type Planorbus contortus (Linn.). 1855. Ann. p. 1926. As genus.). Jugoslav. Type Anisus (B. As subgenus.. (MS) Ch. As subgenus of Anisus.. designated by Herrmannsen in 1852 Schweiz. increasing very slowly in diameter.. 1837 Type Helix contorta Bnthyomphalus Agassiz Gesell. ANATOMICAL CHARACTERISTICS PLATE 3 . II. Type Planorbis contortus (Linn. Ann. Fossil. 1921. Conch. Neue Denkschr. p. Rec. equally visible above and below. 7. Bathyomphalus Thiele.. Nat. of Say. As subgenus of Planorbis. Bathyomphalus Soos. Moll. p.\ll. not present in Anisus. ultradextral with the whorls more nimierous and more closely coiled than in Anisus. but the spire whorls (left side) forming a wide depression. Type Planorbis Type Planorbis As sub- Bathi/omphahis Herrm. contortus (Linn. IX.). Type Planorbis contortus (Linn. Allat. 16.. Prostate long (about 3 mm. Bathyomphalus Soos. 1852. 383. As subgenus. Bathyomphalus Germ. fiir Mollusk. p. Bathyomphalus Moquin-T. . 443. Heft 6. 1935. XXXII. und Musch.Y. 30. 1885.) contortus (Linn. Bathyomphalus D. 1850. Bathyomphalus Wenz. Leben. no. XII. T^-pe Planorbis contortus (Linn. Trab. 1818. XXI. genus. As section of Planorbis.. p.aas. 1883.\ndon. figs. Discoidina Stein. Tyi)e Helix contorta 1905. a false umbilicus. The Molluscan Family Planorbidae BATHYOMPHALUS I. 1923. IV. p. Monogenotype Planorbis contortus (Linn. XVII. 142. Nat.annsen. pp.). Soc. Shell (plate 77. Type Helix contortus Linn. 20. Terr.. 1847. p.

is the same as in A?nsiis. pp.5 — Recent and Fossil 63 (fig. long. The nidamental gland (XG) is long (about 3 mm. p. The spermatheca is cylindrical.) and wide. The prostate shows the great number of short diverticula. Respiratory and Renal Systems. fig. smooth tube (SO).3 mm. The i)rostate duct is a narrow tube about nnn. The stomach (plate 48. There is no ridge on the ventral side of the kidney but there is a rounded bulging on the left side that does not appear on is the right side. The diverticula are long and cylindrical and packed closely on the ovisperm duct (plate 3. The oviduct (OD) is long (about 4 mm.Subfamilies.) and is a small. wide) with a sharply reflected ureter. 8) is greatly elongated. with a large blind sac. The penial complex (fig.5 mm. long. The ovisperm duct lies at each end of the seminal vesicle. long from the prostate to its junction with the sperm duct to form the vas deferens (fig.). The pseudobranch (jilate 3. Hermaphrodite Organs. 0. (about 2 mm. .3 mm. long. 2).3 mm. 0. PRS. 20) shows a laterally jiyriform lumen with the veins at about the center of the section at each end. vergic sac 2 mm. but is smaller and narrower. PD). of about the same length in each division. 7). fig. The kidney (plate 46. wide) and narrowly cylindrical. 8). pear-shaped. 10. OT. 8. There' is one long and narrow retractor muscle attached to the summit of the preputium. fig. The pneumostome (PS) is larger than in Anisus but is otherwise similar. 19) is long and narrow (7 nnn. The uterus (U) is long (2 mm. Soos's figure (1917. the anatomy of the penial complex .) narrow tube which enters the vergic sac without enlargement. Genera. 47) is similar to the ones here presented and is somewhat clearer than those of Germain.) The narrow and cylindrical vergic sac is greatly contracted for about a third of its length near its connection with the preputium. Female Organs. fig. while in the specimens examined it was long and narrowly cylindrical.) and of larger diameter than the uterus (about 0. Internally. The penial complex (p) shows the vergic sac as much swollen. the spermatheca is on a long duct. SPD). 10. P) about as in Anisu>i. The relative lengths of the female and male systems is shown much shorter than was observed in the Poland specimens.5 mm.2 mm. and the penial complex is like the Poland specimens here figured.) and narrowly cylindrical. The vagina (V) is very short (0. The spermatheca in both figures is the same. This junction is placed higher uj) on the uterus than in Planorbis or Anisus. The vas deferens is a long (4. The ovotestis is like that organ in Planorbis and Anisus. A cross section of the kidney (fig. fig. two-thirds as long as the stomach. 1 mm. and Subgenera diverticula 1. Digestive System. long. PD. 532) does not agree with the Poland specimens in all respects. attached to the vagina by a narrow spermathecal duct a little more than 1 mm. The albumen gland is about the shape of that organ in Anisus (see plate 7. 521. In the natural position the prostate lies over the sperm duct and until separated from this organ the prostate diverticula appear to be directly attached to the sperm duct (see fig. the prcputium (PR) being a little more than half the length of the vergic sac (preputium 1. The figure in Germain (1931. Details of the prostate and ovotestis are not shown. 5. fig. 9) is very long and narrow.

the shell is distinctive and the name has been in use for so long a period that it should be retained as a subgenus of the old group Spiralina { = Anisus). Poland. Geological Distribution. This group extends throughout Europe from the British Isles and Portugal eastward to Russia and Siberia. fig. Beitrage zur Fauna unci stiatigraphischen Lage der pannonischen Schichten in der Umgebung des Balaton-Sees. northeast of Warsaw. Marginals (1215) with five cusps are like the intermediate teeth. The pyloric portion of the stomach appears to merge with the intestine. The genitalia strongly resemble those of Anisus septemgyratus. From the Miocene period to the Recent period. The radula (plate 68. placed high up on the reflection. but there is no bending backward with a posterior loop as in the Helisoniatinae. the aperture contains three lamellae. Geographical Distribution. whorls relatively few. Genotype Planorbis krambergeri Halavats. As in Anisus. However. The jaw is like that of Planorbis planorbis. For the anatomical data herein presented. p. as far east as Kamchatka (Eyerdam. a turfy ground 4 km. figs. Germain states that it is absent. ODONTOGYRORBIS Lorenthey. and pyloris. p. Shell (plate 76. Art. collected by A. Cat. The inter- mediate teeth (9-11) differ by having the reflection placed higher on the tooth. 3. Southward the distribution includes the countries bordering the Mediterranean Sea and the Caspian Sea. nearly reaches to the lower margin of the base of attachment. 2). at Drewnica. 1538. and Anisus (Bathyoniphalus) dispar (Westr. Very small. flattened. It should be noted that the radula formula is the same as in Anisus and different from that of Planorbis. Remarks. As subgenus of Paraspira. the reflection very high on the tooth. there is no sharp division into crop. 20. left side almost flat. 14-16). Pars. Odontogyrorbis Wenz. with plate 47. Resultate d. right side slightly concave. 22. (Odontogyrorbis) krambergeri Halavats. Fossil. with four cusps. closely coiled. I. 1. figs.. The kidney shows some differences in cross section (compare plate 46.64 The MoUuscan Family Planorhidae intestine crosses over the oesophagus and extends toward the forward part of the body.) of Europe and Asia. 121. four specimens were dissected. Lateral teeth (1-8) are tricuspid with a small additional cusp above the ectocone. fig. In anatomical features Bathyoniphalus so nearly resembles Anisus that there seems little reason for recognizing the group as distinct. discoidal. 1906 Halavats original designation Planorbis krambergeri Odontogyrorbis Lorexthey. The buccal sac is as in Anisus. wissenschaftl. Ajiparently but two species are included in this group: Anisus ( Bathyoniphalus) contortus (Linn.) of Sweden and Russia. Erforschung des Balatonsees. gizzard. 1) formula is 15-1-15 with 140-145 rows of teeth. Genotype Paraspira 1923. Genus Type by 1906. 1. increasing very slowly in diameter. each side showing all of the volutions. the ectocone split into three cusps. Central tooth bicuspid. 1928). Jankowski. one on the base of the lip and . The internal structure of the penial complex is similar. The outer marginal (15) is very narrow. the superior portion having The about twentv plates. Species Considered as Valid.

no. Syst. Gesell.. Planorbis costulatus Krauss. op. 1870. 1857. 1883. Brown. 49. II. Schneck.. p. 234. Xat. Allij.Subfaf/iilies. Gyraidus W. Stein. Gyrulus Gr. Genera. Giraulus Moquin-Tandon. whence the generic name Odontogyrorbis. (6). cit. 2nd Ed. FAxi unil Siissw. This group is noteworthy for the presence in the aperture of three folds or plicae. In sj'iionj-my. Xon I. Turtons Manual. all but one of which are extinct. Lye.. Ann. Monotype Caillaudia angulata Bo\\rg. 1857. 124). As . p. 1) were broken and did not have the folds in the aperture. p. iipjxT part of the aperture.). (Tableau). and Subgenera — Recent and Fossil of the aperture 65 is two on the thickened. Caillaudia Bourciuign. While the shell does resemble the type of Paraspira [Planorbis leucosfonms ^Millet) no member of the genus Anisus has as yet been recorded from the recent fauna with a dentate aperture. Zool. op. Man. Binney. Odontogyrorbis krauibcrgeri (Halavats) the only si:)ecies here ascribed to the genus Odontogyrorbis. Trochlea 1844. 1837 Designated by 1827. Haldeman. Sci. 351. X"^at. 423.^. Type Planorbis albus Miiller. p. p. Berlins. exphm. Gyraulus Hartmaxx. 1837. The first species originally included. cit. 2). Conch. Britain. Schweiz. Jour. fig. Leb.. 1538) made Odontogyrorbis a subgenus of Paraspira {= Anisus). 1841. fixed by Gyraulus Moquin-Tandon. 1840. G. und Muschl.. Hist. 1850. V. Lorenthey obtained specimens with the lip entire (plate 76. Gyraulus (Agassiz MS) Charpentier. 234. p. (vide Pilsbrj' and Bequaert. X"at. Nnutilina 1855. p. Gyraidus H. Xeue Denkschr. Brown. As subgenus of Planorbis.. Wenz (1923. Planaria (part) Miiller {^Planorbis hispidus Drap. Gaster. Euroji. & F-W Sh. Hist. Gyraulu. p. X. Remarks. p. and Nagyvazsong. In general form the shell is related to Anisus vortex (Linn. 89. but is very much smaller. Type Planorbis albus Miiller.) Dall in 1870. p. Lower PHocene. = . Ann. 1. p. XV. in fact. 1865. II. 48. Fonyod. der Schweiz. 1927. Gattun. fig. 438. Kenese. Halavats original specimens (plate 76. For the present Odontogyrorbis should be treated as an extinct genus related to Anisus. 21. 128. ed.artmaxn. Some of the supposed extinct species are closely related to their recent allies and might be considered geologic variations of the recent forms. Xew name for Planaria. XLII. 216. It is of interest to note that the Pliocene planorbid fauna of the Lake Balaton region consist of sixteen Tyjiographical error for Gyraulus. X. 1883. 99. 1855. Species Considered as Valid. Error for Gyraulus.subgenus of Planorbis. 80. p. L. is Genus Type GYRAULUS Planorbifi albii. The whole edge Geological Distribution. Uebers. Y. Xo type cited.. Soc.'i D. Hungary. As genus.\y. Tyjie Clessin (1886) Planorbis albus Miiller. Amer. As section of Planorbis. 507. X'o type cited. Conch. IX. France. p. and Golze 1782 (Vermes). p. Miiller 1886 2.\ll. figs. Moll. Illust. Gyraulus Gray. plate LI. ancestral species. A. the Pleistocene fauna contains fifteen species and races all of which belong to the recent fauna. Gyraulus Fischer..s (Agassiz :MS) Charpentier. As synonym of Gyraulu.

As subgenus of Planorbis. IV. 33. Gyraulus Martens. Pars 22. Type Planorbis albus Miiller. XXII. figs. Museo Cien. Mus. 2nd Teil. p.. 1931. Gyraulus Wenz. 1928. p. Pub. 118. As subgenus of Planorbis. 6. India. S. p. Biol.. C. Type Planorbis XIII. British Non-Marine Moll. Gyraulus Hannibal. 1915. XVU. Akad. As subgenus or Planorbis. As genus.. p. IV. the aperture oblique. Moll. p. Rec. albus Miiller. Cent. equally visible above and below. Mus. Tyi)e Planorbis albus Miiller. II. Ann. Miscel. Moll. Gyraulus Westerluxd. p. & S. Miiller. Hist. Trab. Example Planorbis deflectus Say. p. Type Planorbis albus Miill. Nat. Hungarici. 151. 144. As 1886. Mai. Tyiie Planorbis albus Miiller. p. Type Planorbis albus Miiller. 1929. As genus. Gyraulus F. Univ. X. Type Pkuiorbis albus Miill. 1917. albus Miiller. As subgenus of Planorbis. Miiller.. p. 1918. Type Planorbis albus Miiller. As subgenus of Planorbis. As genus (genitalia and radula). Mus. Gyraulus Pilsbry and Bequaert. p. Baker. Type Planorbis albus Miiller. II. Handbuch. Gyraulus Germain. Gyraulus Soos. Amer. p. 481. As subgenus of Planorbis. 364.. Bull. Baker. Gyraulus F. Rad. 1921. Musei Nat. p. Zool. Shell (plate 77. Type Planorbis albus Miiller. p. Ill. Type Planorbis albus Miiller. Gyraulus Dall. evidently Planorbis albus Miiller. 1905. Fauna Brit. p. Barcelona. No. As genus. 98. 1926. Gyraulus Thiele. Baker. Type Planorbis albus Miiller. Rec. Arts. Type Planorbis albus Miiller. As subgenus of Planorbis. The Molluscan Family Planorhidae Tryon. 1-3). 481. 75. pp. 124. XXI. Acad. 1922. Ind. Wis. Terr. Moll. As section under genus Anisus. pp. Conch. Conch. 94. p. Chi. Trans. p. I. 124). 1931. Mus. Ind. Mich. XXIV. 1899. Gyraulus Annandale.. 83... 154. 86. Cat. Ind. Cab. 12. 1935. As section of Planorbis. As subgenus of Planorbis. C. Gyraulus Clessin. 1902. 1912. the . 533. et Fluv. usually rapidly enlarging in diameter. p. 30. XVHI. As genus. (see Pilsbry and Bequaert. ultradextral. As genus. Sci.. 1919. Gyraulus Soos. Gyraulus H. Rec.. 2nd Teil. As subgenus of Planorbis.. Gyraulus Preston. Gyraulus Westerlund. V. 297. 1931. London. France. As genus. Handbuch. LIII. Jugoslav. Gyraulus Annand. Type Planorbis albus Miiller. T. Amer. Allat.. 204.. Alaska Moll. albus As genus. Type wrongly given as Anisus (C. Small. C. p. pp. p. Type Planorbis albus Miiller. Moll. Gyraulus Germain. Type Planorbis albus 1902. Proc. 67. Type Thiele. 1927. As section under subgenus Armiger which is a subgenus of Ayiisus. Soc. As subgenus of PUniorbis. No type cited. of comparatively few flattened \vhorls. 8. XXXII. 1927. Area. = Gyraulus F. F. Wis. 1540.. G. pp.66 1884. XV. p. W.\as.) letourneuxi Bourg. 54. Nat. Kozlem. Fos. 1923. 384.. No type cited.. Gyraulus Walker.sil. subgenus of Planorbis. Mus. Type Planorbis albus Miiller. 121. Moll. Gyraulus 1885.. Gyraulus Kennard and Woodward. Syn. Genotype Gyraulus hispicbis Drap. 106.. 361. As subgenus of Planorbis. 52.\le and Prashad. the periphery is usually median and in some species carinate. No type cited. 392. Caillaudia Subgenus GYRAULUS SS.vpe Planorbis 1926. body whorl at aperture usually somewhat deflected. Fauna Palaearc. p. Reg. As subgenus of Planorbis.

shows through the shell. narrow duct about half as long as the spermatheca (SD). and a dependent papilla (PP). narrowed at the lower end where there is a stylet (ST). however. The penial complex (fig. the vergic sac (VS) somewhat shorter than the preputium (PR). The verge (V) is elongate-pyriform. A cross section through the prostate near the anterior end shows the sperm duct behind the prostate (fig. In general form the animal does not differ greatly from the figures of Tropicorbis on plate 78.) and is of the same diameter as the sperm duct which it joins to form the rather short (1. His figure.2 mm. OD) is a short and narrow tube. one at each end of the seminal vesicle (fig. S) joined to the vagina by a short. The ovotestis (OT) consists of a double row of pyriform diverticula attached to a large ovisperm duct (fig. The stylet (fig. External appearance (plate 70. The figures on plate 14 agree with that of Germain (1931. p. 7. Prostate (PRS) relatively long (1. 11). 5). and Subgenera shell in several species is — Recent and Fossil 67 covered with short. a trifle more than 1 mm. 6. Seminal vesicle (SV) less than 1 (fig.^) .) vas deferens. tion to form the ovisperm duct. The vagina (VG) is very short and wide. The spermatheca about 0. GENITALIA. is is elongate-pyriform. 7. over which it lies (fig. 1) the preputium has thick walls. 10).5 mm. The free portion of the prostate duct is very short (0. Soos's figure (1917.5 mm. fig. and the method of junc- Female Organs. the prostate. a muscular ring or diaphragm (D). which forms a loop. At the posterior end of the ovotestis the diverticula are crowded together (fig. the sperm duct. SO).) diverticula attached to a separate prostate duct which is more than twice the diameter of the sperm duct. ANATOMICAL CHARACTERISTICS PLATE 14 {Gyraulus albu. The ovisperm duct (SO) is divided into two parts. fig. SPD). long. The head bears the long. dagger-like with a ridge extending down the center. gland are hidden in the figure by the large prostate. 55) is and its . ST) is dark brown. looped beneath the prostate. 9. Male Organs mm. is not very clear. long. The vas deferens is not enlarged as it enters the vergic sac. The narrow duct of the albumen gland connection with the oviduct is shown. The free oviduct (fig. 6. There is one long retractor muscle (R^I). (fig. p. The vergic sac has thin walls which increase in thickness toward the preputium. the oviduct. 8. Several diverticula had developing egg cells. 545). The albumen gland is squarish or oval in shape and is composed of large gland cells. No pigmentation.5 mm. Hermaphrodite Organs.Suhjamilies. 2) is cylindrically elongated. 7) composed of a few large diverticula or glands three or four times the diameter of the ovisperm duct. however. Sperm duct about 2 mm. Genera. Figure 6 shows the relationship of the nidamental gland diverticula. Animal. of small chameter. The opening of the sperm canal (SC) turns to one side above the stylet and has its exit on the side of the verge. hair-like projections of the periostracum (hirsute). Internally (fig. It contains cavities for the loop of the intestine. The uterus (U) is wider than the vagina and the nidamental gland The uterus and is wide and made up of many large diverticula (fig. 533. long.2 mm.) consisting of about 22 long (less than 0. It NG I . 5). two pilasters. long. filiform tentacles characteristic of the Planorbinae with the eyes at the inner base.

The ureter is short in convexiusculus and obliquus but longer and more tube-like in hirsutus and albus. The penial comjilex is very long and narrow but generally resembles that of hirsutus. A. figs. Mus. fig. p. as in other genera of Planorbinae. 3). 8). fig. The seminal vesicle is practically the same. 2). however. The figures on plate 19 agree with tliose by Annandale and Prashad (Rec. 27) shows a smaller number of prostate diverticula than appears normal for the genus. and the spermatheca is on a longer duct than in the specimens from Poland examined. from India (plate 19. Plate 16 fully illustrates the genitalia of obliquus. and the veins to be placed at each end at about the center. I Respiratory and Renal Systems. with fig. 9 mm. long) and albus (which has the same form as hirsutus) the kidney is short and wide. In Gyraulus convexiuscidus. 2.vius cuius (Hutton) (plate 19. (plate 14. C. This difference in form of the penial complex easily separates albus from hirsutus (compare fig. 40) except that the spermatheca in the specimens examined is on a much longer duct than is shown in their figure. and India. fig. long). Sections through the kidneys at about the middle show this organ to be flattened. 15). The pseudobranch of Gyraulus albus is broad. 3. Figure 15 is an unusual form. and they are figured more irregularly than was observed in the specimens j^ersonally examined. Baker (plate 19.) (1935. 17. Conve. p. the lumen much wider than high. No separate prostate duct is indicated. The prostate shows fewer diverticula. 4) . 8. flattened organ. quite unlike the kidney of the two species previously mentioned. It is similar to that of vermicularis (fig. In Gyraulus deflectus obliquus (DeKay) there are forty diverticula on the prostate which is much longer than either albus or hirsutus. Gyraulus conve.. The rectum passes through the pseudobranch. The kidney (plate 48) is long and narrow in obliquus (fig. fig. possibly pathologic. 10) is higher in proportion to width showing that the kidney is thicker than in the other species. plate 15). The pneumostome is broad (fig. 3. 9) has thirty-two diverticula on the prostate and the penial complex resembles that of obliquus. There is no ridge. figs. XVHI.vius cuius (fig. has from twenty to twenty-two diverticula on the prostate as in albus (plate 15. the nearest relative to albus. Figure 4 shows the pseudobranch cut open to indicate the vascular network of blood vessels. Gyraulus latestomus F. 2. The spermatheca is also on a longer duct than in those species of the genus personally examined.5 mm. The pseudobranch appears to be similar in form in species from Europe. fig. The penial complex has a much shorter preputium with a vergic sac one and one-half times as long as the preputium. PS). Ind.68 The MoUuscan Family Planorhidae similar in general form. America. 3) the pseudobranch is as broad as in albus. In Gyraulus hirsutus (plate 15. 18. 3) it is flat and very thin and the pneumostome is very large.5 mm. American species examined differ but little from the European Gyraulus albus. and flattened. Gyraulus hirsutus (Gould). In Gyraulus deflectus obliquus the pseudobranch was observed to be of several forms (plate 16. 4. 11) are similar in form. figs. plate 14.xiusculus (fig. In hirsutus (fig. It is in each case a broad. 19). somewhat triangular. 2. fig. and latestomus (fig. 12). Soos's figure of Gyraulus laevis Aid. long) and conve. The prostate diverticula are also longer and club-shaped toward the end (plate 16. 13. 14. In Gyraulus latestomus (plate 19. hirsutus (fig. 2) of India has about twenty-five prostate diverticula. 11. but obliquus (fig. 8) it is broad and flattened.

10-13 intermediate teeth. olbu. an unusual condition in the genus. Genera. A. B) where the inner laterals are shown as bicuspid. 11-14 intermediate teeth. 25 specimens from park Collected bv Mr. fig. The cusps of intermediate teeth 10-11 are short. The center tooth varies in size in the 43rd to 62nd rows being a trifle wider at the top than in the first fortytwo rows. Ind.SuhjamUies. Probably the entocones were not observed. and 14-20 marginal teeth. The ducts of these glands are long. The laterals (1-9) are tricuspid with long. For the above anatomical data the following material has been teeth. in Krolikarnia. and Subgenera — Recent and Fossil 69 Digestive System. fig. 8. available: G. about forty on the entire jaw. the mesocone is larger. and 14-22 marginal teeth. Gyraulus albus has the same number. In several membranes examined a very small 4th cusp appears on the 6th tooth and continues to the intermediate teeth. The specimens examined differ from the figure by Annandale and Prashad (Rec. the cusps long and narrow and reaching only about half waj. 10-13 intermediate teeth. The radulae of all of the typical Gyraidus examined have been similar in character. The intestine. 18) the jaw is arched and consists of over fifty plates which are narrower than those of albus and hirsutv^. There are 130 rows of teeth. around the liver. Jankowski. are very small and sharp. 13) is pyriform with the radula sac greatly developed and protruding behind (RL). The entocone is split into two small cusps. and then loops forward to the rectum which has its exit on the pseudobranch. In Gyraulus dejiectus obliquus (fig. . a suburb of "Warsaw. and jndoris plainly indicated. 10-11 intermediate teeth. AIus. 1) is similar to that of albus. for they were present in all of the laterals of the teeth personally examined. Gyraulus convexiusculus has the formula 20-1-20. The radula formula of Gyraulus albus (plate 68. There are 175 to 183 rows. with 1-9 lateral teeth. Poland. 9) is 18-1-18 with 103 to 114 rows of teeth. after leaving the in'loris. narrow cusps.. gizzard. The salivary glands (SG) are elongated.s. and 15-20 marginal ( ) There are 155 rows of teeth. fig. makes a looj) over the oesophagus which extends backward. three in number. The radula of Gyraulus hirsutus (plate 69. and almost as long as the buccal sac. 2) has a formula of 22-1-22. 17) has large plates. fig. The stoiiiacli (plate 48. fig. The buccal sac (of Gyraulus albun plate 70. All species of typical Gyraulus examined have the same type of stomach and intestine. The jaw (plate 50) is horseshoe-shaped and fragmented as in Planorbis and Anisus. and the ectocone is split into two sharp cusps. and the ectocone cusps. The radula of Gyraulus dejiectus obliquus (plate 69. p. 55. The center tooth is bicuspid. with 1-10 marginal teeth. 13j is cylindrical with the crop. The 86th to 103rd rows have a central tooth a little more than half the width of those in the 43rd to 62nd rows. The marginal teeth (12-18) have the reflection high up on the tooth. placed high up on the reflection. cylindrical. fig. Gyraulus latestomus has the formula 20-1-20. Gyraulus hirsutus (fig. The figure of the radula of Gyraulus euphraticu^ (C) by Annandale and Prashad is more correct in the tricuspid character of the lateral teeth. In latestomus the jaw is similar to that of obliquus and has over forty plates. with 1-9 lateral teeth. and 12-19 marginal teeth. are wide and low and 6-cuspid. with 1-9 lateral teeth. The formula is the lower margin of the base of attachment. XVni.

Prashad. 6 specimens from Taylors Lake. Africa. deflect us obliquus. G. and Beq. arcticus he bra if us Bgt. hirsutus. convexiusculus. Asia. Species of Europe and Western Asia alhus Mliller acronicua Ferussac ancylus West. 3 specimens from Cedar Lake. regulosus Lindholm socius West. Baker. Ontario. Ontario. C. intermixtuii Mouss. Below is given a list of some of the species of Planorbids believed to belong in the genus Gyraulus* Some of these species may be found. devians Porro drapanauldi Jeff. Gray aethiopicus Bourg. which Mr. parts of the West Indies and northern South America. near Quebec. on anatomical examination. New Caledonia. The list is probably not complete. collected bj' Dr. infraliratus West.V-C. chedeaui Germain cockbruni Godwin-Au. Specific regions include North America. gibbonsi Nelson kigeziensis Preston kisumieiisis Preston lamyi Germain leucochilus Melville andcrsoni Ancey avakubiensis Pils. tctragyrus West. Wisconsin. to belong to other subgenera or genera.J. A. Fiji. — .70 The Molluscan Family Planorbidae G. Collected by Mr. India. polaris West. Philippines. Umnophilus West. 3 specimens from Calcutta. Collected by Mr. LaRocque. numidicus Bgt. differing in this respect from all other genera of Planorbidae. G. Collected by Mr. crossei Bgt. Europe. Malaysia. LaRocque. capocestianus V. 9 specimens from mouth of Bass Creek. LaRocque. Canada. janinensis Mouss. Australia. laevis Alder M oiler albopersicus Germain borealis Loven hourguignati Moitessier brondeli Raym. H. latutomus. Baker had never fully edited. Brus. Collected by F. Lake Xipissing. A. E. piscinarum Bgt. 13 specimens from Lake Chetek. the author's name follows each specific name with disregard for use of parentheses to indicate change from original generic assignment. A. and other islands of the Pacific Ocean. The genus Gyraulus is practically worldwide in distribution. agraulus Boiu'g. G. Geographical Distribution.sten and Ponsonby niisellus Morelet natcderuiia Krauss and Ponsonby fouladougouensis Germain gardei Germain costulatus Krauss crmvfordi Melville sperabilis Preston schweinfurthi Clessin tilhoi Germain toukotoensis Germain *In this faunal list. Algonquin Park. gredleri Bietz Species of Australia and Tasmania essingtoneiisis Smith gilberti Dunker meridional^ Brazier planissimus Mousson Species of Africa abyssinicus Jickeli adansonii J. B.

Austen compressus Hut ton convexiusculus Hutton cherraeiisis pangongensis Nevill proclivis Martens rotuln Benson saigonensis Crosse saltensis Germain sivalensis Hutton stelzneri Dohrn demissus Westerlund elegantulus and Fischer Dohrn euphraticibs Mousson himlayaensis Hutton hohcnachcri Clessin haltoni Benson hiiptiocyclos stewarti Germain Benson injraUneatus Martens intcrmixtus Mousson issykulensis Clessin sumatranus Martens tondanensis Quoy and Gaimard velijer Annandale yeni Ping and Yen Species of Japan amplificaius Mori biwaensis Preston japonicus Martens noziriensis Mori hiemantium Westerlund infirmus Mori iwaotakii Mori ?)titidanensis pulcher Mori tokyoensis Mori Species of the Philippines Bartsch quadrasi Mollendorff Species of America arcticioi (Beck in Moller) boetzked (Miller) burealis Lo\'en cyclostomus F. are quite unlike and the genera should undoubtedly be separated on conchological grounds alone. ]\Iore than ninety species of Gyraulus are listed on the previous pages (excluding those from America). All of those listed have been included in the genus Gyraulus by competent conchologists. particularly in the presence of a horny stylet on the verge. . Baker Geological Distribution. C. Baker Intistoiinis F. A7iisus. Gyraidus. In fact. especially the genitalia. Some of these may be found to belong in other genera. Remarks. The shells of Anisus and Gyraulus. Completeness is not claimed. based on their anatomy. Genera. and Bathyomphalus form a natural group. dcflectus (Say) Baker deflectus obliquus (De Kay) hirsutus (Gould) hornensis F. and Suhgeuera — Recent and Fossil 71 Species of Fiji Group singularis Mousson Caledonia i-us-siteri Species of montrouzieri Gassies New Crosse Species of India and Asia associatus Westerlmui ocutua Clessin barrakporensis Clessin cnntori Benson labiatus Benson laidocensis Nevill liratus Westerlund nialacaensis Germain Godwin. it is probable that several additional species should be added to those listed. C. when examined anatomically. C. however. According to Wenz the genus Gyraulus dates geologically from the Middle Eocene period. The genus Gyraulus is closely related to Anisus in its anatomical features.Subjani'dies. Also.

As section of Gyraulus in synonymy. pp. Prostate (PRS» more than half as long as the sperm duct with eleven long and cylindrical diverticula. 204. As subgenus of Gyrajilus. 83. Alaska Moll. Pars 22.. British Xon-Marine Moll.\rd. not hirsute.. Moll. Type Planorbis parvus Say. Zool. I. Type Planorbis parvus Say.) of the same diameter glands which in length. 4-6). 392. Internally (fig. As subgenus of Planorbis. As section of subgenus Gyraulus. 6) Seminal vesicle (SV) short (0. Torquis Dall. C. pp. Torquis W. GENITALIA. No. Wis. and cylindrical. The vas deferens is at first a large duct but becomes smaller in diameter toward the penial complex (VD). p. Torquis Germain. Rec. Cat. The vcrgic sac is longer than the preputium.. Torquis F. or excavated (appearing as though reamed out). pyriform. Genotype Gyraulus (Torquis) Say. enlarged at the upper end..5 as the sperm duct. As subgenus of Gyraulus. Not differing from Gyraulus in general form. smaller than Gyraulus. The vergic sac is long and cylindrical and is enlarged at the upper end (VS). 373. p. Gyrcudus (part) Martens. Contr. joining with the sperm duct to form the vas deferens. Between the vergic The penial complex sac and the preputium there is a large papilla and below this a swelling of the wall of the preputium representing the better-developed muscular ring or diaphragm of typical Gyraulus (not shown in the figure).. Univ. A cross section through the prostate shows the cylindrical or pear-shaped diverticula on a separate duct which has a greater diameter than the sperm duct (fig. The verge (V) is long and narrow. As section of subgenus Gyraulus. 94. Type Planorbis parvus Say. 86. very narrow below where there is a horny stylet (ST). Wis. 1905 original designation Planorbis parvus Say 1899.3 mm. 1923. Biol. Torquis Kennard and Woodw. There are a number of small stand out like pustules.. ANATOMICAL CHARACTERISTICS PLATE 17 (Gyraulus parvus) . 6. 1905. the lip often slightly thickened within. Miscel.72 The Molluscau Family Planorhidae Subgenus Type by TORQUIS Dall. 34. XXI. Shell (plate 77.. C.\lker. 1918. 12) the preputium has two vertical pilasters. Male Organs mm. 1928.. p.) and narrow. PR) is very long (about 1. Includes Planorbis parvus Say. As subgenus of Gyraulus. Trans. 12. 6. the base (left side) more or less concave. Type Planorbis parvus Say. Torquis F. The free portion of the prostate duct is very short (PD). 1926. Fossil. Mus. 8. 1921.. Sperm duct a trifle more than 1 mm. VS. in length. 150. and connected with the vagina by a duct about as long as the spermatheca . and Arts. Ultradextral. It is (fig. Type Planorbis parvus Say. Baker. The opening of the sperm canal is on the side as in typical Gyraulus. 1926. figs. Ind. Mich. Cent. Baker. Amer. with the whorls less distinctly spirally striated. XXII. Female Organs. 6. There is one large retractor muscle attached near the junction of the vergic sac with the preputium (fig. 67. Torquis Wenz. p. S) is small. Animal. p. SS. 10). 1627. Fresh-water Moll. Sci. Acad.. pp. about 1 mm. Mus. (fig. RIM). The vas deferens is not enlarged as it enters the vergic sac. The spemiatheca (fig.

10). Several other species of the subgenus Torquis have been dissected and may be compared with parvus. Its duct is long and looped (fig. Figure 5 on plate 18 and fig. The oviduct is short (about 0. and is composed of large diverticula. 8) it is flattened and somewhat triangular. 11). also fig. fig. longer than wide (about 1 mm. fig. 5) it is like that of vermicularis but the rectum has a slight ridge which was not observed in any of the other species examined. OT. ( . Hermaphrodite Organs. 2. and ten diverticula in the ovotestis (plate 15. 4). the ureter short and recurved. long. long). The ovisperm duct is small and tube-like and is longer between the seminal vesicle and the ovotestis than between the seminal vesicle and the oviduct (fig. is squarish in form. The duct to the albumen gland is not so long fig. The ovotestis is rather short and is composed of Init few club-sha})e(l diverticula arranged in a double row (fig. No figures of the genitalia of the Torquis group iniblished jireviously. wide). In Gyraidus circum- . flattened. sperm duct. The rectum pierces this pad as well as the pseudobranch and has its exit (A) above the leaf-like pseudobranch. C. the two groups are very similar in the form and position of the ticula in the prostate The genital organs. 1. fig. 9. and is pierced by the rectum. 6. and the rounder spermatheca on a shorter duct. and Subgenera fSD). A section through the penial complex (fig. The verge is very long and slender and has a large stylet (fig. The junction of oviduct. The pneumostome is very large. fig. 6. 2) The penial complex is the same in both species which are of about the same size. however. In the penial complex the vergic sac is shorter than the preputium. 1) there are nineteen diverticula on the prostate and the ovotestis diverticula are few in number. only about half as long as the preputium. 7. 8) is flattened. The jiseudobranch of Gyraidus parvus fig.Subfamilies. gland 1. hence no comparisons can be made. There is a heavy swelling at the upper part of the preputium where the papilla and muscular ring are located. the preputium being long and cylindrical and the vergic sac very short. In Gyraidus similaris (F. have been Respiratory and Renal Systems. The prostate has twelve diverticula.5 mm. In Gyraulus circumstriatus (fig. and ovisperm duct is shown in fig. In the four species of this group studied there was a large. Genera. the fewer diverticula of the prostate. parvus) is very long and narrow (4 mm. The albumen duct. fig.5 mm. fig. The genitalia of the subgenus Torquis differ from those of typical Gyraidus in the form of the seminal vesicle. Baker) (plate 18. SO). muscular jiad extending backward from the free portion of the pseudobranch to the place where the rectum begins on the intestine.). The junction of the oviduct and sperm duct to form the ovisperm duct is shown in fig. 11). DA). 0. 3) the pseuclobranch is much narrower (the animal was much contracted). 3) shows the relationship between the several organs. leaf-like. In general. penial complex is somewhat different. 4) is similar in form. 8 on plate 15 show the form of this pad. In Gyraulus similaris (plate 18. In Gyraidus vermicularis (plate 15. (plate 17. wide. The spcrmatheca is the same. Gyraidus circumstriatus (Tryon) (i)late 17. Several of the diverticula of the ovotestis were filled with ova ready for extrusion (fig. The vagina its is — Recent and Fossil tlie 73 narrow and about as long as spernuitheca and The uterus and nidamental gland are relatively short and wide (U). In Gyraulus vermicularis (Gould) there are sixteen diver- fig. The kidney (plate 47.

but not enough material is available for study to confirm this suggestion. fig. both lots collected by Dr. C. the cusps not reaching the lower margin of the base of attachment. Intermediate teeth (7-8) with the ectocone split into two small cusps. The 9th tooth is also transitional with the ectocone split into three small cusps. The intestine does not form a posterior loop. Baker and A. collected by F. 14 specimens from Winnebago Lake. 4) the formula is 22-1-22 with 1-7 laterals. California.siyiiilaris (F. Wisconsin. Henry Co. The radula teeth of the subgenus Torquis do not differ in general characteristics from those of typical Gyraulus. 16. D. California. The subgenus Torquis is found throughout North America from Mexico northward. Colorado. and Meach Lake. Gyraulus laevis Alder. Illinois. 17 specimens from Smartweed Lake. Canada. The digestive tract shows division into crop. 3). collected by Dr. Cross sections through the kidney of the four species show a similarity of conditions (fig. 2 with figs. There are 120-125 rows of teeth. fig. 3. For the above anatomical data the following specimens were studied Gyraulus parvus (Say). Gyraulus circumstriatus (fig. Wisconsin.: 74 striatus The Molluscan Family Planorhidae (fig. Croix. C. long). In Gyrauliis vermicularis (Gould) (fig. The kidneys in Gyraulus and Torquis differ little in structure. fig. Bishop. Baker. Gyraulus circumstriatus (Tryon). 17 specimens from Wainwright Park. Geographical Distribution. narrow jaw with about thirty-seven plates.. It is also apparently represented by Gyraulus santacruzensis Germain from the Island of St. Toland. The buccal sac is like this organ The jaw (plate 50. 5 on plate 69. Compare figs. Lake. 19). tricuspid. and North Star Lake. 4. In vermicularis it is short and wide as is also the case in similaris. but crosses over the oesophagus and extends forward as in the genus Planorbis. with 1-6 laterals. a common species of Europe. There are 1-6 lateral teeth. Gyraulus vermicularis (Gould). San Francisco. C. 8-10 intermediates. fig. Frank Smith. LaRocque. West Indies. 6 circumstriatus . collected by Dr. 9 similaris). . Some of the other species listed under the distribution of typical Gyraulus may belong to Torquis. Digestive System. and pyloris and there is a large blind sac. Baker) (fig. and 11-22 marginals. parvus) is horseshoe-shaped with many small plates as in Gyraulus typical. 22) has a small. the mesocone large and spade-shaped. Baker). 2) the kidney is much shorter and wider (less than 2 mm. collected by F. The radula formula of Gyraulus parvus (Say) is 14-1-14 with 105 rows of teeth (plate 69. 17-18) are not as typically horseshoe-shaped as in the subgenus Torquis. Swales. fig. In Gyraulus similaris (F. also belongs to this subgenus. 7 vermicularis. The jaws of Gyraulus (figs. Hanna. 10 specimens from Mt. in typical Gyraulus. The center tooth is bicuspid. Gyraulus circumstriatus (Tryon) has the formula 14-1-14 with the same number and position of teeth as in parvus. with spade-shaped cusps. as in typical Gyraulus. 5) the formula is 15-1-15. There are 130 rows of teeth. Gyraulus . There are approximately thirty-six to forty plates on the jaw of parvus and Gyraulus vermicularis has about forty-seven plates (fig. Typical marginal teeth (10-13) have the entocone split into two small cusps. and Hat Creek. G. 1. gizzard. 5 parvus. 7-10 intermediates and 11-15 marginals. and the ectocone split into three small cusps. Alberta. C.

p. Baker. referred to on a subgenus Torquis is similar in anatomical details to typical Gyraidus. Nat... of Hartmann in 1842. As subgenus of Planorbis. Akad. Fauna Palaearc.). Rad. Planorbis crista (Linn. 1929.. Archiv. Nautilus crista Linn. 13. Armiger W. Armiger Germain. Armiger Lindholm. It is. 6. Armiger H. As subgenus of Gyraulus. Torquis apparently dates geologically from the Oligocene period. 86. Bull. V. 65. pp. Nautilus crista Linn. Sci. Schneck. p. Aryniger We.. XXI. Mu. Jugoslav. Ideogenotype Plnnorbis nautilus Stein Planorbis cristata Drap. like parvus and vermicularis. p. Em-op. 385. Gatt. 253. 204.. III. Nautilus Gray. which have narrower and smoother whorls than the species of typical Gyraulus. pp. 121. Armiger Westerlund. I. Zool. Type Planorbis crista (Linn. As genus. p. 172. Cat. and Subgenera — Recent and Fossil 75 in tliis Species Considered as Valid.). like alhus and de flee tus. Rec.. As subgenus of Planorbis. Armiger Hartmann. 153. Type Plariorbis crista (Linn. Gasterop. Leben.). Drap. however. und Musch. Contr.sterlund.. the for some small details in genitalia. Quest France.alker. 1847. Mich. pp. Type Nautilus Type evidently p. 1905.ska Moll. 1842. Ind. p. Nat. 81. Mus. V.). of Wisconsin. 219. 1923. Type Turbo nautilus Linn. Genotype Gyraulus (Armiger) crista (Linn. 1902. 1840 Remarks. Tyjie Armiger Nautilus crista Linn. 94. XIII. Baker) Gyraulus (Torquis) vcrDiicularis (Gould) (hjrnulus (Torquis) vermicukiris Gyraulus (Torquis) circuinstrint as (Try on) Gyra ul us ( Torq uis ) circ u ni s[ ria tus u'nlkcri Vanatta hendcrsoni Walker Geological Distribution. Armiger Dall. 8. Soc. Oligocene of Dry Hill. 1625. Type designation by Lind- Armiger Hartmann. table. Ala. Monogenotype Armiger cristatus T)rap. Syst.. 83. Reg.Subjcunilics. Torquis siliceus Pilsbry and Brown. p. Miscel. 58 year. Erd-und Siiss. 1926. 1931. a convenient name for the smaller species with 'reamed out' left side. C. C. Univ. Fossil. fiir Mollusk. Armiger Wenz. No type cited. Pars 22. Armiger F. No. Fresh-water Moll. Tj'pe Planorbis crista (Linn. pp. ubers. Proc. 181. Mus.). 1850.\As.. As section of subgenus Gyraulus.seo Cien. 1903. Type Turbo nautilus Linn. As subgenus of Gyraulus. Except Type by designation 1840. Barcelona. p. As subgenus of Planorbis. ])revious page. C. As genus on conchological = grounds.. 151. cristatus = = holm 1885. As section of subgenus Gyraulus. 1928. Antigua. Zool. Heft 6. 1918. The American species included subgenus are: Gyraiilus (Torqids) aeruginosus Morelct Gyrauluti (Torquis) arizonensis Pilsbry and Ferriss Gyraulus (Torquis) allissimus (F. 386. 1923. Goiera.. Baker) Gyraulus (Torquis) carus Pilsbry and Fcrriss GtjmuJus (Torquis) parvus (Say) Gyrauhts (Turquis) mntacruzensis (Germain) Gyrnuluj< (Torquis) siniildris (F. Genus ARMIGER Hartmann. in 1926. ii. Vide Germain. Nautilina Stein. Soc. crista Linn. Spiniformis Germain. Trab. i*. . Berlins.

6). crista 1931. The albumen gland (fig. Type Anisus (Armiger) crista (Linn. 22. Male Organs The (1 free portion of the prostate duct is very short. Allat. As subgenus of Planorbis. a dependent papilla (D). il. Fauna Takern. 30. The vergic sac is enlarged at the end w^here the vas deferens enters it as an enlarged tube. (Linn. ANATOMICAL CHARACTERISTICS PLATE 18 . Die Mollusk. bent upon itself in a loop posterior to the prostate. with few rapidly increasing.). costate whorls. de France. Armiger A.76 1929. As subgenus under Anisus. Soos. PA. IV. 481. p. elongated and composed of large gland cells. Kozleni. The prostate has nine . but a small.^in. cylin- mm. long. His figure (1935. The oviduct (OD) is short and greatly swollen below the junction with the sperm duct to form the ovisperm duct. 540. The general form is like that of Gyraulus. Germ. in the center and not at the side as in Gyraulus (figs. The Molluscau Family Plauorbidae Armiger Odhxer. It is joined to the large. II. drical. As genus on anatomical grounds. and also a muscular ring or diaphragm (not shown in figure). small duct. Prostate 0. PA). The carrefour connects with the oviduct. fig. cylindrical diverticula placed on a separate prostate duct. small. taj^ering gradually to the short. (Linn. Moll. External form like that of Gyraulus. 28) differs in several respects from the ones here presented. the uterus short and .). The verge (V) is elongated. 7) Seminal vesicle (SV) with very few large glands or diverticula. GENITALIA. SC). Ultradextral. The opening of the sperm canal is just above this papilla at the end of the verge. pointed fleshy papilla (fig.5 mm. wide vagina (VG) the duct being very short. fluv. Internally (fig. Animal. ovotestis Hermaphrodite Organs. p. 8) is large (1 mm.. Vas deferens a long 7) with a vergic sac one and one-half times as long as the preputium. long. As subgenus of Gyraulus. 11. Shell (i)late 76. Handbuch. terr. Sperm duct 1. Armiger Thiele. Teil 2.). Penial complex (fig. tapering gradually to the end. p. The preputium is of larger diameter than the greater part of the length of the vergic sac. The uterus (U) and nidamental gland are wide. very fine duct (DA). Artniger. The space occupied by this organ is shown in the figure (AL). Female Organs. bulbous carrefour (CF) by a long. There is one retractor muscle (RM). (plate 18. which does not bear a horny stylet.5 mm.). Soos's paper in 1935. j). No type indicated. Type Armiger Type Plonorbis crista 1931. 10. p. 10. 1935. the gland rather long. The spermatheca (S) is long and pyriform in shape. There are two }ulasters. fig.). The ovisperm duct (SO) is twice as long between the oviduct and the seminal vesicle as between that organ and the ovotestis. consisting of thirteen long. the costae usually projecting at the periphery. No figure of the whole genitalia of Armiger was available before the publication of A. 9) the j^enial complex is formed like that of Gyraulus. XXXII. as in Gyraulus. The (OT) consists of many club- shaped diverticula placed in a double row. The intestine extends the whole length of the albumen gland which covers the greater jmrt of the intestine. fig.

convex above. 20) is narrow and characteristically horseshoeshaped with the lateral processes elongated. The spermatheca in Soos's figure is also on a longer duct. In 1929 (p)). Genera. J. 3). even in the lateral teeth. was only 11-1-11 while the larger form has a formula of 14-1-14. The stomach region resembles that of Gyraulus (fig. The kidney (plate 47. The teeth resemble those of Gyraulus except that there is always an accessory cusp above the ectocone. 5). These represent the variety i}nbricatus. and the marginals (12-13) six-cuspid. Lake ]\Iendota. The jaw bears a larger number of plates. The separate prostate duct is not shown in Soos's figure. by Dr. 21) shows it to be flattened. The outer marginals are vestigial. provided most of the anatomical data. Respiratory and Renal Systems. 13). gizzard. is 16-1-16 with 150 rows of teeth. The formula. 20) is short and wdde. There is no stylet at the end of the verge and the opening of the sperm canal is at the center beside a small papilla and not at the side above a stylet as in Gyraulus.Subfamilies. ii. [Morrison. collected by A. The buccal sac is a distinct division into crop. The genitalia of Armiger and Torquis may be compared on i^late 18. Wisconsin. also. The jaw most nearly resembles that of Gyraidus circumstriatus (fig. The seminal vesicle is also different in form of glands. 22. This is. 540) says 'Appareil genital comme chez les especes stylet first du sous-genre Gyraidus. The section is also similar to that of Gyraulus vermicidaris (fig. For the above anatomical data ten specimens were dissected. (plate 50. Jankowski. P. fig. a suburb of Warsaw. 22) except that it The radula formula . is Digestive System. 30) Odhner called attention to the absence of a stylet in Armigcr. figured the verge of Anniger crista to show the absence of the and the i)resence of a small penial i)apilla. like that of Gyraidus. The center tooth (fig. ending in the rectum on the pseudobranch. It bends around the liver and turns forward. 7). the indication of this anatomical difference between Armiger and Gyraulus. In the region of the stomach (plate 48. The rectum is a large tube bordering the upper side of the pseudobranch and the anus is above the pseudobranch. The penial complex differs in several respects. somewhat resembling the kidney of Gyraidus vermicularis (fig. There are twenty-two plates in the upper or superior jaw and fourteen to sixteen plates in each of the side jaws. leaf-like. The is bicusj^id and resembles that of Gyraulus vermicularis laterals are four-cuspid (1-9). fig. apparently. 15) there and pyloris. with the veins near the center of each side of the oval lumen. E. the intermediate teeth (10-11) five-cuspid. The pseudobranch (plate 18. near [Madison. collected from a swamp on University Bay. Poland. Germain (1931. 6) broadly triangular. The intestine makes a loop forward and then backward under the oesophagus. where the difference in the structure of the termination of the verge is i)lainly indicated. Odhner noted a difference in the size of the teeth and formula of crista. is fig. A section through the kidney near the middle (fig. Typical crista from a pond in Krolikarnia. and Subgenera — Recent and Fossil 77 divertieuhi which upi)ear widely fjpaeecl while there were thirteen crowded diverticula in the specimens personally examined. These differences in anatomy are sufficient to remove Armiger as a subgenus of Gyraulus and indicate its right to full generic rank.' But this is not the case. those from the Takern having much smaller teeth than specimens from near Stockholm. fig.

The genus contains several forms which should probably rank as species. From these observations he concludes that the development of the stylet (penisstachel) may be checked in the later fall. 20. 30) genus. gives the group full generic rank on the basis of the lack of a stylet on the verge which is characteristic of Gijraulu^^. 10). Another species in eastern Asia has been named annandalei by Germain. but the subject is one of great interest and dissections should be made of many species at different ages to determine whether the stylet is absent in young and immature Gyraulus. There appears to be some variation in the genitalia and also in the radula. 21. and Atropoarmiger (vide Lindholm. Soos (1917) and A. 24). This is undoubtedly true. Odhner found the formula to be 11-1-11 and 14-1-14 in crista from two localities and the writer obtained the formula 16-1-16 from crista . Odhner (1929. Lindholm (1926) considers it a genus. p. from Algeria. but the lack of a penial stylet removes it from that group. He found young Gyraulus borealis of two whorls without a stylet on the verge while those of three and one-half whorls had a well-developed stylet. this Species Considered as Valid. Dybowski and Grochmalachi erected three subgenera based on spiny variation in Armiger: Nautiloarmiger. is shown (fig. Specimens of Gyraulus laevis of three whorls had a stylet. 20) does not fully agree with Odhner. Haas. but these were either full grown or nearly ( grown. Thiele (1931) considers it a subgenus of Anisus. 30) states his belief that Armiger represents a stage in the evolution of Gyraulus. L. crista (Linn. A good figure of the verge of Armiger crista. The value of recognizing subgenera for these variations may be questioned seriously. in which no ribs project from the full shell. showing the small penial jjapilla at one side of the sperm outlet. By the form of the penial complex tliis group is distinct from Gyraulns and should be considered as of full generic rank.) crista spinulosum Armiger imhricatus (Miiller) Armiger annandalei (Germain) (Clejsin) Remarks. pp. with strong ribs projecting from the shell. This group of small snails is highly variable specifically and a number of names have been bestowed upon the variations. p. Armiger crista is reported from the trans-Mediterranean region. possibly through some unfavorable climatic condition. and imbricatus (Miiller). Tunis. Armiger spinulosum (Clessin) is i^robably a race of crista Linn. Armiger is Palearctic in distribution. The only species definitely assigned to genus in the present volume are: Armiger Armiger crista (Linn. Odhner (1929. in a review of this work 1930. 1632-1633) Armiger known from the Middle Miocene period. 1926. 386) considers Armiger a distinct p.). etc. It has been present in all specimens personally examined. A species {Armiger annandalei Germain) has been described from eastern Asia. Soos (1935) list it as a subgenus of Gyraulus. In 1922. Haas (1929. p. believing that more study should be given the subject before conclusions are made. Geological Distribution. Cristoarmiger. Two forms appear distinct enough to rank as species.78 The Molluscan Family Plauorbidae Geographical Distribution. According to is first Wenz (pp. occurring in the northern United States and Canada. northern Europe and Asia.

Animal. Acad. Variation of Planorbis andecolus Orb. 33.-section of Planorbis figs. Monotype Planorbis andecolus Orb. Taphius Tryon.) Lake Titicaca. Taphius Fischer. Geographical Distribution.. spire greatly depressed below the level of the body whorl. Conch.) Lake Titicaca. p. As subgenus of Planorbis 1883. Hist.. the spire whorls bluntly angulate. Adams. 1899. Ann. Type Planorbis andecolus Orb. — Recent and Fossil be individual. IX. Mus. Zool. p. Lye. T. Amer. and Subgeneracollected in Poland. 479. Type Planorbis andecolus Orb. As . all the whorls visible and nearly in the same plane. as pronus and subpronus. 507. As section of Planorbis 1855. B.. N. Some of the species. Venezuela.. p. Papers. Species Considered as Valid. p. Only five species are here referred to Taphius. \). 1886. and A. Type Planorbis andecolus Orb. probably of Pleistocene age. Baker. no lamellae within the aperture. expanded. Ind. in diameter. 81. Rec. de Conch. 1924. plate 4. and A. 396. XXI. XVII. 1905. whorls few. Type Planorbis andecolus Orb. The characteristic of the animal and its anatomy have not been described. Syst. The systematic position of the genus Taphius is greatly in doubt because its anatomy is unknown. Adams. Type Planorbis andecolus Orb. Y. and S. 79 may all The group (k'nus TAPHIUS H. Mentions Planorbis subpronus Martens. 86.. Proc. raindly increasing body whorl wide and much enlarged. pp. and A. As subgenus of Planorbis Taphius D. Taphius D. Cab. Nat. aperture large. S. 43. South and Central America from southern Mexico to Peru and Bolivia. 62. Type Planorbis andecolus Orb. Genera. Type Planorbis andecolus Orb. Sci. As subgenus of Planorbis Taphius Martens. p. Peru pronus (Martens) Venezuela s}ibpronus (Martens) Mexico Geological Distribution. Taphius Taphius Taphius Taphius Taphius andecolus (Orb.). Not definitely known. the 13-15). . 262. Ad. 7. Ill.. Sinistral. 1855 p. 1931. Remarks.Subfmnilies. 1930. Genera. Biol. Tj'pe Planorbis andecolus (H. error for Orb. Mus. 210. Taphius H. 351.. As subgenus of Planorbis Taphius Clessin. Cent. As genus Taphius Thiele. II.\ll. Oc. umbilical side flattened. Conch. Cermain (1921) makes it a subgenus of Planorbis allied to Helisoma. Typical andecolus bears a striking resemblance to the tyi')ical group of Helisoma and Dall (1905) made Taphius a synonym of that group. p. Handbuch. p. Nat. 106. Alaska Moll.. 1921. 1884. Peru titicacensis (Clessin) Lake Titicaca. Taphius H. Peru moiitanus (Orb..all.. As subgenus of Planorbis Taphius Pilsbry. These differences merits additional anatomical study. Mich. Shell (plate 77. 49. As sj'nonym of Helisoma Taphius Germain. Moll. T(tph{}(s pronus has been found fossil in Lake Valencia. Man.\ms. Univ. 1870. resemble certain species of Tropicorbis and Taphius may be another genus of Planorbidae peculiar to both Central and South America. p.ype by original designation Planorbis nndecolus Orb. Part 2. Phil.

As 1931.. Acad. Type 1921. Nat. 70 (non Gyrorbis Fitz.\as. 99. Sci. 1627. Fossil. Ind. 55. 1899. 480... Papers Mus. Obstructio Haas. Type Planorbis liebmanni Dkr. p. Tropicorbis maya Morelet. 1941. The labial lamellae differ principally from those of the genus Planorbula in having the lower palatal lamella pointing slightly downward on the left side while in Planorbula this lamella always points upward. 1940. 106. Anatomy described. in the group Hcddemanina the upward end forming a vertical and transverse lamella combined. Planorbis liebmanni Dunker. Rec. Genotype bis) liebmanni (Dunker). Handbuch. iii. Tropicorbis Pilsbry and Brown. As genus. As section of Planorbis. and A.. Biol. Type incorrectly given as Planorbis tnaya Morelet. 1899. Teil p. 10-12. rather is it allied to Tropicorbis. Mus. . pallidus) whorls in view equally on both sides. and A. Proc. p. orbicular. Includes 1914. What the group may be and where it may be placed can only be determined by an anatomical study. II. p. Mex. Tropicorbis Wenz. As genus. Baker. 66. 86. of few whorls which may increase regularly in diameter (plate 77. figs. As section of Planorbis. Mich. 212. 1923. Nautilus. plate 4). Sci.. now referred to Tropicorbis. 1884. and S. figs. C. Tropicorbis Thiele. 398. p. Phil. As genus. 31. Nautilus. S. o\^. Menetus M. Phil. Type wrongly stated to be Planorbis 1934. B. described. 104. aperture unarmed or with one large parietal lamella with a small infraparietal below it. distinctly curved (plate 76. As subgenus of Planorbis. Planorbula Martens (part). 390.. but pronus certainly does not resemble any form of Helisoma. Field Mus. Anatomy of Planorbis obstructus Morelet and Planorbis havanensis Orb. Cat. Amer. genus. p. The basal lamella in Planorbula is normally more massive and is. C. Type Planorbis liebmanni Dunker. Etudes Moll. Nautilus. XXI. Moll. With our present limited knowledge. . Adams). Example Planorbis orbiculus Morelet. idtradextral. p. 49. p. 43. it seems best to regard Taphius as a genus related to Tropicorbis. Conch. Gyrorbis Fischer and Crosse. LIII. Adams). referred to Tropicorbis. p. Zool. ]>. Planorbis liebmanni Dunker (= Pkmurbis orbiculus Morelet) 1880. Tropicorbis Germain. Example Planorbis heloicus Orb. 91. accompanied by a good plate. Proc. is given by Pilsbry (1924. Cent.80 The Molluscan Family Planorbidae Pilsbry (1934. Nat. \). et Guat. Genus TROPICORBIS Pilsbry and Brown. Baker. As subgenus of Planorbis... XXIV. 49. No. Oc. p. Oh^tructio H. p. Pars 22. Hist. 47) associates Taphius with Helisoma and refers Taphius pronus to Helisoma together with caloderma. Of medium size. Gyrauhis (Tropicor210. somewhat overlapping on the left (spire) side. p. Baker. 7.. Tropicorbis H. 1936. Menetus Tryon (non H. Acad. cit. IV. p. Shell. 1930. 8. 106.. 1914 Type by original designation. Pilsbry. 1939. Includes Planorbis obstructus Morelet and Planorbis dcntiens Morelet.artens species now (part) (non H. Nat. Tropicorbis F. As subgenus of Gyraulus. 9). An excellent account of the shell variation of Taphius andecolus. Zool. 52. in some species. 2. terr-fluv.. p. Ser. Tropicorbis F. 1833). Type by original designation Planorbis janeiremsis Clessin. and four lamellae within the outer lip. Univ. figs. 7-9 orbiculus) or the body whorl may increase rapidly in diameter (plate 77.

figs. These are attached to the upper j^art of the neck. When in locomotion the foot is rather long and narrow. GENITALIA. fig. ) ANATOMICAL CHARACTERISTICS PLATE 12 (Tropicorbis havanensis) . 4). 7. The verge (V).Sub fcun Hies. The vas deferens (VD) is moderately long (3. 3. This apparently takes the place of the papilla in Gyraidiis. as dot-like markings (plate 70. D. Male Organs The penial complex (fig. The sperm duct is over 5 mm.5 mm. (fig. Genera.) and bears about fifty short. havanensis (Pfr. The I'jrostate (PRS) is very long (3. CF). It is peculiarly enlarged at the junction of the oviduct and sperm canal and might be considered a carrefour. which have small branches. rounded before and tapering to a point behind. and are particularly plain when the animal is removed from the shell (plate 12. havanensis). CF. These are placed on a separate duct. long. the larger vesicles being nearer the ovotestis. A cross section through the prostate (fig. 4. These sometimes show on the base. black. blackish or brownish. fig. through the shell. 9. ends in a . T. 8. There are several series of muscles on the opposite side of the penial complex which we have called supporting muscles (SM). there being three branches all conis . and even the tentacles may be drawn backward.) and is a small tube throughout its length except that it is slightly enlarged where it enters the vergic sac (fig. rounded. as there is in Gyraulus. fig. VD). These may be what we have called supporting muscles. PD. the prostate duct. markings which show plainly through the shell (plate 78. and conspicuously placed at the inner base of the tentacles (plate 78. 4. When at rest (plate 70. and Suhijeuera. The vergic sac is always of less diameter than the preputium. figs. sac-like diverticula. Tropicorbis nigrilabris (Lutz) ). long and is very narrow. very long and cylindrical. but there is no second ring below this ring. They vary in size anteriorly. fig. It is beset with numerous protuberances or glands many of which have the form of club-shaped diverticula. There is a heavy muscular ring or diaphragm which separates the preputial cavity from the vergic space (fig. 11. the preputium (PR) cylindrical and slightly longer than the vergic sac ( VS) In another specimen (fig. See figs. 2). There are also several small muscles below the retractor muscles which are attached to the mantle and not to the columella muscle. T. The Tropicorbis group is noteworthy for the peculiar pigmentation observed in some species. 1) the preputium has two or more pilasters. The upper set. The whole seminal vesicle is about 2 mm. this consisting of dark.5 mm. 10) the preputium and vergic sac were more nearly equal in length and the upper part of the vergic sac was slightly enlarged. 1. nected by cross muscles. the free portion of which is very short and ciuickly joins the vas deferens.-Recent and Fossil 81 Animal. may also help in retracting the penial complex following coitus. Tropicorhu havanensis) the foot may be hunched up. The retractor muscles (RIVI) are somewhat complicated. SPD) shows the relationship between these two separate ducts. 11) Seminal vesicle (SV) of greater diameter than the ovisperm duct. The tentacles are very long and filiform and the eyes are large. 3. but that feature is attached to the female system and is a small swelling into which the albumen duct enters (fig. The velar area is well developed. 5. Internally (fig. 1. 11) is much elongated. 4. ).

havanensis and obstructus as figured in this volume. Some of the diverticula of the prostate are folded over at the end in riisei (figs. and ovisperm duct.) Hermaphrodite Organs. Tropicorbis nisei (Dunkerl (plate 13. the lobulate seminal vesicle. are shown in figs. shown = . in the long. nidamental gland. 3). together with the albumen gland connections.82 The Molluscan Family Planorhidae small papilla and the sperm canal has a central outlet (fig. Two other species of Tropicorbis have been dissected by the writer. may bear as many as fifty diverticula {havanensis) or as few as twelve diverticula (paUidu. a related species. and sperm duct. This figure differs in a few respects from the genitalia of riisei. near Santa Alarta. 3). p. Pilsbry (1934. long and smaller in diameter than the nidamental gland but much wider than the sperm duct. one loop of which passes through a trough-like space on the under side of the albumen gland. The verge appears as in the other species studied. The albumen gland is short. sperm duct. The spermatheca (S) is pear-shaped and less than long. 6). The 1 mm. It lies over the intestine. The retractor muscle complex is complicated in riisei but quite simple in obstructus. Female Organs. It is believed to be like havanensis. There is complete agreement. The penial complex shows a very long and narrow vergic sac. however. the long and narrow oviduct. 9) has twenty-four diverticula on the prostate and Tropicorbis obstructus (]\Iorelet) has eighteen diverticula on its i)rostate. It is desirable that the genitalia of the type species {orbicidus liebmanni) be known. The junction of the prostate duct and the sperm duct to form the vas deferens is in fig.s) The penial complex may have the vergic sac shorter than the preputium {havanensis) or much longer than the preputium {pallidus). 53) has figured the genitalia of Tropicorbis pallidus (C. In obstructus a number of the diverticula of the ovotestis contained eggs almost ready to be discharged (plate 13. fig. from Bolivar. 8 iriisei). 7 and 8. The duct is a narrow tube. particularly. long and narrow (about 3 mm. considerably longer than the preputium. It is evident from the knowledge afforded by the four species of which the anatomy is known that there is considerable variation in the details of structure of the genus Tropicorbis. The spemiathecal duct is longer than the spermatheca. Adams). narrow verge with tenninal oj^ening. is uterus (U) narrow and slightly longer is The nidamental gland (NG. 4. Hundreds of shells of orbiculus have been seen but no material containing the animal has been available. There is one retractor muscle as in obstructus. fig. The walls of the vergic sac are thin but those of the preputium are rather thick. than the spermatheca and its duct. The ovisperm duct is very short between the seminal vesicle and ovotestis but three times this length between the seminal vesicle and the oviduct. The penial complex is about the same in the three species. long) and of larger diameter than the uterus. and the rounded or pear-shaped spermatheca on a long duct. and enters the short vagina (VG) near the female opening. Colombia. The prostate. The spermatheca is rounder and less pyriform in riisei and obstructus and the ducts are longer than in havanensis (compare the figures). somewhat pear-shaped with large gland cells (fig. . The junction between the oviduct. very narrow. The ovotestis (OT) consists of many clubshaped diverticula placed in a double row. B. There are twelve long and somewhat irregular diverticula on the prostate. 5). The oviduct (OD) is about 2 mm.

fleshy organ doubled or folded upon itself. The outer three marginals are vestigial and are without denticulations. the pseudobranch is figured as much elongated. and Subgenera — Recent and Fossil 83 Respiratory and Renal Systems. and four to six or seven small cusps on the outer edge representing the ectocone. There is a large blind sac. the reflections becoming very long and narrow. 11). 53. . with no trace of a superposed ridge' (p. The 7th tooth is an intermediate. This appears to be a pathologic condition. Baker has described the jaws of pallidus and kiihnianus (1930. 4 is a thin. In his figure C. fig. longer than the buccal sac. 12). ending some distance above the anus. The kidney (plate 46. Pilsbry's figure of the pseudobranch of pallidus (1934. Stomach region with distinct division into crop. Pilsbry says 'The kidney is long. with an entocone of four or five small cusps. The jaw of riisei (plate 50. 12) shows the kidney to be rounded with the lumen rounded.). 13) shows that in this species this organ is much more compressed and much wider than in riisei. The jaw of obstructus is similar to that of riisei (fig. leaf-shaped. the cusps spade-shaped and reaching to the lower margin of the base of attachment. The buccal sac is pyriform with a large and conspicuous radula sac which protrudes from the posterior end of the buccal sac as a rounded projection. The salivary glands are cylindrical. fig. as in other groups of the Planorbinae. These are enlarged at their junction with the superior jaw and taper to a point at the lower end. and two lateral pieces which assume the form of a question mark. p. The rectum (Rl lies above the pseiidobranch and has a conspicuous crested ridge extending nearly its whole length. 11. the entocone broken up into four small cusps and the ectocone with two small cusps on the outer edge above. Digestive System. 47. and joined behind. 52). The digestive organs are alike in all species examined.5 mm. A cross section near the middle (fig. There is a large pneumostome (PS). A cross section of the kidney of havanensis (fig. the entocone larger than the mesocone and ectocone. Laterals (1-6) squarish with three cusps. fig. The jaws of Tropicorbis are unlike those of the other genera of Planorbinae (excepting Australorbis) and resemble those of the subfamily Helisomatinae. The marginal teeth become narrower toward the edge of the membrane. The radula formula of havanensis (plate 68. and pylons. pp. Of pallidus. The pseiidobranch (of havanensis. fig. the central lumen is irregularly oval with the two veins placed below the lumen at the outer ends of the section. 10. the two veins being at the lower corners of the section.Subfamilies. 8) is 19-1-19 or 20-1-20 with 109 to 112 rows of teeth. There is no ridge on the kidney but there is a distinct vertical ridge in the mantle to the left of the kidney. 48). 10) plate 12. finely striated along the face. The 6th tooth has a small interstitial cusp on the entocone and two very small cusps above the ectocone. fig. The crest is present in all species examined. a wide and low superior part. The 8th to the 16th or 17th teeth are marginals (8. 12) is composed of three pieces. B. This sac is longer in obstructus than it is in riisei. band-like. riisei) is rather long (about 2. the rectum ending near the pseudobranch. Genera. H. 1 the folded nature of the pseiidobranch is better shown as is also the crested ridge. In riisei (plate 13. a single spade-shaped mesocone. B» shows a simple folded pouch. the ureter sharply reflected. The intestine makes a backward loop around the liver and extends forward. The central teeth of the different rows vary in width at the upper gizzard. The central tooth has a broad base of attachment and is bicuspid. and flat.

In obstructus (plate 68. The radula of kiihncrianus is described on page 47 of the same paper. The formula is 18-1-18 with five laterals and there are 106 rows of teeth. .84 The MoUuscan Family Planorbidae part of the tooth. it would seem that its place is with the Planorbinae. 5. The nature of the prostate. Collected by Dr. p. Pilsbry (1934. C. The center tooth varies in width as in the other species. with many diverticula in a single row. modified only on the entocone. Oligocene or Lower Miocene to Recent time. The jaw is quite differ- ent and resembles this organ in Helisomatinae and Planorbulinae. as in havanensis. C. of the islands of the Geological Distribution. All central teeth examined exhibited this variation in width. E. and kiihnerianiis. The formula is 19-1-19. obstructus. Thus the radulae of five species of Tropicorbis are known. Faust. upward of thirty-three species and races being now recognized (see a later page for the list of species). which is nearest to orbiculus i^licbmanni) the type of the genus. Geographical Distribution. The penial complex differs from that of Anisus and Gyraulus in lacking a penial stylet. 12 specimens obstructus. it outranks any other group of the family found in America except Hclisoma. 7. a distance of about 7. The first thirty-four rows measure 5. The teeth of the radula also differ in having interstitial cusps on the intermediate and marginal teeth. 10 specimens riisei. Hoffman. In riisei (plate 68. Barceloneta. In number of species represented. fig. fig. The group is also found in many West Indies. New Orleans. Collected by Dr. The outer marginals are vestigial.000 miles. plate 28. From Louisiana and Texas in the United States. W. Baker (1930. B. These cover fairly well the different variations of the species of the genus. 7) the formula is 19-1-19 and there are 124 rows of teeth. havanensis. . Seven to twelve are typical marginals with a variable number of small cusps. There are four lateral teeth. All points considered. 53) remarks that the group resembles Gyraulus but differs in the unarmed verge and the different jaw and marginal teeth. the largest number of characteristics agreeing with those of this subfamily. . riisei. E. but the teeth are somewhat smaller. rows 35 to 89 measure 6 microns and rows 90 to 109 measure 5. fig. 8) figures the radula of pallidus from Bolivar. Faust 10 specimens. The 15th to 18th teeth are vestigial. The center teeth vary in width as in havane7isis.4 microns. 6) the formula is 18-1-18 with 105 rows of teeth. There are seven tricuspid laterals. Collected by Dr. New Orleans. A. Colombia. H. The genus Tropicorbis is one of the most widespread of any group of American Planorbidae. the 8th tooth is intermediate. places the genus in the subfamily Planorbinae.4 microns. and there are nine marginals becoming narrower toward the edge of the membrane. The genus Tropicorbis presents some puzzling features of its anatomy which render its systematic position debatable. The kidney is also like other members of this subfamily. 6). Remarks. The material personally examined is as follows: havanensis. The center tooth is like that of havanensis but the cusps do not reach the base of attachment. it extends southward through Mexico and Central America into South America as far south as the Argentine Republic. Puerto Rico. with six laterals and 106 rows of teeth. on a separate prostate duct. pallidus. The 5th and 6th teeth are intermediate with interstitial cusps between the entocone and ectocone and small cusps above the ectocone (figs.

B.sis (Clessin) LATEORBIS pallidus (C. there are variations in the shell which seem to warrant a division of the genus into three subgenera or sections. The African forms may be placed in Afroplanorbis of Thiele. Type Planorbis orbiculus Morelet Shell orbicular. between the South American Tropicorbis and certain quite remarkable and has been emphasized by Pilsbry (1934. Plauorbis jancircnsis Clessin increasing slowly in diameter. 7. The species and races believed to group under the three subgenera of Tropicorbis are tabulated below. The basal lamella is also usually more massive in Planorbula than in Tropicorbis (plate 76. the enlarged. \\\u\e the anatomy of the several group forms is similar.. PIanorl)ulinae. These are indicated below. B. in a separate subfamily. Genera. B. Subgenus TROPICORBIS SS. witii Menctus. however. at least. original designation. Baker. Adams) pedri7ius (Miller) cano7iicus (Cousin) irigyrus (Philippi) may a (Morelet) havaneiisis (Pfeiffer) philipjnanus (Danker) heloicus (Orbigny) tepicensis (Martens) gracilcntus (Gould) peregriniis (Orbigny) chile nsis (Clessin) 77}eridaensis (Preston) denticns (Morelet) d('ntie7is ca7i7iarum (Morelet) obstructus (Morelet) ob>itructu>< donbilli (Tristram) obstruct)is auodontus (Pilsbry) dentifcrus (C. Adams) deciivis (Tate) paparyensis (F. 1939 Type by Shell orbicular. Adams Shell with whorls rapidly increasing in diameter. 7) whorls Subgenus LATEORBIS Type F. Aperture never with lamellae. New Subgenus F^lanorbis pallidas C. Subgenus OBSTRUCTIO Haas. fig. Adams) nisei (Dunker) straniiiieus (Dunker) pete7iensis (Morelet) isthmicus (Pilsbry) decipie7ii< (C. Tw^o African species (adowensis and pfcifferi) have been dissected and the genitalia are very similar to the same organs in Tropicorbis. This relationship is discussed at greater length under the description of the relationshij) is The species in Africa genus Afroplanoi'bis. body whorl notably Species Considered as Valid. Aperture with lamellae during some stage of the growth of the shell (plate 76. until more is known about other species of the African fauna. Subfamilies. the last whorl not notably expanded. figs. 54). 8. These lamellae have the same general position in the two groups but the lower palatal lamella in Planorbula points upward while in Tropicorbis it points slightly downward. C. has a very different form of genitalia which removes it from the Planorbinae and places it. p. TROPICORBIS orbiculus (Morelet) OBSTRUCTIO jnneiren. the whorls increasing gradually in diameter. and Subgenera — Recent and Fossil 85 Some of tlie species of Tropicorbis have been heretofore placed in the genus Planorbula because of the presence of lamellae within the aperture. Apertural lamellae never present at any stage of growth. 9). B. Baker) nigrilabris (Lutz) albica7is (Pfeiffer) bourcardianus (Preston) ce7itimetralis (Liitz) fieldi (Tryon) . Planorbula.

XII. Rad.. and A. Field Mus. Phil. XXXVII.\as. From the figure the diverticula appear to be arranged in fan-like manner. 9. composed of five main branches each of which branch again two to four times. Planorbulina H. It is about as long as the preputium and appears from the figure to be slightly enlarged at the end. 151. Animal. 55. Type Planorbis Type Planorbis sudanicus von Martens. 25. 179. Tvpe Planorbis boissyi P. p. 33. Africa. lips sharp. of few or whorls which may be closely coiled or rapidly increase in diameter. 3). 1940.. Ann. p. N. As genus. usually in the same plane as the last whorl. Type Planorbis p. Biomphalaria Connolly (part). V. 195 (non Haldeman). the name used being tl'Orbigny in 1826 in Foraminifera. XXIV. the glandular follicles short and projecting from the side of the duct. p. usually flattened on both sides. Ind. 1925. Royal Soc. there being no reference to this name in the work mentioned.. 480. Sperm duct a narrow tube about as long as the ovisperm duct and seminal vesicle combined. p. 2.. States that citations from Jickeli's paper (1874) are erroneous.. 1931 siidanicus 1855). thirteen diverticula showing at the upper or outer end of the prostate. 221.. 1939. 2. The spermatheca is an elongated pouch connected with the rather long vagina by a duct half as long as the spermatheca. 431. The vas deferens is a long. Male Organs (fig. p. Ajroplanorbis Thiele. Planorbula. Nautilus. There is but one series of diverticula but there is no indication of a separate prostate duct. Planorbulina was used by many figs. not much greater in diameter than the vas deferens. sudanicus von Martens. von 1934. Small to medium size. Handbuch. Female Organs. Acad. Seminal vesicle occupying about half the length of the ovisperm duct. (non Adams. Citation for Planorbulina Jickeli (1874) Fauna der Land und Siisswasser Mollusk Nord-ost-Afrika. p. XXI. aperture rounded. p. Ajroplanorbis pfeifferi (Krauss) ). Akad. apparently as long as the sperm duct and ovisperm duct combined. Sencken.. The . Dresden. p. XXXIII. 64. p. Gesell. 1939. Abh. orbicular. Planorbulina Germain. Natur. AIus. Not observed. 1855). 1902. 86. As subgenus of Planorbis. Notes.. The vergic sac is a narrow tube. As section of Planorbis.. Ajroplanorbis Ha. 100. and M. Rec. fine duct. ANATOMICAL CHARACTERISTICS PLATE 71 GENITALIA. Nat. The penial complex has a sac-like preputium with a narrow retractor muscle attached to the summit near the junction of preputium and vergic sac. p. Teil Martens. Prostate short.. siidaniciLs Anatomy 1931. 1923. Citation from Jickeli 1874. Martens Reg. 1936. Fauna Palaearc. of Planorbis pfeiffcri Krauss. 86 The Molluscan Family Plaiwrbidae Genus AFROPLANORBIS Thielc. Sci. Planorbula Connolly. Carol. part p. Ajroplanorbis Pilsbry. Proc. Planorbulina Haas. South African Mus. No. 1. Nova Acta Acad. Shell (plate 81. Adams. the aperture with or without lamellae. No type cited. H. Zool. Jugoslav.\s. Malac. Menetus Westerlund. 483. Type by original designation Planorbis Mcnetus Westerlund (non H. 120. 1885. Trans. Leop. The features of the interior of the penial complex are neither figured nor described. 221 ('pour genre Planorbulina)') la Planorbis alexandrinus Ehrenberg {Segmentina sousS.

54) calls attention to the similarity of the shells. 8) shows a wide gizzard. The seminal vesicle is placed nearer the ovotestis than observed in the species of Tropicorbis examined.) paetcli (Jickeli) is Geological Distribution. The group as Tropicorbis. The above anatomical notes are drawn from the papers of Connolly (1925) andPilsbry (1934). Pilsl)ry's figures of Planorbis adowensis Bourguignat (plate 71. Ovotestis apparently a double series of clubshaped diverticula. The retractor muscle is inserted at the junction of vergic sac and preputium. Internally (fig. the smaller number of prostate diverticula. The albumen gland is a large. a narrow and elongated pyloris. The pseudobranch of Planorbis pfcifferi (fig. p. figs. The oviduct is shorter than the uterus and rapidly narrows to meet the sperm duct. The anatomy of Afroplanorbis sudanicus (von Martens) is unknown. . 12) the preputium has three vertical pilasters and the verge is very long and slender. Genera. The nidamental gland is about as long as the uterus and is swollen to twice the diameter of the uterus. sudanicus (von Martens) pfcifferi (Krauss) alexandriensis (Ebrenb. The salivary glands form a short loop behind the buccal sac. 10) is more pyriform than that of pfcifferi and the duct is longer. and the intestine looped about the stomach. The ovisperm duct is very narrow and its free portion is about as long as the sperm duct. 10-12) are essentially the same in general generic characters as in pjeifferi described above. 11) shows a wide. Geographical Distribution. Pilsbry (1934. The chief difference between Tropicorbis and Afroplanorbis appears to be in the longer seminal vesicle of the latter. The branched diverticula of the prostate are believed to be attributed to faulty drawing rather than to real differences in structure.) Afrdjihniorbi. The penial complex (fig. saclike preputium and a slender vergic sac equal in length to the preputium. Connolly (1925. The prostate (fig. Not ascertained. but the shell so closely resembles that of Afroplanorbis adowensis (Bourg. a narrow blind sac. half as high as long. 7) appears to be folded as it is in Tropicorbis. He also dissected Planorbis adowensis (his figures are copied on my plate 71) and found the anatomy to be very much like that of Tropicorbis. The figure of the digestive system (fig. Species Considered as Valid.) adowensis (Bourg.'^ salinariDu (Morelet) Afroplanorbis (Bourg.) that they appear to be congeneric. elongate-ovate organ. and Subgenera — Recent and Fossil 87 uterus swells behind the vagina to more than twice the diameter of the vagina. Hermaphrodite Organs.) Afroplanorbis Afroplanorbis Afroplanorbis sudanicus tanganikanus bridouxianus (Bourg. and Mich. and the greater difference in diameter between the preputium and vergic sac. plate 8) figures the genitalia of Planorbis pfcifferi Krauss and his figure differs little from that of Pilsbry except in specific details. Afroplanorbis appears to be distributed over northern and central Africa and to include a number of species. probably as old Remarks. The spermatheca (fig. 10) shows about sixteen short diverticula which are unbranched.) boissyi (Pot. The following are the species believed Other species may be added when their Afroplanorbis Afroplanorbis Afroplanorbis Afroplanorbis anatomy becomes known. to be referable to Afroplanorbis. the greater size of the uterus and nidamental gland.Subfamilies.

disjunctis. 1886. laevigata. marginibus distantibus.). 1899.' Westerlund. firma. cornea (subtus paullo pallidior). is unknown and until that species has been examined anatomically it would seem unwise to use this name so inclusively. Type Planorbis libanicus Westerlund. C. Remarks. As section of Planorbis. Alaska Moll. as far as known to the writer. Teil 2. spira magna. spiraliter lineata. Testa supra late concavo-umbilicata. The close relationship between Tropicorbis of Central and South America and the African species here referred to Ajroplanorbis is. originally designated for 1902. supra late profundeque concavo-umbilicata. media (d. libanicus W. it would then take precedence over Ajroplanorbis which would become a synonym of Biomphalaria. alt. an interesting parallel in zoogeography. Westerlund mentions in the description of the species (libanicus) that it differs from the subgenus Menetus and that it should become the type of a new subgenus. smithi Preston. which he later calls Heterodiscus. As section of Planorbis. as suggested by Connolly (p. infra subplana. Handbuch. subtus pone suturam impressam obtusissime angulatus. 5-6. primi lenti. which is unfortunately preoccupied. sub lente distincte sitiraliter lineata. preoccupied. 83.88 The MoUuscan Family Planorhidae The African group would appear properly to take the name Afroplanorbis proposed by Thiele. pfeifferi. 170-171. Type Planorbis libanicus Wester- 'Shell n: t. preferably as a genus closely related to Tropicorbis. All of the information concerning this group name and its type species is given above. infra subplana. utrinque subaeque lata. Planorbis adowensis and P. Tj'pe Planorbis Westerlund. No specimens for figuring have been obtainable and no figure has been published. 14. 14 mm. Akad. Heterodiscus Dall. As subgenus of Planorbis. subaequalem formantes. ultimus major. quod a subgen. PI. 5^2-6. Malak. sub lente forti densissime at distincte spiraliter lineata anfr. Havniensis)..). 151. GeselL. 484j. Should the anatomy jirove to be like that of the two species here referred to the genus Ajroplanorbis. p. utrinquc orbiter subaeciua. p. 120. 'Hab. subcylindraceus. p. p. 55). Meneto differre A'idetur. A similar relationship is found in the Ancylidae and in some groups of land shells. 'Haec species forte typum novi subgeneris format. Non Heterodiscus Sharp. However. p. Genus SYRIOPLAXORBIS F. spiram magnum. lund. interiores utrinque perlente accrescentes. Mons Libanon (legit beat. as pointed out by Pilsbry (1934. Deutsch. p. Type Planorbis libanicus Westerlund. ultimus subcvlindraceus (typ. Nach. Westerluxd. the anatomy of the type of this genus. ad apert. Hetcrodiscus HETERODISCUS libanicus Insecta. It has no relationship with Planorbula Haldeman which is purely an American genus belonging to a different subfamily. Jugoslav. Heterodiscus Thiele. 483) Connolly uses the generic name Biomphalaria Preston for all of the African species once referred to Planorbula. subtus subplana. Baker New name for HETF]RODISCUS Westerlund. 479. supra convexus. In a recent work (1939. rotundatus. Rad. sub lente tenue distincte ocris.' Westerlund.. extrorsum paullo planulatus. nitida. 1905. basali oblique surrecto. spira magna. 1931. . pp. 5 mm. 120. convexi. Diam. Tropicorbis appears to be generically distinct from the African species and can not be included in either Ajroplanorbis or Biomphalaria. . apertura oblique rotundato-lunaris. Original description of Planorbis libanicus Westerlund: 'Testa magnitudine medisupra late profundeque concavo-umbilicata. Evers. anfr.

aperture gaping. As genus. Phil. which is now placed in the genus Afroplanorbis. Subdiscoidal. Acad. but shallow. Abh. Genera. as indicated above. (8). E. 120) suggests that it is a modification of the Planorbis adoiuensis type and might be placed in the same section of ( . 431. Mus. and Subgenera — Recent and Fossil 89 In Westerlund's list of the subgenera of Planorbis (1902. Monotype Planana albertensis J. Sci.a. Ge. 55.SubjainUies. Biomphalaria Thiele. Only the type species is known.. Proc. The habitat of Planorbis libanicus is not so far removed from the African region as to preclude the possibility that this species might group with Afroplanorbis. 120) 'The prominent character of this species is the deviation towards the left of the last half-whorl. H. labrum greatly receding below (Preston). p.synonym of Planorbis Miiller. LIII. 479. Ann. Africa. Only an examination of the animal of libanicus would positively settle this question. Biomphalaria Ha. plate Tyjie Biomphalaria smithi Preston. To the Mountains of the Moon. VI. Lake Albert Edward. Planaria albertensis J. Shell (plate 81. as adowensis and suflanicus. Bull. p. 86.' Viewed as a dextral species. last whorl very large. Natur. Type Biom- phalaria smithi Preston. some species. As . Nat. 1910. planulate with concave spire. 1827. The anatomy of the group is at present unknown. included in this genus by Haas (1936) appear to belong elsewhere. Type Biomphalaria smithi Preston. the type of Menetus being Planorbis opercidaris Gould. p.\ert. the type of Menetus is given as Planorbis boissyi P. Species Considered as is ultradextral. S. p. 26a. Mag.. is the sole Remarks. fig. ix. Biomphalaria Pilsbry. 121. Planorbis ruppelli Dunker.. As section of genus Planorbis. Species Considered as Valid. p. p. No. BIOMPHALARIA Preston. Pilsbry 1927. 4). N. p. Monotype Biomphalaria smithi Preston. N.. p. Biomphalaria smithi Preston representative of the genus here recognized. 1936. They do not have the characteristics assigned to Biomphalaria. Biomphalaria Pilsbry and Bequ. hence the new name Syrioplanorbis (Heterodiscus) would become a synonym of Afroplanorbis. S. 1776). Animal. figs. p. the last part of the whorl drops below the periphery as in some specimens of Gyraidus defiectus. As genus. Belgian Congo. 26. 120). 260. S. The . Planaria Pilsbry and Bequaert. Biomphalarin Preston.. 1927. Tvpe wrongly given as Planorbis choanomphalus von Martens. This genus is unknown anatomically and its true position in planorbid nomenclature can not be definitely assigned until the animal has been dissected. the \'alidity of the generic concei:»t is not wholly certain. 115. 1934. Moore (non Planarin Brown. 25. Planorbis choanomphalus Martens and Planorbis katangae Haas. 1927. op. Monotype E. Teil 2.sell. or Miiller. Genus Type by 1901. H. Sijrioplanorbis libanicus (Westerlund) is the only species attributable to this genus and. E. Of this species Pilsbry says (1927. umbilicus open. in a measure. Handbuch. 535. 1910 S)7iithi original designation Biomphalaria Preston Moore. Moore. Plnnorhia J. and M. 1931. Amer. Secken. The comparison of Heterodiscus with Menetus would indicate that the latter group was somewhat related to the African species and the Latin description would fit. p. cit. Geographical Distribution.

p. As genus Planorbina Thiele. 29-31. 47 lists Brazilian species of this species listed belong to Tropicorbis 1905. Moll. Type Planorbis olivaceus Spix (non Planorbina Hald. In both references by Pilsbry the suggestion is made that the shell might have been modified somewhat after the manner of Taphius.. Large. Genus Type by 1883. Type Planorbis olivaceus Spix (non Planorbina Haldeman. Alaska Moll. Its best place in classification at present appears to be near Afroplanorbis as a distinct genus. 55).. pars 22. Handbuch. guadaloupensis Sowb. long and is an enlargement of the ovisperm duct on which there are many elongated follicles placed laterally. Type Plaiiorbis guadaloupensis Sowh. The shells as figured by Pilsbry (1927. and Biomphalaria might bear the same relationship to Ajroplanorhis that Taphius bears to Tropicorbis. Cat. p. No. 1921. The shell is sinistral. In a later publication (1934. 1899. Acad. 1934. 1938. Univ. 1855). the two or three anterior diverticula single and unbranched and the three posterior diverticula bifurcate. Australorbis Scott.. reproduced on plate 81 of this work) are strongly suggestive of Taphius. Genotype Planorbis guadaloupensis Sowb. Zool. As section of Planorbis Planorbina Germain. Biol. (fig. 1930. the mantle mottled with brown or cinnamon. (non Planorbina Haldeman. Type Planorbis guadaloupensis Sowb. Proc.90 The Molhiscan Family Planorbidae the genus Planorbis.. 11. Shell (plate 77. Fossil. as examples Planorbina Dall. and A... As section of Planorbis Menetus Martens. p. 81. No. 6. in front of the seminal vesicle there are numerous nodes which are probably continuations of the seminal vesicle. The others. pp. B. 210. 1855). 1843). 55. Adams. Xotulae Naturae. and A. 43. 6. p. Some of the 1940.. 86. XXI. Mus. Mich. 1918. Papers. 1923. 1843). 121.. long) of the same caliber throughout. As subgenus of Planorbis Planorbina Wenz. Resembling Tropicorbis in general form. Oc. Australorbis glabratus (Say) ). The prostate is 8 mm. 54. 1934 original designation Planorbis guadaloupensis Sowb. The sperm duct (SPD) is a very long and narrow tube (29 mm. Mich. 390 (non H. eighteen in num- GENITALIA. 1843). (^ Planorbis glabratus Say).. Sci. Phil. figs. Teil 2. long and is composed of twenty-three or twenty-four diverticula. and Planorbis guadaloupensis Sowb. smoothish. de Conch. Contribuicao ao estudo do genero Australorbis Pilsbry. 10) The seminal vesicle (SV) is 3 mm. 1482. Baker. Male Organs . Zool.. Mus. p. Type Planorbis olivaceus Spix (non Planorbina Haldeman. Rec. (non Planorbina Haldeman. Ind. As genus and subgenus Planorbina H. As genus. of slowly widening whorls which are rounded or angular laterally (Pilsbry. 509. Amer. pp. The color or blackish. p. Cent. As section of Planorbis Planorbina Walker. Nat. Adams. behind and 4 mm. Martens gives no type but says subgenus is peculiar to South America and mentions Planorbis cumingia7ius Dkr. 94. 1843). 9. Type PI. p. p.. Xo. 480. Type by original designation P/a«or£)is olivaceus Spix (non Planorbina Haldeman. p. is Animal. For a distance of 2 mm. 84. 55) this relationship is again emphasized.. On p. grayish ANATOMICAL CHARACTERISTICS PLATE 9 {Australorbis glabratus) . 41. Univ. 1843). pp. Pub. Miscel. 1934. 1843). biconcave. and other genera. = Planorbis glabratus Say Menetus Flscher. p. AUSTRALORBIS Pilsbry. As section of Planorbis Australorbis Pilsbry. Man. Mus. (non H. 1931. As genus Australorbis Martins.

The spermatheca is pear-shaped (S) and the duct (SD) is narrow and as long as the spermatheca.5 mm. the secondary diverticula being of unequal length. There is no penial papilla. . cylindrical. and Subgenera — Recent and Fossil 91 Inwc three iiiaiii l)ranches from a single stem. It is about twice the diameter of the sperm duct and is about 18 mm. not in nudtiple series. A single branched diverticulum is shown in fig. In cross section (fig. Hermaphrodite Organs. A wide retractor muscle (RM) comi^osed of several smaller branches. A part of the uterus projects behind the nidamental gland for several millimeters. 5). The ovotestis is composed of multiple diverticula as seen in situ (fig. The very narrow vergic sac (VS) is 3 mm. long. There are several sets of muscles. There is a flattened muscular ring or diaphragm (D). The ovisperm duct (SO) is very long (17 mm.7 mm. heavy pilasters in the preputium tapering to a point at the upper part of the preputial space. There is a rounded carrefour (OF) extending from the oviduct to which the duct of the albumen gland is attached. All diverticula arise from the sperm duct in a single series. On the same side of the preputium there is a wide muscle about midway of its length and two smaller muscles near the male opening which appear to be supporting muscles since they are attached to the mantle above the penial complex. Several of the diverticula contained one large ovum ready for exi)ulsi()n. ber. saclike preputium (PR) about 4 mm. wide and of about the same diameter throughout its length. after coitus. long.) and is a simple tube except for the portion occupied by the seminal vesicle. Some of these may aid in retracting the I'jcnial complex . It tapers to a point and the sperm canal is placed at the center (fig. which is observed to be fan-shaped in section. as in Helisoma. Internally (fig. Genera. 6. 10. The verge (V) is long and very narrow. On the opposite side of the preputium there is a narrow muscle attached to the upper part near the insertion of the vergic sac. is attached to the ui:)per part of the preputium and to the columellar nmscle. The oviduct (OD) is a narrow tube extending from the ovisperm duct backward. The space for the passage of the intestine (IX) is quite conspicuous. long and 0. The i)enial comj^jlcx (fig. of less diameter than the sperm duct. The uterus is the same width as the vagina and gradually enlarges in diameter to meet the nidamental gland (NG) which is very long and wide (about 8 mm. It is not much greater in diameter than the vas deferens. 9) there are two wide. Both gland and duct measure a trifle more than 3 mm. long or slightly shorter than the preputium. Female Organs. The space between the pilasters is marked by many cross muscles forming a reticulated pattern (PL). and a very wide muscle or set of small muscles which are attached to the roof of the mantle cavity and are presumed to be supporting muscles (SM). The organ is not multiple in the same sense that Helisoma is. each branch again dividing two or three times. There is a rounded enlargement of the vergic sac at the end receiving the vas deferens. 10) consists of an elongated. in length. The vagina (VG) is very short and wide. The diverticulum enters the sperm duct directly and is not placed on a separate duct as in Tropicorbis.Subjnniilies. 1) upon which a number of short divertictila are attached (2). The albumen gland (fig. The vas deferens (VD) is a narrow tube about 10 mm. 8) is rectangular in form and is composed of large follicles. OT). wide). in length. 4) the ovotestis appears to consist of a main diverticulum (fig. 1.

sac is short and high. 2. 1) is long and narrow (16 mm. somewhat as in Tropicorbis. There is a large pneumostome (PS). The stomach region (plate 48. and projecting over the left margin. figs. The kidney (plate 45. especially in the form of the spermatheca. In section (fig. 9). Cross sections show several . p. the veins still disproportionate in size and the ridge very wide. wide). The pseudobranch rounded. The position of each section is shown at A. the gizzard rather rounded. It is finely striated on its anterior face. 13). In section C (fig. with the pulmonary vein (AP). The salivary glands are narrow and nearly three times as long as the buccal sac. With these exceptions the genitalia of glabratus {guadaloupensis) here figured are like the figures of the race christopherensis given by Pilsbry on page 57. flattened. 11) near the base of the kidney. B. fig. large and round. A frilled crest or ridge extends down D Digestive System. and the ridge wide. is almost as long as the preputium in the specimens examined. The ridge is a very characteristic feature in the kidney of Australorbis Respiratory and Renal Systems. A low. fig. E. Section E (fig. 10) agrees with the specimens personally dissected. The low ridge is seen at the left.5 mm. The form of the long ridge to the left of the kidney is shown at the left in this section (RD). though wide. 9) the lumen is somewhat smaller and there is the same disparity in size between the two veins. in fig. In section B (fig. 55). 7) through the pericardium shows a large lumen nearly filling the entire section. 9. The side jaws are shorter in comparison with the length of the superior jaw than in most species The buccal . C. containing many glandular diverticula. 10) the lumen is much smaller. then runs forward to the rectum which bears a plaited crest or ridge (plate 48. There are no veins. the veins flattened. The ureter is nearly 2 mm. The pericardium is 3 mm. wide ridge extends from below the pericardium to the lower end of the kidney. The radula sac or pouch is large and rounded. the specimens examined was not short. They are joined behind as in Helisoma. long and 2 ram. not flattened. 8) the lumen is smaller. 10) resembles that of Promenetus. long and is a narrow tube directed upward at a right angle from the kidney. 1. Jaws three in number as in Tropicorbis (plate 50. smaller and oviform. but low. The veins are very large. The ridge (RK). and projecting over the left margin. not much elongated. D. is higher than in section D. also.interesting features. the length being about three times the height. as Pilsbry found it. and. The vergic sac. the pyloris long and the blind sac rather short and placed as in Helisoma. fig. is rather long and the center of the rectum (R) which it crosses as a simple ridge to the anal opening (A) and extends across the pseudobranch to the lower edge. the spermatheca was not really adnate. the whole organ is seen to be flattened. more nearly equal. The intestine makes a loop around the stomach and another looji around the liver. p.. In section A (fig. The superior jaw is very high. The figure by Scott (1940. therefore. The side jaws are much shorter than the width of the superior jaw and are very narrow. long. wide. 92 The Molluscan Family Planorhidae The above description agrees The spermathecal duct in substantially with that of Pilsbry (1934. on the other side. rather squarish in shape. the lumen very small. on one side and the renal vein (RA).

) Subjar/illies. Species Considered as Valid. tricuspid. p. subequal. Venezuela. pp. the lower outer corners of the base of attachment expanded to the extent that the base is twice as wide as the summit of the tooth. Lateral teeth (1-13) asymmetrical. Scott (1940. are separated by a considerable space. In specimens personally dissected the marginals began on the 16th tooth in two specimens and on the 18th tooth in one specimen. Puerto Rico. This genus probably dates fi'om the Oligocene of the "West Indies and South America. 55) figures the radula of the race christopherensis which agrees in all essential details with the figures on plate 68. the mesocone remaining a large sharp cusp. and Puerto Rico. but the entocone splitting into three to five small. there are three to six very small cusps in addition. Puerto Rico. The ectocone is always separated from the mesocone by a distinct gap. formiUa is 31-1-31 to 33-1-33 with 182 rows of teeth (plate center tooth (60) is wider than high. 5) The ( Geographical Distribution. 68. p. oblique. The small cusps above the ectocone are placed higher up on the tooth than in other genera. These differences are unimportant. Wm. The material used for this anatomical study was received from Dr. The ectocone continues as a large cusp but above it. Hoffman and was collected at Lares. 10-12) figures the radula of specimens from Caracas. The outer marginals are very narrow and oblique and the cusps appear as minute serrations. raclula . The on plate 50 agrees with that by Pilsbry (1934. and extending below the base of attachment. the 15th to 17th teeth being intermediate in character.) Australorbis Australorbis Australorbis Australorbis Australorbis Australorbis glabratus christ ophcren. The Venezuela form with a formula of 45-1-45 may be representative of the race oUvaceus.fis Pilsbry glabratus oUi'accus (Spix and Wagner) glabratus refulgcns (Diinker) glabratus higubris (Wagner) glabratus blauncri (Germain) antigucnsis (Sowerby) itn 7)i A ustralorbis unis ( Liit z Australorbis bahiensis (Diinker) Australorbis tenagophilus (Orbigm-) or Miocene period Geological Distribution. Avstrcdorbis is a South American group (Pilsbry I. There are two sharp. A. and the Caribbean islands. showing great variation in the form of the cusps. It is known in South America from Venezuela southward to Argentine Republic. He gives the formula as 30-1-30 with seventeen laterals. subequal cusps arranged obliquely. the cusps sharp. the two groups of cusps. Pilsbry (1934. Even here. mesocone and ectocone. 57). . Marginal teeth 16-26) elongated. and Subgenera— Recent and Fossil figure 93 Planorbidae. ]\Iany names have been given to variations within this genus and consequently there are many synonyms. on the' margin of the tooth. of Genera. The following species and races appear to be recognizable: Australnrbifi ginbratus (Say) (= Quadaloupensis Sowh. however. spade-shaped cusps which extend below the lower margin of the base of attachment. The intermediate teeth (14-15) differ from the lateral teeth in having either an extra interstitial cusp between the ectocone and mesocone (14) or an interstitial cusp as in 14 and an extra cusp above the ectocone. The fig. Except for its occurrence in Haiti.

by Dall or Germain are well stated and do not need to be repeated here. p. Conch. 1875 Type herein nominated Planorbis calculus Sandberger 1875. and Planorbis cal- . No type 1905. nearly equal. Pilsbry 's discussion of the reasons for rejecting the types proposed for Planorbina. The kidney is of the Helisoma type. In this respect. Australorbis is apparently an aberrent genus of the family.. cited Land imd Slissw. the branching being on one side only. 56) that it might be considered a subgenus of Tropicorbis but this procedure would appear untenable because of the peculiar prostate and ovotestis as well as the ridge on the kidney. Pilsbry has suggested (1934. Pilsbry did not examine the ovotestis or the kidney. cited as examples Plaiwrbis loryi Coq. This characteristic. Australorhis is so peculiar in certain of its anatomical characteristics as to puzzle the systematist regarding its proper position in the classification of the family Planorbiclae.94 The Molluscan Family Planorhidae Remarks.. p. may show ancient affinities with the Helisomatinae. each diverticulum arises from the ovisperm duct as a single tube. In its narrow penial complex withit is related to Tropicorbis. The prostate diverticula are also attached directly to the sperm duct instead of to a separate prostate duct out penial gland different as is the case in Tropicorbis. The ovotestis is also of a multiform nature. der Vorwelt. Haldeman's Planorbina is now considered a synonym of Anisus. the group appears to be related to Tropicorbis and to be correctly placed in the subfamily Planorbinae. The ridged kidney is an exception in the Planorbinae. 81. in fanformation. the Brazilian species Planorbis olivaceus Spix and Wagner being designated as the type. Each diverticulum arises from the sperm duct as a single tube. selected Planorbis guadaloupensis Sowb. Anisop. as type of Planorbina. culus Sandb. in 1921. Alaska Moll. Genus ANISOPSIS Sandberger. for which no type was designated by Haldeman. Germain. 958. So also in the ovotestis. as well as the apparent multiple appearance of the prostate and ovotestis. The name Planorbina Haldeman was used for this group by Dall in 1905. neither olivaceus nor guadaloupensis answer to Haldeman's description 'whorls numerous. with a distinct ridge.sis Sandberger. and also the form of the shell (with overlapping whorls). The prostate. however. In spite of the multiple nature of the diverticula of the prostate and ovotestis. In the Helisomatinae. excludes it from the subfamily Segmentininae. the branching taking place after the diverticula are above the sperm duct. both the prostate and the ovotestis have the diverticula radiating from the ducts like the spokes of a wheel. But as Pilsbry states in his description of Australorbis. When the characteristics of the prostate are closely examined it is seen to be uniserial in form in spite of the branched nature of the diverticula.. Anisopsis Dall. the few diverticula in this group being attached directly to the sperm duct. the prostate of Australorbis resembles that of the genus Drepanotrema. is very from any member of the Planorbinae and superficially resembles some of the Helisomatinae in the presence of multiform diverticula. The absence of a penial gland removes it from Helisomatinae and Planorbulinae and the absence of a flagellum.' the whorls in these species increasing in diameter to such an extent that the last whorl is notably wider than the preceding whorl. p.

ultimus permagnus. with a distinct. gracilis. 10 ». It also appears to have been comparatively unnoticed by recent students of the Mollusca. Planorbis loryi Coquand (Sandberger. p. 81) suggests its relationship to Helisoma but the left side is different from any species of that genus known. Sandberger suggests its resemblance to Planorbis complanatus (Linn. prima partem su})eram excavatam. The name Anisopsis appears to be a nude name.' These localities are in the Upper or White Jura. who states that the figure is much enlarged. Cajac. The only valid reason for associating the two species mentioned above with Anisopsis is the presence of the stars in the Aliddle ]\Iittlerer) Jura and the Upper (Oberer) Jura in the table. p. les Rousses. suturis tenuibus profundis disjiincti. as far as the writer is aware. The original reference is in a stratigraphic table on page 958 of Sandberger's work in which Anisopsis appears between Bathyomphalus and Anisus. lacking the distinct carina and deep umbilicus of the American Valvata. Remarks. Taf. and Planorbis calculus Sandb. 7a (stark vergrossert). spiram eminens (Sandberger. 1. fig. gracilis.Subfamilies. 'Testa calsuperne excavata. increasing rather rapidly in diameter. The second species. From the figures there appears to be a carina at the periphery. . layers e and Horizon and Distribution. Sandberger's description of this species is as follows: 'Testa calculiformis. The only American reference observed is that by Dall in 1905. culiforinis. su])ernc profunde excavata.' The horizon and : localities for this species are given as follows by Sandberger (p. Middle or brown Jurassic period. The group Anisopsis. Very rare. Apertura trapezoidea. The figure of this species does look much like Valvata tricarinata on the right side but on the left side the figure is too wide. p. and Subgenera — Recent and Fossil 95 Shell (of Anisopsis ccdcidus (Sandberger) plate 81. 14). The exact size is not given by Sandberger. ad margineuni sui)erum necnon ad iimbilicuin carina cincta. Dall (1905. inferne depressa. Genera. It is also larger. 11). no type hav- ing been formally designated for it. sharp carina in the middle of the whorls on the right side. Charix bei Nantuar. inferne latissime uinbilicata.' The shell is ultradextral. 7. Alferme und Vigneules am Bieler See in den gleichen Schichten. ( Planorbis calculus Sandberger occurs first on page 14 and should be taken as the type of Anisopsis. quadrangulates. fig. may not be congeneric. as examples. latissime umbilicata. but the figures do not resemble either of these species. The left side also has carinated whorls. Anfractus 3^/4. which may be designated as a genus of the subfamily Planorbinae. The whorls are few in number. superne excavati. 41) 'Villers le Lac (physaschicht b). may The two species referred to this genus be retained to contain these early fossil species. costulis trans versalibus subtilibiis ornati. indicating that these two species are the ones intended for this group name. loryi appear- . costulis transversalibus tenuibus muniti et carinis duabus acutis insignes. Anfractus quinque. On page 15 Sandberger mentions the likeness of this sj^ecies to the tricarinate Valvatas of North America typified by Valvata tricarinata. since no others from these horizons occur in the body of this work. 41) is more planorboid in form and somewhat different in shape from calculus (see plate 81.. who cites Planorbis loryi Coq. valde iniquilateralis. fig. altera basin cingente.) (=jontanus Lightfoot) and Planorbis carinatus Miiller. in f.

Synop. Brit. p. Vol. Man. both American and European. The radula teeth are modified by additional cusps or by the presence of many interstitial cusps. Trochorbis. Segmentina x A . With short flagella: Scgme7itina. Lapsus for Segmentina. p. 1817 XII. and Platijtophius. 337. genus of Planorbis Segmentina H. p. Berlins. Hippeutis. The prostate Type genus Segmentina Fleming. p. Monotype. No type cited.'\ldem. Drepnnotrema. Helisoma. 1842. Proc. Treat. Intha. Without penial gland: Polypylis. is Genus Monotype Xautilus 1817. Hippeutis. Isl. 137 Hemithalamus 'Leach' Turton. Zool. 1847. of course. is the only sj^ccies unciucstionably assigned here.. Sandberger. Mon. Without lamellae or barriers within the aperture: Hippeutis. With both long and short flagella: Drepanotrema. 509. be placed by shell characters alone. Miiller Hemithalamus Fitzinger. Syst. New diverticula are simple sacs placed in a single row along a prostate duct as in the subfamily Planorbinae. Schnecken Hippeutis u.' p. Ill. Type Nautilus lacustris Lightfoot. As sub- = ed. Helicorbis. SEGMENTINA Fleming.. Pingia. No type cited.ay. VII. With long flagella: Pingiella. Polypylis. Zool. Miiller Segmentina Fleming. Brit. p. Conchology. The assignment of Planorbis loryi Cociuand seems indefensible. Moll.. Drepanotrema. 1819. Subfamily SEGMEXTININAE F. Land Fresh-water Shells Brit.\n. penial gland and an external duct: Intha. 78. Trochorbis. p. 1850. 110 Segmentaria Sw. F.\inson. Edinburgh Encyclopedia. F. Fresh-water Univ. (proofs). U. Such are Helicorbis. 1847. 509. Moll. 1840. Miiller Segmentina Fleming. With lamellae or barriers within the aperture: Segmentina. Acrorbis. C.ach. Verz. lists the subgenera Menetus. p. As synonym of Seg7)ienti7ia Segmentina (part) Stein. Encyc. 'Conchology. groups which we now know are confined to America and have no representatives outside the Western Hemisphere. The groups included in this subfamily With With are as follows: penial gland but without duct: Segmentina. 1833. p. Musch. Pingiella.. The jaw is fragmented as in Planorbinae. Type Nautilus lacustris Lightfoot Hemithalamus Le. The penial complex has one or two flagella placed at the end of the vergic sac and in addition several genera have a penial gland in the preputium. Proc. Baker. In synonymy of Segmeyitina nitida 0. 1818. All of the extinct Planorbidae. the genotype. 116. 14. Platytajyhiu. The kidney is without a ridge. Acrorbis. Nautilus lacustris Lightfoot Planorbis nitidus O.96 The MoUuscan Family Planorhidae ing more like some of the species of Planorbis or Anisus. Supplement to 4-6th ed. Soc. Intha. lacustris Lightioot ^^ Planorbis nitidus O. are badly in need of a thorough taxonomic revision.-^. for example. Anisopsis calculus (Sandberger). F. As subgenus of Planorbis Segmentina Gr. Several groups have not been examined anatomically and their position doubtful. 309. 1831. The fossil group Paraplatiorbis must. Soc. IVIany of the names of groups of the Planorbidae have been used rather loosely by European writers. As genus Discus Gray. and Carinifex among the European fossils. Polypylis. Malac. S. Acrorbis. Species Considered as Valid.

1915. 121. Monogenotype Nautilus lacustris Lightfoot {^Planorbis iiitidus Miiller). Typo Planorbis nitidus Miiller. 163. Type Planorbis nitidus Miiller. Zool. Syst. 128. Zeits. Ind. Brit. Hist. Amer. The embryonic whorls have spiral rows of small pits. Cat. 85. p. 140. T. Trochorbis. Univ. LIII.'VLL. and Pelec. 2. 1661. Tj^pe Planorbis nitidus Miiller.. Type Segmentina nitida (Miiller). 1931. The internal lamellae are composed of enamel-like substance. Pub. ed. p. 107... Type Planorbis nitidus Miiller. p.. \).. p. Nat. whorls few. H. Akad. Mus.aas. and palatal.. Soos. Xo. 97. and Polypylis Segmentina L. As genus Scgmentina Binney. 1884. 89. Heft 6. 509. II. Museo Cien.. Type Planorbis nitida Miiller. Naturw.Subfamilies.. Includes species now referred to Helicorbis. 1931. SegmcHti)ia — Recent and Fossil Fluv.. As subgenus of Planorbis Segmentina Dall. As genus Segmentina Walker.ale and Prash. Xo type cited. p. Type Phnwrbis nitidus Miiller. figs. 4). N.. As genus Segmentina We. XXI. parietal. XVII. Fluv. 97 1865. p. Rec. Ill. XIII. Tj'pe Planorbis nitidus Miiller. 1926. Moquin-Tandon. p. Fauna Pal. Reg. Jugoslav. Syst. pp. II. 13. Moll. France. fiir Mollusk. II. Conch. Genotype Segmentina lacustris (Mantagu). Mai. As genus Segmentir^a Germain. As p. Type Planorbis nitidus Miiller. 1912. As genus Segmentina Thiele. 388. and Subgenera 1855. Soos.. Type evidently Planorbis nitidus Miiller. Shells N. Alaska Moll. 1885. No type cited. Rad. Man. p. As genus Segmenlina Fi. Mich. 136. Hist. Ficinus. XV.) nitidus (Miiller). 1883. As subgenus of Ajiisus Segmentina A. 542.. As genus Segmentina Hannibal. 481. 6.ype Planorbis nitidus Mi'ill. p. Allat. Rec. As genus Segmentina Wenz. As genus Segmentina Annand. 1935. Archiv. 142. Moll. Ann. As genus Segmentina H. T. p. 1923. Fauna Brit. causing the preceding whorl to project far within the width of the aperture. 424. FW. Type Planorbis nitidus Miiller. As subgenus of Planorbis Segmentina Preston. and F-W. 1921. As genus Segmentina Westerlund. p. p. 1918. 151. Miscel. As genus Segmentina Kennard and Woodward. basal.. p.. notably overlapping the body whorl embracing most of the preceding whorls.\d. p.. pars 22. 363 1886. Barcelona.. X. Terr. Cab. 252.ype Planorbis nitidus Miiller. Mus. France. smooth. Handbuch. Type Segmentina lacustris Lightfoot. 1-3). 124. India. and S. XVIII. Appendicular id 1870. 1917. N. As genus Segmentina Pilsbry and Beqluert. A. H. fiir gesamm.steklund. 423. 58 year. Nat. As subgenus of Planorbis Segmentina Glessin... and glossy. ultradextral. Ind. Lye. genus 1867. Terr. Trab. 1927. 154. Ann. Y. p. 352. L.scher. As genus Segmentina Germain.. each set usually composed of three large transverse lamellae. p. XXXII. 1919. Soc. Musei Hungarici. Moll. Mus. IX. Type Planorbis nitidus Miilleir. Type Planorbis lacustris Lightfoot. Teil 2. 1929. Type Nautilus lacustris Lightfoot. Lond. fig. 1926. XXX. de Conch. As subgenus of Planorbis Segmentina Tryon. Cavity of the last whorl restricted by several barriers. 56. Fossil. Conch. 29.. p. 34. S. p. Type Anisus (S. p. N. Shell (plate 79. Proc. Non-Marine Moll. . As genus Segmcntina D. Mus. V. periphery rounded or carinated. (])art) pp. Bull. As genus Segmentina Lindholm. p. There may be other smaller lamellae (plate 76. Kozlem. 1902. Cast. Genera. 1905. Small.

Male Organs The verge (V) is long and very narrow. figs. There are two short. narrow flagella (FL) at the end of the vergic sac at the point of insertion of the vas deferens. The preputium (PR) occupies half of the length of the complex and is elongatepyriform in shape. The vas deferens is of medium length (about 3 mm. 2) the preputium presents some striking features. Internally (plate 2. This penial gland has a shallow cup. 3). fig. which joins the sperm duct to form the vas deferens. There is a single heavy ridge on one side (plate 2. enlarged at the base. sac-like. long. With a short foot.) with twenty -two to twenty-five club-shaped diverticula of varying lengths. The spermatheca (plate 3. the eyes sessile at their inner bases.5 mm. rounded in front. tentacles fihform. rather wide retractor muscle. There is a single. The vagina is narrow and about 0.) and of small diameter. The free portion of the prostate duct is about 0. PA) which is also long and narrow. the upper part swollen.98 The MoUuscan Family Planorbidae Animal.3 nnn. in length. The nidamental gland is very long (about 3 mm. This ridge merges into a long. sac-like glandular organ which is reflected toward the base or opening of the preputium (SB). fig. In the specimens examined the posterior end of the prostate duct is free of diverticula for about 0. fig. The color of the animal is brownish black much paler when immature. jiointed behind. very narrow pilaster which extends the whole length of the preputium from the external opening upward to the muscular ring (D) separating the preputium from the vergic sac. Sperm duct long (4.7 mm. (plate 3. fig. 1) is very long (about 3 mm.7 mm. about three times the diameter of the ovisperm duct with several protuberances near the posterior end. Female Organs. On the right side of the preputium there is a long. The oviduct is about half the diameter of the is in length. and is as long as the preputium. 1) which extends vertically up a large portion of the preputium and is in the nature of a pilaster (PL). The vergic sac (VS) is very long and narrow. A cross section through the prostate and sperm duct shows the relative position of these parts of the animal (plate 3. . fig. 3). 4 of plate 2. 2) is long and sac-like is attached to the vagina by a long narrow duct which is somewhat longer than the spermatheca.) and of small diameter. and envelops the uterus. placed on a separate prostate duct. At its distal end it bears a fleshy papilla (plate 2. 2) Seminal vesicle sac-like.). In cross section this gland presents the appearance shown in fig. the bottom of which is paved with small protuberances resembling pavement (plate 2. It is about 2 mm. Prostate moderately long (1. long and of larger diameter than the sperm duct. 5. as shown in the figures. almost tube-like. Edge of mantle thickened. fig. The large gland is free from attachment to the wall of the preputium and is connected only with the large left pilaster. fig.5 mm. The and uterus about twice the diameter of the vagina and is a trifle over 1 mm. 6. The spermatheca is constricted about midway of its length. the fleshy sides rolled upward. ANATOMICAL CHARACTERISTICS PLATES 2 and 3 . GENITALIA. but no supporting muscles. the sperm canal (SC) being in the center of the verge at the end. The penial complex (plate 3.). This papilla is an extension of the verge only from one side.

^larginal teeth (14-21) with the reflection very low and wide. it is a long (2 mm. It is similar in form to that of Planorbis. the reflection low and wide and placed high up on the tooth. Some small species. wide. nitida (Miiller). 2) consists of rela- than twenty) large club-shaped diverticula. The kidney (plate 47. and pyloris. Radula (plate 69. 23. fig. Poland. Respiratory and Renal Systems. Hermaphrodite Organs. but anteriorly. There is a conspicuous radula sac which projects notably from the rear end of the buccal sac. the mesocone a trifle larger than the three cusps on each side. below the middle) it is seen to be rounded. the salivary glands short. then runs forward to the rectum. few (less The buccal sac is much elongated. fig. 5. Geographical Distribution. The flagella are shown somewhat longer than those in the figure on plate 3. There are six teeth which may be called intermediate (8-13). Soos's figure (1917. 543. dagger-like and subecjual in size. Twelve specimens of Segmentina nitida were dissected. the short ureter reflected at right angles to the kidney. 44. smooth.SubjaniiUes. The pseudobranch (plate 3. They are placed in a double row on the ovisperm duct. which is oval in sha]K'. the ectoconic cusps a trifle smaller. the flagella being more like the figure on plate 3. 56) is practically the same. The central tooth is wdder at the lower edge of the base of attachment than at the upper margin of the tooth.). L. p. The albumen gland fig. The tively ovotestis (plate 3. The teeth are all crow^led on the membrane. by Westerlund and others. Germain's figure (1931. gizzard. The two short cusps do not reach the lower margin of the base of attachment. and folded. The stomach region (plate 48. The pneumostome is very large (fig. PS). 4. The ovisperm duct between seminal vesicle and ovotestis is very short. fig. some of them even overlapping. fig. as . The teeth are very uniform in size and shape. with the veins placed at the upper corners of the lumen. The outer marginals (22. Digestive System. 17) is the same.) and narrow. collected from marshes in a meadow in Czerniakow. 16) is very narrow. p. 23) are vestigial. since the cusps are a trifle smaller than those of the lateral teeth. Jankowski. elongate-ovate. fig. ribbonlike. There are seven cusps. The formula is 23-1-23 with 164 rows of teeth. and the few diverticula on the prostate. There is a short blind sac. fig. a suburb of Warsaw. In cross section (fig. 24) is horseshoe-shaped. The jaw (plate 50. the elongated spermatheca and duct. showing the few large ovotestis diverticula.5 mm. and Subgenera — Recent and Fossil is 99 rather nidamcntal gland and about small and oval in shape. and composed of many small plates (about sixtv). 4) is rather small. Typical Segmentina is distributed throughout Europe and northern Asia. by A. Genera. 7cuspid. is recognized. Lateral teeth (1-7) long and narrow. fig. The figures on plates 2 and 3 agree in the main with those already published by European malacologists. 22) is very long (8 mm. with several races. betw^een the seminal vesicle and the oviduct. and looped behind. A single species. 1. 7). The intestine makes a sharp turn about the stomach and another around the liver. There is no superposed ridge. long. narrow tube. divided into crop. a trifle wider than high.

is the only species definitely considered. As subgenus of Planorbis 1850. Hist. As subgenus of Planorbis 1905. Cab. Helicorbis) As typified by Planorbis nitidus Miiller it is a well characterized genus. V.. As genus Hippeutis Westerlund. Xon Seg- II. Since the genus Segmentina is not found America. Conch. may be true Segmentina. Anim. the few large ovotestis diverticula. 119. France.^rpentier. pp. p. 1837 Type designated by Clessin in 1886. but also by its genitalia with the two short flagella on the verge. (Segmentina plus Hippeutis) Hippeutis Moquin-T. subgenus of Planorbis 1844. 426. planatus Drap. Hippeutis D. 82. Pknwrbis complanatus Drap. Leben. Man.. kempi Preston and angusta Species Considered as Valid. 22. London. p.. Hippeutis Clessin. Hippeutis H. and the papilla at the end of the verge. Fig. X. Hippeutis Gr.4Ndon. mentina Fleming. The genus Segmentina is distinguished not only by its glossy with the embracing body whorl and internal barriers. der Schweiz. 154. 151. Rad. Segmentina 7}iti(Ja (Miillerl.. The generic name has been used in a rather wide sense and has included one group which belongs in another subfamily {Planorbula) and several other related groups which are separated generically from Segmentina {Popypylis. Hippcutes M. p. or Helicorbis. p. S^vst. Moll. The Indian and Chinese species included by authors in Segmentina probably all belong to Poli/pylis.-vnnib. South African species may belong to an undescribed group.\nn.100 The Molluscan Family Planorhidae Jickeli. V. Land FW. the genotype. Allg. IsL. Naturw.. 2.\y. (= Mai. Type Planorbis jontanus (Lightfoot). all of the twelve specimens examined being more or less heavily infested. Genus HIPPEUTIS (Agassiz AISl Charpentier. 243. 51. As subgenus of Planorbis 1886. p. Fluv. Alaska Moll. pp. Neue I. 1885. in Remarks. Erd-und Siisswussser-Gast. Geological Distribution. in Turton. Type Planorbis jontanus (Lightfoot). Terr. Gr. Arrang. Species quoted from northern Africa probably belong to other genera. No type cited.. 87. Fauna Palearc.\ll. Proc. Pingiella. The ovotestis appeared to be most affected and in several specimens this . As Hippeutis Hartm. Akad. Oligocene period to Recent fauna. Type Planorbis comHelix jontana Lightfoot). p. de Ch. Type Planorbis fontanus (Lightfoot). 86. As subgenus of Pla^wrbis 1912. 2. as Ajroplanorbis. {= Helix jontana Lightfoot). The liver was also affected. As subgenus of Planorbis . Jugoslav. Conch. Muschl. Soc. 121. Berlins. organ was almost obliterated. Hippeutis Type Planorbis com- Westerlund. 2. E. Segmentina nitida appears to be a favorite host for parasitic worms.. p.. Shells Brit. 65. No. the ductless penial gland in the preputium. pp. 423. No mention is made of this peculiar gland in any w^ork consulted. IV.. Ed. Segmentina (part) Stein. Syst.\y. 78. one of the two species originally included 1837. The prostate is on a separate duct as in the Planorbinae. it is extralimital to the scope of the present work. Binnen. As subgenus of Planorbis Denksch. As subgenus of Planorbis 1902. Schneck. 84. Ed. Pingiella.ssiz' in J. pp. Hippeutis 'Aga. Typographical error for Hippeutis 1850. 34. Only an anatomical examination can correctly place the many species referred to segmentina. Moll. 1837. Schweiz. Reg. These special features of the anatomy distinguish Segmentina from all shell other genera of Planorbidae. I planatus Drap. 1855. XVII.

). LIII. Penial complex (plate 2. Heft 6. Genotype Hippeutis complaHelix jontana Lightfoot. As genus = 1926.7 nun. p. = No tj-pe cited 4-6). 1633. and Subgenera 1915. where the small depression may be 15) . 58 year. 6. No. 1927. Museo Cien. outer lip thin. Hippeutis 1923. As subgenus of Planorbis p. Sperm duct iSPD) long (over 1 mm. Ideogenotype Planorbis complanatus Drap. 69. the preceding whorl forming a sharp extended ridge on the parietal side of the aperture. which extends crosswise of the preputial cavity (SB). Type evidently Helix jontana Lightfoot. As section of Phuiorbia 1918. Small. 101 Type Prestox.) (fig. Pub. Fauna Brit. Allat. 8. Vergic sac (VS) shorter than the preputium. Fossil. sac-like penial gland. Handbuch. Planorbis jontanus (Lightfoot). II. figs. Ind. Zool. 1931. shining. Soos. Hippcutiii — Recent and Fossil Gast and Pelec. ) Seminal vesicle rather short (about 0. Archiv.Subfamilies. There is a single wide retractor muscle (RM) attached to the preputium at the lower part of the bulbous swelling.. body whorl very wide and overlapping most embryonic whorls have spiral punctures as in Segmentina.. Preputium (PR) strikingly pyriform. p. Teil 2. p. 12 (SB) and with the end stretched out in fig. Moll. As subgenus under genus Anisus Hippeutis A. Mus. The Shell (plate 79. Univ. p. Syn. As genus natus (Drap. It is shown folded in fig. 481. 9..) 1923. Type Planorbis complanatus Drap. norbis jontanus (Lightfoot). Type Helix complanatus Linn. p. tapering to a narrow neck near the outlet. As subgenus of Planorbis Hippeutis Thiele. France.. There is a depression at the end of the gland but apparently no cavity or cup. N. placed midway between ovotestis and oviduct. 8). P/«nor6zs jontanus (Lightfoot). Nat. Type Anisus (H. fiir Mollusk. lens-shaped. 12). India. Hippeutis Germ. the diameter of the organ about the same as the ovosperm duct. The body of the animal is clear and translucent. sac-like. XXXII. XIII. Hippeutis Lixdholm. 14) less than 1 mm. Aperture triangular. the preputium has two large fleshy pilasters (PL).ain. p. 253. Not differing from that of Segmentina in general form. Moll. Prostate short (about 0. Trab. Walker. 1935. p. pp. XXI. Fluv. ultradextral. Cat. of much smaller diameter than the preputium. but of smaller diameter. fig. As genus Hippeutis Germaix. Type Planorbis jontanus (Lightfoot). As subgenus of Planorbis Hippeutis ^as. 1929.) complanatus (Linn. Non-Marine Moll.6 mm. Male Organs . Pars 22. Mis. Mus. Amer. p. GENITALIA.. with many pustule-like vesicles. ANATOMICAL CHARACTERISTICS PLATE 2 {Hippeutis complanatus (Linn. 94. 541. H. There are no barriers or lamellae within the aperture. The square end of the gland is folded backward upon itself.) and of large diameter. Brit.) with ten long and narrow diverticula.) described. As subgenus of Planorbis 1926. Vas deferens (VD) about as long as sperm duct. Hippeutis Kenxard and Woodward. 124.. 159. Barcelona.. Kozlem. to the left one of which is attached a large. There are two short flagella (FL). the of the inner whorls and forming a sharp keel at the periphery. 387. Mus.. Mich. Animal. p. 1931. Rec. 128. As subgenus of Segmentina.. Type Pla- Hippeutis Wexz. Free portion of prostate duct short (fig. Genera. As subgenus of Planorbis Hippeutis Pilsbry and Bequaert. Internally (fig. long. 29. Bull.). FW. Type not cited but Planorbis complanatus (Linn. Terr.

fig. bulbous carrefour (CF) which receives the duct from the long and narrow albumen gland (AL. fig. wide vagina (V) by a long. 17. and shorter than the spermatheca and its duct. much swollen and about half as wide as long. (fig. and there . 12. PES). A few points. The intestine loops around the stomach and liver. . 10). The nidamental gland (NG) is longer than the uterus. however.102 The Molluscau Family Planorhidae plainly seen. which is twice the length of the spermatheca. ing in resi)ects Odhner most (1929. No mention is made of a separate prostate duct or of a gland in the preputium. Female Organs is and also fig. 13) figures the genitalia of this species agree- Respiratory and Renal Systems. The pneumostome is very large. are figured differently from the position observed in the specimens studied. The retractor muscle (RM) is attached to the preputium at the point where the gland joins the pilaster (fig. The ovisperm duct (SO) is very long (about 2 mm. Digestive System. cjuite different from the long.) and about the same diameter as the uterus. The verge is swollen above and tapers to a narrow diameter for about half its length (V). with is small. There is no superposed ridge. fig. The with the material personally examined. tube-like duct. The uterus (U) is twice as wide as the vagina. the lumen rounded with the veins placed at the upper corners of the lumen. there being no interstitial cusps between the larger cusps. 7). The two short flagella (FL) are hollow and under high power show a minutely punctate surface at the upper rounded end (fig. fig. The kidney (plate 46. 22. The pseudobranch (plate 2. the greatest width being almost one third of the length. ovotestis (OT) has about thirty clubshaped diverticula jilaced for the most part in a double row (fig. The oviduct (OD) is short (about 0.5 mm. ovate. sac-like joined to the short. fig. as well as others of this subfamily and of Planorbinae. SB). plate 46. There is a large. 32. plate 47). 17) is rather short (1. but extends horizontally into the mantle cavity. 15) The spermatheca (S) is long.) and of small diameter. Odhner's figure of the radula of Hippeutis complanatus differs somewhat from the radula as personally examined. An elongated object in the upper part of the prej^utium is shown in Odhner 's figure which might represent this organ. The formulas of Odhner's specimens agree exactly with ours. 13) and })ierced above by the rectum (P). rather narrow. There is a narrow muscular ring or diaphragm (D) between vergic sac and preputium. The ovotestis diverticula are shown extending in pairs from a short duct which enters the ovisperm duct. A cross section below the middle (fig. 12) is elongated. The stomach (plate 48. as in Segmentina. the ovotestis diverticula arise in pairs directly from the ovisperm duct. In this species.) and wide. each diverticulum entering the duct separately (see plate 2. The ureter is not sharply reflexed as in Segmentina. 11). Hermaphrodite Organs. 18) shows the kidney to be flattened. 15. fig. The kidney of Hippeutis is quite different from that of Segmentina (compare fig. The prostate is shown as having thirteen diverticula rather widely spaced on the sperm duct. fig.5 mm. 7). The sperm canal has a central outlet but there is no papilla. p. crowded diverticula personally observed (see plate 2.

From the Lower Eocene period (Wenz).) Hippeutis junodi (Connolly) Hippeutis benguelensis (Dunker) Geological Distribution. . Poland. They are joined behind as in Scgmcntina. The specimens were obtained by Mr. 16 km. narrowed in front with a large radiila sac behind. The kidneys of the two genera are quite unlike.) Hippeutis raymondi (Bgt. Geographical Distribution. Eighteen species and races of fossil Planorbis are referred to this genus by Wenz. The above anatomical information was obtained from specimens collected in a pool on a meadow in Jabtonna. The species cited from South x\frica. the mesocone large and almost reaching the lower margin of the base of attachment. Remarks. however. Genera. The whole genitalia of Segmentina are much elongated while these organs in Hippeutis are short and heavy. the group palearctic group.Subfmnilies. and China probably belong in other genera. They are not like those of Scgmcntina nitida figured on plate 69. The center tooth is higher than wide with two wide cusjis which do not reach the lower margin of the base of attachment. The following species are apparently referable to Hippeutis: Hippeutis Hippeutis Hippeutis Hippeutis complanatus (Linn. As in Segmentina a number of species have been described with many varieties. Species from northern Africa and eastern Asia have been assigned to Hippeutis but until these species have been examined anatomically this reference remains open to doubt. The jaw is fragmented as in Segmentina and is made up of many small plates. with 150 rows of teeth. The radula formula is 16-1-16. The penial gland differs in shape but both are attached to the right pilaster. Italy. 7. The Asiatic forms probably belong to Benson's genus Helicorbis. The entocone is shorter than the mesocone. The radula teeth of Hippeutis resemble those of Bathyomphahis contortus shown on plate 68. The cctocone is shorter than the entocone and is placed high on the margin of the tooth. Europe from Great Britain.) riparius (Westerlund) syraciisanus (Cafici) diaphaneUus (Bgt. north of Warsaw. fig. 1. The entocone and mesocone are of equal length and the ectocone is shorter with two small cusps above on "the outer margin of the tooth. Eastward to northern Siberia. It has been recorded from northern Africa and be circum-]\Iediterranean in distribution. India. A. In the marginal teeth the reflection is wide and low and is placed high u\) on the tooth. has fewer diverticula. The radula resembles Bathyomphahis rather than Segmentina. The shell is wholly without barriers or lamellae of any kind within the aperture and this characteristic at once marks Hippeutis as a separate genus. The buccal sac is elongated. Jankowski. A may Species Considered as Valid. The salivary glands are short and do not extend far beyond the posterior end of the buccal sac. and Sweden south to Portugal. and Subgenera is — Recent and Fossil 103 a larj^e blind sac. and Albania. The prostate. Hippeutis complanatus resembles Segmentina nitida in its anatomy. Internally the penial complex of the two groups differs in details. Parasites were not observed in any of the material examined. fig. Norway. The laterals (1-9) are squarish and tricuspid.

The embryonic whorls are punctured. 362. on a separate prostate duet (PD). Polijpylis Pilsbry. rather narrow.. Zool. Internally. Sci. as shown in the small figure in fig. fig. Type Segmentina (Dunker). and one or two in the upper margin. Mus. As synonym of Segmentina Mus. 10) is long and narrow (0. Animal.104 The Molluscan Family Planorhidae Genus POLYPYLIS Pilsbry.) convoluted. p. 96. 10. both included in Polypylis 1923. Polypylis Haas. XXII. rather narrow and elongated (1 mm. rather long (2 mm. consisting of twenty-eight to thirty-four long and narrow diverticula. Ind. Rec. Several of the diverticula are bifid at the end. 166. Rec. H. N. Proc. The parietal lamina is obliquely transverse. tube-like duct somewhat shorter than the sperm duct. basal long. The penial complex (plate 8. Mus. There is one large retractor muscle (RM) which is attached to the preputium near the junction of this organ with the vergic sac. figs. 1922. 4). XXIV... Not examined. As subgenus of Segmentina (part) Prashad (non Fleming). The vas deferens (VD) is a small. There are over thirty diverticula on the prostate of calathus and the seminal vesicle is very large with projecting vesicles. Type Segmentina largillierti (Dunker) (= <S. The narrow part of the flagellum has about the same diameter as the vergic sac. Nat. are very thick pads in calathus. narrow muscular ring or diaphragm (D) separating the preputial . p. hemisphaerula (Benson) ) 1906. The free prostate duct is very short. The vergic sac (VS) is two-thirds as long as the preputium and is very narrow. ANATOMICAL CHARACTERISTICS PLATE 5 Seminal vesicle (SV) almost twice the diameter of the ovisperm duct. There are several or many barriers (Pilsbry). 1906 Type by original designation Segmentina largillierii (Dunker)(= S. in the penial complex of hemisphaerula (plate 8. largillierti 1921. long). The vas deferens enters the vergic sac at one side of the flagellum. Notes. 7-9) less compressed than Segmentina (nitida) and not carinate.. p. Sperm cluct (SPD) about 3 mm. included in Polypylis Annandale (non Fleming). As subgenus of Segmentina Shell (plate 79. the others transverse. Male Organs in most of the diverticula (fig. 164. Field Germain. 7) . p. Prostate 1. Segmentina calathus (Benson).5 mm. GENITALIA. Mus. Miscel. Rec. p. There is a single flagellum (FL) which is long (over 1 mm. long.3 mm. The preputium (PR) is sac-like. There is a small. Polypylis 1939. fig. Segmentina Mentions Segmentina Mentions Acad. Ind. the tip of which is bent over (fig. but glossy with deeply clasping whorls. The vesicle gradually decreases in diameter to the ovisperm duct which is not sharply distinguished from the vesicle. a shorter one in the outer wall.. also. LVIII. XXI. 585. XXIV. Phil. with but few pustulose excrescences. hemisphaerula (Benson)). Planorbis calathus Benson and Planorhis caenosus Benson.) and narrow and has a bulbous enlargement at the end which is almost half as long as the flagellum. 8a. The pilasters. long). long. fig. In Polypylis calathus there are two short flagella and the vergic sac is longer than in hemisphaerula. Ind. 9) the preputium has two pilasters extending the whole length of the sac.

squarely truncated at the lower margin. B.'i (Benson). wider than high with the two lateral veins of large diameter. as in Segmentina. Collected by Dr. without a ridge. Burma. 24). . The oviduct (OD) is short (1 mm. The above anatomical details were obtained from the following material: Polypylu hemisphaerula (Benson) from Peipino.) and narrow. The uterus (U) is not much longer than the vagina.) and very narrow.5 mm. tube-like and at the end there is a fleshy papilla and the sperm canal has its outlet at the side of the verge just above this papilla (plate 5. from below Machligaon. It resembles the kidney of Polypylis calathus (plate 47. S. H. In calathus the formula is 26-1-1-26 with 205 to 210 rows of teeth.7 mm. Four specimens. segmented with many plates (plate 50. Polypylis tnia (Annandale and Rao). There is a large carrefour (CF). The jaw is like that of Segmentina nitida. The ovotestis (fig. 7). OT) consists of many very long and club-shaped diverticula placed in a double series. all 6-cuspid. 7. Peiping. 6) There is no penial gland in the penial complex. Mongyai. fig. Respiratory and Renal Systems. As many as thirty-eight pairs of diverticula were counted in one specimen. 1 plates. One specimen e. Female Organs is and Hermaphrodite Organs. the cusps short and the reflections wide and low and placed high up on the teeth. long) and of about the same diameter as the uterus.. fig. China. There is a large pneumostome. The pseudobranch (plate 5. The verge (.s calathu. The buccal sac is pyriform. The kidney of Polypylis hemisphaerula is very long (7 mm. Collected by "Dr.^^) is long (0. long) and ovate in outline (fig. 8). fig. Polypyli.xamined. fig. fig. Rao. in length. H. 16 but the blind sac is somewhat larger. 2). from weedv tank near inspection bungalow. A cross section is similar to that of calathus (plate 47.7 mm. and Subgenera — Recent and Fossil 105 cavity from the vcrgic cavity. The stomach I'esembles that of Segmentina nitida fig. 3) is long and narrow and soniewliat folded. 23) than that of calathus. The spermatheca (S) is rather small attached to the long and narrow vagina by a narrow duct (SD) about 1 mm. X.Subjamilies. (plate 5. long.) and much swollen (XG). Rao and Mr. Digestive System. The radula formula of hemisphaerula is 25-1-25 with over 200 rows of The teeth are like those of Polypylis calathus (plate 69. . Andamans. S. In outline the section of hemisphaenda more nearly resembles that of Segmentina nitida (plate 47. 24) which is long and narrow and 9 mm. with a large radula sac behind and two small salivary glands like those in Segmentina nitida. sent by the Fan Memorial Institute. fig. In hemisphaerula the lumen is rounder and the veins relatively smaller. The ovisperm duct (SO) is very short between the ovotestis and seminal vesicle and very long (2. One specimen examined. fig. teeth. the lumen oval. Shan States. Genera. 25) which shows the kidney to be flattened. China. The nidamental gland is long (2 mm. Chopus.) between the seminal vesicle and the oviduct. The albumen gland is large (1. The rows of teeth are all in straight lines. Port Blaie. fig. Polypylis calathics has a similar jaw with many small (plate 48. The vagina rapidly enlarges from its external opening to the point of juncture of the spermathecal duct (VG). H.

Its full distribution and the species belonging to it can not be fully known until more of the species from Asia have been anatomically examined. 105. being flattened with an acute periphery. Subgenus of Planorbis. 361. Hippeutis Ann. differs generically from Segmentina and Intha in the absence of a penial gland in the preputium. fig. 5). 1837). as suggested below. Example: Polypylis calathus (Benson). XXIV.. The lamellae within the aperture in Segmentina are three in number. Annandale (non Charp. Helicorbis Kennard and Woodward. Example: Polypylis hemisphaerula (Benson). is here nominated as type of the genus. No 1906. too few of the Asiatic species have thus far been examined anatomically to warrant separation into subgenera at the present time. p. p. Group II. The following species appear to be correctly placed in this genus: Polypylis Polypylis Polypylis Polypylis hemisphaerula (Benson) calathus (Benson) lucida (Gould) nitidclla (Martens) Polypylis swinhoci (H. quite unlike the wide space in the aperture of Polypylis (plate 76.. Brit. 106 The Molluscan Family Planorbidae Geographical Distribution. 1922.. p.. XXII. Species Considered as Valid. There is some variation in the form of the flagella in the species examined. Non-Marine Moll. and upper labial. Helicorbis Dall. Considered synonym of subgenus Hippeutis. Adams) Polypylis taia (Annandale and Rao) Polypylis usta (Gould) Geological Distribution. XIX. {kg. Handbuch. The species Planorhis (Helicorbis) umbilicalis Benson. figs. all whorls visible on upper (right) .) Charp. 4). The difference between the form of flagellum in the type sjiecies and the flagella of the other species examined from India and China might indicate a division into two groups. With one swollen flagellum on a long duct. Mus. Polypylis Group Genus HELICORBIS Benson. Polypylis is found throughout much of eastern China and India and on some of the islands lying off the coast of China and India. the body whorl wide and overlapping or embracing the previous whorl.-vnd. Treated as synonym of Hippeutis. while in calathus and taia there are two short flagella as in Segmentina. I. 1837). Hippeutis 1926. Shell (plate 78. all transverse to the whorl (plate 76. one of the original species listed. The space between the lamellae is narrow and three-cornered. Rec. XXIV. 1931. parietal. 481. 1837. 584. Rec. Syn. Places Planorhis umbilicalis in this genus. Ind. 69.\shad (non Charp. p. Considers group equal to Hippeutis 1921. 126.. Remarks. 1855 1855.. p. Mus. Mentions Planorbis iimbilicalis Benson as belonging in genus Hippeutis. Teil 2. Nautilus. narrow flagella without swollen end. 5-7) Ultradextral. The shell of Segmentina differs notably from that of Polypylis. . Helicorbis Thiele.^le and Pr. type cited. while the perijihery of Polypylis is rounded and the shell is high. Jour. With two short. Asiatic Soc. Ind. p. Not ascertained. basal. fig.. Polypylis taia (Annandale and Rao) However. Bengal. In hemisphaerula there is but one flagellum on a long duct. Helicorbis Benson.

584). Hippeutis complanatus (Linn. 'The radula is very minute and the dental formula is approximately 14-12-1-12-14(26-1-26): The central tooth is comparatively large and bicuspid with sharp cusps. 8. does not rei)resent that species but is a good representation of the new genus PingicUa F.. The periphery in these figures is rather well-rounded. subfamily Segmentininae are in need of careful revision. 15. The figures in Hanley and Theobald. Germain (1921. 7. figs. of the family. Just what species may be included in this genus is not definitely known. but differ in details. and 9 show the periphery as slightly carinated. The penis is massive with a lateral opening and without anv stylet' (Annandale and Prashad. 8. 1921. Taf. and the line of demarcation between the laterals and marginals is not very sharp. Clench' and Beciuaert (1939) give the best figures observed (plate 1. and 10) showing well the slight angulation of the l^eriphery. Baker. at least as far as the shells are concerned. 'The external soft parts generally resemble those of Gyraulus except that the branchial process and the pulmonary siphon are a little better developed' (Annandale and Prashad. 20. p. The radula is figured on page 583 of the work cited (fig. p. Not clearly known but including eastern and central China and India. 19. p. C. Genera. that Helicorbismearnsi Bartsch is a synonym of umbilicalis (see plate 2. however. 177) has called attention to the discrepancy in Clessin's figure and gives good figures of umbilicalis on plate II. 1921. Geographical Distribution. The vas deferens is very long and coiled. Lips thin. Radula. XVII. the spire depression forming The whorls are rounded or at most only subangulate. There are no internal barriers or lamellae within the aperture. Conch. The embryonic whorls are marked as in other genera a false umbilicus. figs. 6 . but — Recent and Fossil 107 mostly concealed on lower (left) side. Not ascertained. plate 40. Remarks. 585). The teeth are like those of Bathyontphalus and Hippeutis. Animal. the central cusp much the largest' (Annandale and Prashad). It is evident that the species referred to the The figure of Plajwrbis umbilicalis in Clessin. Cab. Species Considered as Valid. The penis sac is a well-developed ovoidal structure with an elongated tubular preputium in continuation of the sac. The laterals and marginals both have a peculiar twinned structure. not sharply carinate. The Segmentina-like shells of India and China referred to Hippeutis are apparently not like the type of that genus.Subfamilies. ANATOMICAL CHARACTERISTICS 'The male genitalia generally approximate to Simroth's Typus I. figs. and 21. 9. We can not agree. Conchologia Indica (1876). The twinned nature noted for some of the teeth was not particularly apparent in the radula of Hippeutis complanatus examined. Both agree in . GENITALIA.). and Subgenera side. the Philippines. 10-12) it is too sharply angulated on the j^eriphery. The following are believed to belong here: Helicurbis umbilicalis (Benson) Hclicorbis caenosus (Benson) Helicorbis mearnsii^Sirtsch) Helicorbis papyraceus (Benson) Geological Distribution. 16B). and probably other islands off the cast coast of Asia. figs. fig. The laterals are tricuspid.

Type Trochorbis 1906. Helicorbis. Type Planorbis trochoideus Benson. diaphana. inconspicue radiato-striata. figs. type cited. Germain (1921) omits all reference to it. versus umbilicum inconspicuum. p.ard and Woodward. 6) Until more is known of the genitalia of Planorbis umbilicalis. Brit. convexa. Planorbis umbilicalis) is insufficient for the purpose of making comparisons with Hippeutis complanatus. as does Thiele (1931). Hemisphaerula. subtrochiformi. trochoideu. places a related species in Helicorbis. 1850. penial gland. p. XXIV. penultimo valde convexo. is lenticular with a sharply angular peripheral keel and the aperture is longitudinally triangular. spira parvula. The published description of the anatomy of this group (the type.s (Benson). marginc suj^criori arcuato. 1837. 105. anfractibus 3i/{>. 84). however. Bengal. Kennard and Woodward make it a synonym of Hippeutis (1926). made . the group Helicorbis had better be considered a separate genus related to Hippeutis. which is recognized as a genus. Nautilus. The shell of complanatus. inferiori recidente. leviter excavato. 3 mm. or prostate. as no mention is made of flagella. 90.108 The MoUuscan Family Planorhidae lacking internal barriers or lamellae in the aperture. As subgenus of Planorbis. 16) place Planorbis umbilicalis in Hippeutis. In Benson's paper (1855. 1855 Type by original de. Bartsch. which is like that of Hippeutis (see plate 2. The name Helicorbis has been almost unnoticed since its publication in 1855. p. Considered a synonym of Segmentina. arete convoluta. 352). interne acute carinato. XIX. on the contrary. versus periphaeriam vix declivi. p. has a higher shell with rounded or subangular whorls ancl the aperture is oblique and higher than wide. Dall (1905) overlooked it in his Alaska Mollusks but later (1906) he considered it equal to Hippeutis Agassiz. 'The infrequency and irregularity of the internal laminae in this species. 126) three species are mentioned under Helicorbis. medio. Helicorbis may bear something of the same relationship to Hippeutis that Polypylis bears to Segmentina. The penis (verge) is said to have its sperm canal opening on the side. versus suturam obtuse angulato. hemisphaerula and nitida are ruled out by the remark in the paragraph above concerning Secjmentina (of which nitida is the type) which refers to the internal divisions or barriers. Syn. extus depresso. 5. Genus TROCHORBIS Benson. p. Jour. 126. apertura oblic|ua subsagittata. wider than high. As subgenus of Planorbis. No Description of the type of the genus Planorbis trochoideus Benson: 'Testa vix perforata. Non-Marine Moll. .. nitida. jn'ominente. Asiatic Soc. concava.signation Trochorbis trochoideus Benson 1855. Trochorbis Dall. intus laminis rarissimis jileurumque minuto. There remains the species umbilicalis which has not previously been used as a type for any group and this is here nominated for the type of Helicorbis Benson. p. Bequaert and Clench in a later paper on Philippine Lymnaeidae and Planorbidac (1939. Trochorbis Benson. Diam. in a paper on Philippine mollusks (1907. causing them to look more like accidental thickenings of the shell. sutura profunde impressa. is the type of Pilsbry's Polijpylis. Trochorbis Kenn. the latter treated as a subgenus of Planorbis. axis vel 2' (Benson. although the anatomical differences between the last two groups are greater than appears to be the case of the first two groups. subtus piano. also. p.. 1927. ultimo majori.

India. p. p.. the last whorl very large and strongly overlapping or clasping the preceding whorls. Shell (plate 78. p. Major diameter of shell 7.. Germain (1921. 8-10). the genotype. 126) erected the group Trochorbis for it. Subfamilies. 105) considered Trochorbis a synonym of Secjmentina. 1850. Species Considered as Valid. C. Trochorbis trochoidcus Benson. The Asiatic mollusks allied to Segmentina have shown such diversity in anatomical characteristics that until the animal of trochoideus is examined it seems best to treat Trochorbis as a separate genus. p.^ (Hippeutis) peipinenf^is Mori. Biol. to 8. Conchologia Indica. The is shell treated as dextral (ultradextral).8 mm. 352) Benson noted the trochoid form and small umbilicus of Pkuwrbis trochoidcus and in 1855 (p.. Genera. The embryonic whorls are pitted. 175) does not mention the group name proposed by Benson and places the species trochoidcus in the genus Segmentina. figs. is the only species assigned to this genus. Kennard and Woodward (1926. is near the group of shells called Polypylis. XIV. figs. with the internal lamellae far within the aperture. Ani-fu. Foot and pneumostome lead color. flattened. p.6 mm. Wrongly interpreted as a land shell.^ New Genus Ping and Yen Type Pyramidula (P alula) Pyramidula (Patula) Ill. the base flattened. Ultradextral. This name has l)een overlooked by most writers on Asiatic mollusks. Mem. 90) consider it an absolute synonym of Segmentina. Bull. U one can judge from the very excellent figures of both species in Hanley and Theobald. Aperture without internal lamellae. Ping and Yen. Its place. In Preston's Fauna of British India (1915. with a narrow spire dejiression exhibiting all of the whorls. the species previously referred to that genus now being placed in the genus Polypylis of Pilsbry. According to Benson. 1-3. Barrackpore. lenticular. Animal. B. 1938. Animal. p. 25. Col. 2. body yellowish finely mottled with black. Kidney pigmented (from alcoholic specimens). 352). Bengal. Dall (1906. 6. . Baker. Genus Univ. Trochorbis trochoideus greatly resembles Plartorbis calathus Benson which belongs to the genus Polypylis and from the shell characters appears to be only specifically distinct from calathus. plate 39. figs. The anatomy of this species is at present unknown. Ser. as far as can be judged at present. the apertural lamellae are so weak that he overlooked them in his first description of the species. Fan Memorial In. with sharp i^eripheral carina.. Typical Segmentina is not found in eastern Asia. 1-3 (calathus) and 4-6 (trochoideus) The species trochoideus has a much smaller shell than calathus and the periphery is more acute. Kvoto Imp. and Subgenera — Recent and Fossil 109 me lar overlook this structure in my original description. Sci. Geographical Distribution. Xo. PINGIELLA peipineiisis F. Remarks. the depression about one third of the diameter of the shell. 125) the species appears as Planorbis iSegmentina) trochoideus without reference to its use as type of Trochorbis. plate 14. 288. Trochorbis is not recognized by Annandale and his co-workers in the studies of the mollusk fauna of the Indian Empire. The species is singufrom the nearly total absence of umbilicus' (Benson. p. In 1850 (p. fig. peipinensi.

There is a single. short and wide. 6. plate 5) attached to the vagina by a very short wide duct. 8.5 mm. The sac-like flagella (FL) are turned horizontally to the tube-like duct. 6) there is a short. narrow. Free portion of prostate duct very short. The duct of the flagellum is narrow but is of greater diameter than the vas deferens. The retractor muscle is attached to the preputium near the small liranch of the pilaster . There is a very long and very fine. 7. placed over the stomach. short (little more than 0. Another specimen had a spermatheca long and sac-like (fig. D). The spermatheca is globe-shaped (S) with a short.).. long (3. 3.110 The Molluscau Family Planorhidae ANATOMICAL CHARACTERISTICS PLATE 4 GENITALIA. fig. Vas deferens long (4 mm. The very long duct (DC) which appears on the outside of the preputium. ( The verge ) . 1 elongated. placed on tube-like ducts longer than the whole penial complex. with several but mostly tubular. and DC). There are two flagella. Seminal vesicle (SV) large. 9) . which is in the center of the verge (figs. Tlie other end of the penial duct passes on the outside of the small pilaster and enters the round. V) is long and narrow. narrow duct about as long as the greater diameter of the spermatheca (SD).) vergic sac (VS). C. enters a canal in this small branch (fig. Internally (plate 4. sac-like preputium (PR) and small. tween the vergic space and the preputial space (figs. Vagina rather short but longer than wide (VG). Sperm duct. (fig. Albumen gland fig. vesicles. This spermatheca contained a spermatophore and the specimen was evidently in gravid condition. Nidamental gland (NG) veiy long (3 mm. with a small penial papilla at the end. Oviduct (OD) short and narrow. fig. There is a thickened muscular ring or diaphragm be2. Female Organs. Penial complex (fig. GL). flat gland (GL) which is doubled backward at the end (fig. 2. The sac-like portion of the flagella is large and the cavity is lined with elliptical cells which show on the exterior surface as small. 5. 7. Male Organs tortuous. PL). muscle (R]\I) attached about midway of the i:)reputium at the point where the penial duct has its exit. wide retractor ( DC ) . about twice as wide as long. The form of the spermatheca evidently varies with the breeding condition of the animal. 8. 6. more than double the diameter of the ovisperm duct. long with twenty to twenty-two long diverticula. RM). G (fig. one branch forming a wide. 5. Uterus (U) short and as narrow as the vagina. 6. 4).) and much swollen. The lower end of the verge is notably flattened and broad. (plate 4. wide pilaster which bifurcates about midway of the preputial cavity. swollen or pear-shaped at the upper part but narrowed and tubular below. the intestine looped beneath it. Prostate (PRS) about 2 mm. 6). disc-like organ (G). just beside the sperm canal exit. C) which terminates on the underside of the reflexed end of the gland (fig. rounded projections (fig.) and tubular. 5) with a large. narrow. The vergic sac is placed on the side of the upper part of the preputium. fig.5 mm. turban-like termination of the small pilaster (figs. The other branch is short and narrow and ends above in a rounded. tube-like external penial duct about three times as long as the penial complex when stretched to its fullest extent.

sac-like glands. Hermaphrodite Organs. The terminal end of the verge is also peculiar. the radula sac large and near the middle of the posterior end. The ureter is directed horizontally into the mantle cavity. the ducts are small The jaw and tube-like. The salivary glands extend considerably beyond the rear end of the buccal sac and are united behind. The portion between the seminal vesicle and the ovotestis is very short (less than 1 mm. It is horseshoe-shaped as in the other groups of this subfamilv. the latter with a turban-shaped termination. Northern and eastern China and Japan. received from the Fan Memorial Institute of Peiping. These differences are sufficient to stamp the species as belonging to a new and hitherto unknown generic group. is the intestine Digestive System. presumably of this species. It differs from Segmentina in the possession of an external duct and from Polypylis by having a penial gland which is absent in that genus. Species Considered as Valid. Unknown. Pingiella peipinensis (Ping and Yen). but not as long. The anatomical details have been obtained from three specimens from Peiping. is the only representative of this monotypic genus. is Geographical Distribution. resembling that of Hippenfis. Tiie kidney is long and narrow. figs. fig. The Respiratory and Renal Systems. and the very long. Chi Ping. is fragmented with about eighty plates. The buccal sac is elongated. the penial gland. resembling that of Segmentina. contained eight eggs. the genotype. Radula formula 26-1-26 with 160-170 rows of teeth. China. not reflected. 1. There is no superposed ridge. making a loop around the stomach and another around the liver and then passing forward to the rectum and pseudobranch. and Subgenera ovotestis — Recent and Fossil consists of 111 many long and ovisperni duct (SO) is of small diameter. P) small and inconsjncuous. 7. Remarks. The laterals have six short. The teeth resemble those of Segmentina and Polypijlis (see j^late 69.) but the portion between the seminal vesicle and the oviduct is very long (about 3 mm. pear-shaped. An egg capsule on one of the shells.Subfatnilies. tube-like external duct with its peculiar gland terminations. This genus is remarkable for the peculiar characteristics of the penial comj^lex. the veins at the upper part of the triangle. The blind sac is very short and pointed below. It probable that this distribution will be extended when more material has been examined from China and India. bifurcated into a large and a small branch. A section below the middle shows a flatter form with the lumen almost as high as wide and somewhat triangular. The stomach is long and narrow as in Segmentina. Genera. 8). Geological Distribution. The pseudobranch (plate 5. through the courtesy of Dr. The (OT) narrow diverticula placetl in a double series. This triangular form is caused by the acute angle of the shell on the periphery. A section of the kidney near the middle shows a wide and low lumen about three times as wide as high with the veins at the upper corners. spade-shaped cusps and the marginals have seven short cusps. . The flagella are notable in having the long ducts and the small.).

As sub- genus the body-whorl. places the species in the genus Anisus. The anatomy. with a few irregular.^le. The genus is dedicated to Dr. Sperm duct (SPD) long (slightly more than 1 mm. research professor in the Fan Memorial Institute of Biology. an accomplished student of the Asiatic molluscan fauna.112 The Molluscan Family Planorhidae It is quite probable that other species belong in this genus and will be discovered when examined anatomically. The shell is very minute and has few whorls. 2. 3). narrow (fig. in length.) and of large diameter. No figure of the shell is available and the single specimen used for dissection had the shell badly broken which made it unfit for illustration. 361. when stretched out). about three times the diameter of the ovisperm duct. 1922 p. Peiping. GENITALIA. which increase in size rapidly. glandular projections. Thiele. Mus.^nd. Chi Ping. especially the genitalia. XXIV. with six large. Ind. 361). There is a short. club-shaped diverticula. ( Genus Type As genus. however. b}^ original designation Intha capitis Annandale 1922. Anisus (I. The lower surface is narrowly umbilicate. Animal. Prostate duct (free portion) about as long as the prostate. though relatively smaller than it completely embraces. The pseudobranch is poorly developed' (Annandale. The head of the animal bears short. The aperture is large and very oblique but with a cordate outline. Together both organs measure 1 mm. It is less than 1 mm. The external surface is practically smooth' (Annandale. bulbous preputium (PR) and a small. 362). Mori 1938. slender vergic sac (VS). AXATOMICAL CHARACTERISTICS PLATE 8 11). following the classification of Thiele. pointed. 1931. Rec. The shell is ultradextral. but internal ridges are completely absent. Dr. Type hitha capitis Annandale. p. long. Those of the spire are cylindrical.. p. differing widely in its internal organization. 482.and occludes the rest of the shell in such a way that the spire is entirely concealed. subgenus Hippeiitis. There is a well-developed simple callus on the inner lip. indicates that this species can not correctly be referred to either Anisus or Hippeutis. Male Organs . blunt tentacles (plate 8. The outer lip arises in the middle of the upper surface and forms a small lobe at its point of origin. Shell. fig. except in so far as it can be detected by transparency. this is Tn in Hippeutis. Intha IXTHA Annandale. 6).) capitis (Annandale). Seminal vesicle (SV) very large. He. but the body whorl is flattened below and has the form of a flattened conoid slightly truncate above. Prostate short. Intha Ann. The penial complex has a large. p. 288) has called attention to the fact that this species is a planorbid and not a land shell. somewhat shorter than the preputium (fig. China. p. Vas deferens (VD) of small diameter and very long (about 5 mm. Ping has supplied the author with a number of mollusks for anatomical study from near Peiping. 'The animal is remarkable externally for the large upper and lower lobes into which the mantle is divided. Handbuch. Teil genus under genus Anisus.

There is a short duct. The buccal sac resembles that of Segmentina. 27) the lumen is long. The ovotestis (OT) is large with the diverticula long and clul)-shaped. the cup being free. tapering. fig. 8). FL). 11. the intestine making the same loops around the stomach and liver as in the genus Segmentina. external to the 'preputial cavity. The relationship of the gland and duct to the preputial sac is well shown in fig. Lateral teeth (1-5) with two flatly . There arc more than sixty plates on the jaw. Annandale says of the genitalia: 'The male organ resembles that of Planorbis. The radula has the formula 15-1-15 with over 125 rows on a liroken membrane (iilate 69. and Female Organs circular. The verge (V) is narrow. fig. The kidney (plate 47. depressed. about 1 mm. which enters a round. FL). In section (fig. sac-like. 1) Hermaphrodite Organs. The spermatheca (S) is large and almost and is joined to the short. 23) is fragmented as in Segmentina. 8. except that the penial bulb is relativelv verv large' (op. The ovisperm duct (OS) is very short between the seminal vesicle and the ovotestis and about 1 mm. The jaw (plate 50. The pseudobianch (fig. There is a large carrefour (CF). The stomach is narrow as in Segmentina. Small glands on the interior of the fiagella show through as white circles. 7.. long between the seminal vesicle and the oviduct. Digestive System. A spermatophore of peculiar form was found in the spermatheca No figure or descri]ition has been published of the genitalia of this genus. Genera. P) is a rounded. about as long as the preputium. Internally (fig. fleshy ring or diajihragm between the vergic and jn-eputial cavities (fig. the preputium has a large penial gland (GL) which is attached to the preputium only at its base. simple lobe. Respiratory and Renal Systems. 2. The albumen gland (AL) is large. as shown in the figure (fig. D). 26) is wide and fiattened and not as long as in Segmentinn or Poh/pi/Us. fig. long. pierced above by the rectum. 11. 8. with a wide cup-shaped depression in which the walls are ridged (probably glandular). the cusps not reaching the lower margin of the base of attachment. There is no superposed ridge. long) which are sausage-shaped and are attached to the vergic sac by a very short duct. 362). and about twice as long as the uterus. 4). with the exit at the end in the center. The oviduct (OD) is short and wide. VS. At the upper end of the vergic sac there are two very large fiagella (about 1 mm. Center tooth bicuspid. wide vagina (V) by a duct which is shorter than the diameter of the spermatheca. Subfamilies. p. The nidamental gland (NG) is very wide. and Subgenera — Recent and Fossil 113 penial gland duct on the outside of the i)reputiuni. The uterus (U) is wide and shorter than the spermatheca and its duct. 11). 9). emerging from the latter about midway between the outlet and the summit of the preputium. cit. There is a wide retractor muscle which is attached to the upper part of the preputium. in fact they are almost sessile. There is a very small papilla at one side of the opening (fig. It is more or less circular in shape. They are placed in a double series. The two fiagella are joined in one tube before thev enter the vergic sac (fig. has a space for the passage of the intestine (fig. oval in outline with the veins about in the center on each side.. (fig. fig.

11. a single raesocone. Known at present onlv from the tvpe He-Ho and Inle valleys (3000-3800 ft. The teeth are all in a straight row. and F. op. M. 397. Drepanotrema Pilsbry.\ll. and F.. 1934. Planorbis of 1899. and F. and the striking penial complex with a penial gland which has a small. As section of Planorbis. S... It appears to be another peculiar variant of the Segnientina group of Planorbidae. Figures genitalia of D. Spiralina 1899. Ill. the mesocone a trifle larger than the other cusps. subgenus of Planorbis. Miss. Species Considered as Valid. p. cited as example.. As genus.. They are very uniform in size and shape but become narrower toward the outer edge of the membrane. Type Planorbis yzabalensis C. 1930. Type Drepanotrema yzabalensis C. As subgenus of Planorbis. Intha capitis Annandale is the only species recognized for this monotypic genus. Acad. Drepanotrema M. Type Planorbis yzabalensis C. and F. Rec. S. 75. (= Pkinorbis anatinum d'Orbigny) 1880. No. S... p. II. and S. p.. As synonym 1899. Conch. p. Moll..). Type Planorbis yzabalensis C. 481. Zool. 397. In this respect it resembles some of the Helisomatinae (typical Helisoma) but differs in the form of the prostate and ovotestis. p. p. (part) Cent. Handbuch. Cites Plaiiorbis yzabal- 1905. Man. Type Planorbis yzabalensis C. 8. Marginal teeth 7-cuspid (6. 82. Type Planorbis yzabalensis C. 86.) and D. southern Shan States. 1820). 106. 1884. Bathijomphalus Tryon (non Agassiz. Ill. and three ectocone cusps. 1837).114 The MoUuscan Family Planorhidae entocone cusps. Sci. and F. Annandale in Inle Lake. Univ. H. cultratum (Orb. Amer. Drepanotrema Tryox. As subgenus. 58. Drepanotrema D. Oc. Mich. de Conch. 1883. There are no intermediate teeth. pp. Alaska Moll. Drepanotrema Germain. and F.. Mus. . and S. 8. Moll. Drepanotrema Fischer. and F. B. Includes Drepanotrema cultratum cit. cit. 1884. 509. Papers. Scient. Teil 2. Type Planorbis yzabalensis C. 12) with three small cusps on each side of a mesocone cusp. Mus. Genus DREPANOTREMA Fischer and Crosse. Hippeutis ensis. Martens (non Agassiz. lucidum (Pfr. kertnatoides Orb. received from Dr.artexs. As genus. 43. Ind. Remarks. p. 106. Hippe litis. Burma. Rao. Mex. and sumichrasti.. Ayiisus Tryon (non Studer. Section of Gyraulus. Conch.. 1880 Monogenotype Planorbis yzabolensis C. p. Drepanotrema Fischer and Crosse. with the few prostate diverticula of large size.. Type Planorbis yzabalensis C. pp. Recent and fossil. 59. the two enormous flagella. 1884. Drepanotrema H. 1921... N. Intha is peculiar in the form of its shell and also in the characteristics of its genitalia. 38. 1931. and F. Biol. type of Drepanotrema. 395. p. S. XXI.. For the above anatomical details a single specimen was dissected. 107. Conch. collected by Dr. The teeth are essentially like those of Segmentina. 51. As subgenus of Planorbis. Proc. Planorbis anatinus Orb. Nat. op. cited as example. Burma. locality. 1837). alt.^rtexs. and F. and S. p. Phil. Geographical Distribution.) southern Shan States. As section of subgenus Hippeutis.. Drepanotrema Thiele. 210. p. short duct entering the circular ring between the vergic sac and the preputium. Ill. Baker.

placed at the inner base of the tentacles. Small. there are seven diverticula still more widely sjiaced and the prostate is about 1 mm. . The penial comi~)lex is much elongated in all species examined. but in some others (as hoffmani F. and lucidum) Seminal vesicle (SV) large. 1). the vergic sac is shorter than the preputium. fig. long.6 mm. 16-18). narrow tube. the flagellum is very long (1 mm. figs. from the sperm duct and are not on a separate duct as other 2. rounded before and sharply pointed behind. 7). plate 10. The body is ANATOMICAL CHARACTERISTICS PLATES 9. 7. figs. in lucidum. varying in length in the three species examined. swollen. and Subgenera — Recent and Fossil 115 Shell (plate 79. It is about half the length of the ovisperm duct region in hoffmani but only about a third of this lengi:li in lucidum. Anatinum is much like hoffmani. 2) the body is not much pigmented. 1. The embryonic shell is punctate. the body whorl large and expanded. the flagcllum is The . and about twice the diameter of the long and narrow vergic sac. lucidum and a little over 1 mm. the flagellum is as long as the preputium (0. (plate 9. 4). figt. The vas deferens of the three species is always a long. fig. long. varying in in mm. hoffmani. whorls rounded or carinated. The prostate (PRS) varies greatly in the three species: in hoffmani it is 3. In hoffmani (plate very short and is bifid at the end. the preputium is about three times the diameter of the vergic sac. cylindrical. of few whorls. figs. long and is made up of twenty-three short. 1) the l)ody is dcei^ly jngmented with black. A strong retractor muscle is attached to the end of the vergic sac near the insertion of the flagellum (plate 10. which is nearly twice the length of the preputium. fig. fig. There may be two dark lines of color extending down the neck and top of the head. 5) the preputium is elongated. 10. Genera. plate 11. 6. The sperm duct (SPD) also varies in length. which forms a reversed cone. length in the different species. The tentacles are long and filiform. flagella of the three species differ considerably.3 in Segmentina and in hoffmani. In some species (as melleus Lutz. there are but eight diverticula on a prostate about 1 nun. iiltradextral.. 1). C.5 mm. two to three times the diameter of the ovisperm duct. 51.Subfa/fnlies. which is considerably longer than the preputium (plate 11. Baker. the preputium and the vergic sac were about equal in length (plate 11. the right side with whorls sunken below the upper surface or showing a wide. in lucidum (plate 11. in anatinum (plate 9.) and is bifid at the end. 10. There are no barriers or lamellae within the aperture. All diverticula arise directly GENITALIA. widely spaced diverticula. long and narrow. In lucidum. members of the subfamily. 5). Animal. respectively anatinum. club-shaped. or more) and is not bifid at the end. These lengths do not include the sperm duct portion under the prostate diverticula. 2. 8). In anatinum (plate 9. much elongated. embracing the previous whorl. There is also a large nerve at this point (N). fig. being 5 mm. In another specimen. slightly concave depression. in anatinum. becoming smaller in diameter toward the ui^i^er part. plate 10. fig. plate 78. fig. Male Organs fig. fig. 4. fig. the eyes large and conspicuous. in anatinum. 3. 11 2. In hoffmani (plate 10.

2 the spermatheca is pyriform and is joined to the long and narrow vagina by a duct twice as long as the spermatheca. plate 11 {lucidum). the vergic sac being but a trifle shorter than the preputium. in anatinum (plate 9. Pilsbry 's figures are the only ones previously published on the anatomy of Drepanotrema. Several specimens of lucidum had the male genitalia completely everted and lying on the body of the animal (see plate 11). Pilsbry (1934. The nidamental gland is long and wider than the uterus in all three species. The albumen gland is much elongated (plate 10. 59. as shown in fig. 6. The ovisperm duct is much longer in lucidum than in either hoijmani or anatinum (see plate 9. Baker). This is shown to better advantage in fig.) (plate 76. a small muscular ring or diaphragm between the preputium and the vergic sac. fig. fig. 2) the spermatheca is wider than in hoffmani but not as globular as in lucidum and is attached to the narrow vagina by a duct almost three times as long as the spermatheca. the penial complex is simple in all species studied and resembles the condition observed in hoff)nani (plate 10. 2. Pilsbry also figured the penial complex of Drepanotrema cultratum (Orb. The flagellum is somewhat more than half as long as the vergic sac. 6). 3). much wider than in hoffmani. These also differ in the three species. 4). much shorter in lucidum (about 2 mm. There are three rather heavy pilasters.116 The MoUuscan Family Planorhidae Internally. the preputium is fully everted and the verge is extended to its fullest extent. only the preputium is everted. 8 on plate 10 [hoffmani) the vas deferens. As shown in fig. fig. vas deferens. and is connected with the vagina (which is shorter than in hoffmani) by a duct a trifle longer than the spermatheca. The uterus (U) is very long and narrow in hoffmani. fig. 2. the organs are diagrammatically shown so that the relationship of the different parts may be seen under this condition. Female Organs. and a very long verge completely filling the vergic sac (V). The flagellum. fig. In hofjmani (plate 10. 4. fig. F) figured the penial complex of lucidum (from sketches by H. and retractor muscle remain in the body of the animal just behind the male ojiening. fig. and narrow in anatinum. shorter and wider in lucidimi. 7. 6. 1 plate 10.). the retractor muscle was inserted at the end of the vergic sac near the flagellum and not at the summit of the preputium. 2. in lucidum the spermatheca (plate 11. In fig. plate 11. 9). In all specimens of Drepanotrema examined. . B. 1. flagellum. fig. fig.) and very short in anatinum (less than 1 mm. . p. There is a small and inconspicuous carrefour (CF) in all species. 3) is globular. The verge is narrow at the end with a central sperm outlet (plate 10. 10) which is similar to that of lucidum. This figure differs from the specimens here examined in having a shorter flagellum and in having the penial retractor attached to the preputium instead of to the vergic sac. fig. The oviduct (OD) is very long and narrow in hoffmani (about 5 mm. In fig. and retractor muscle may be so closely pressed together and bent downward at the summit of the preputium that relationship of the muscle is difficult to see clearly unless the parts are separated. vergic sac. The ovotestis (OT) consists of a few clubshaped diverticula placed in a double series. In fig. Hermaphrodite Organs. All specimens were torn apart to insure accuracy in this statement. long). fig.

and five small cusps on the outer margin of the tooth where the ectocone would normallv be. fig.' . the pyloris tapering to the intestine. which forms a loop over the oesophagus and another loop around the liver as in other genera of the subfamily. figs. Lateral teeth (1-6) very wide. fig. with 194 transverse rows. the lumen wider than high. Above the entocone there is a small additional cusp. 1. The center tooth is as in lucidum and anatinum. Tlie pseu(l(jl)rancli (P) is a somewhat i)ro('ess in lucidum (plate 11. the formula is 27-1-27 with 230 rows of teeth. (plate 68. fig. The stomach (plate 48. and above the ectocone there is another small cusp on the first four laterals and two cusps on the 5th and 6th laterals. Baker (1930. The lateral and marginal teeth are similar to those of the other species figured. the lumen regularly long-ovate and the veins smaller and more 'regular in form (fig. and ectocone. 11) is pyriform. lucidum. A cross section near the middle (fig. B. The intermediate teeth (7-10) are very wide with the three main cusps well-developed and with a small cusp above the entocone and a small interstitial cusp between the mesocone and the ectocone. Above the ectocone there are four small cusps on the outer margin of the teeth. The center tooth are longer. with three long cusps. 15) shows the kidney to be much flattened. 3). 4). About seventy plates were counted on a jaw of lucidum (plate 50. 14) is very long and ribbon-like (15 mm. H. There is a very small cusp on each side of the center cusps high up on the reflection. The kidney of lucidum is shorter and a section near the middle shows it to be more flattened than hoffmani. Dr. The ureter is very short and curves upward at right angles to the long axis of the kidney. Genera. In hoffmani (plate 68. fig. the formula is 28-1-28 with 240 rows of triangular teeth. The pneumostome is very large (PS). 2) and anatinuni (plate 9. The buccal sac is similar to that organ in Tropicorbis. 3) is 18-1-18 with 158 rows of teeth. The blind sac is two-thirds as long as the stomach. p. long). 16). In lucidum (plate 68. The kidney of Drepanotrema hoffmani (plate 46. 27) and about as many on a jaw of hoffmani (]ilate 50. ahenum are also much as in D. 2).Subfamilies. Center tooth squarish with two long. fig. The rectum is very large and the anus discharges just above the base of the pscudobranch. The jaw is horseshoe-shaped and is composed of many small plates arranged somewhat as in Planorbis. 3) but in ho)J))iani appears to be narrower and more cylindrical (plate 10. and the veins flattenecl and elongated in section. fig. 50) has examined the radula of ahcnum and says 'The radula and jaw of D. The marginal teeth (11-18) are narrower than the intermediate teeth. 26). fig. and Subgenera — Recent and Fossil 117 Respiratory and Renal Systems. The radula formula is 30-1-30. mesocone. three cusps between the entocone and mesocone. The marginal teeth are slightly broader at the tip of the cusped reflection and may develop two interstitials between the entocone and mesocone. The lateral is like that of anatinum except that the large cusps and marginal teeth are similar to those of anatinum except that the cusps are longer. spade-shaped cusps reaching two-thirds of the way to the lower margin of the base of attachment. there is a long entocone and mesocone with a small cusp above the entocone. fig. entocone. fig. The radula formula of anatinum. Digestive System.

the periphery from rounded to carinate. varying principally in the number of interstitial and accessory small cusps. (Moricand) labrosum Drepanotrema (Fossulorbis) kermatoide (Orbigny) c. G.oricand) Drepanotrema (Fossulorbis) (Cooper) anitense Drepanotrema (Fossulorbis) (Tristram) Drepanotrema (Fossulorbis) Pilsbry Drepanotrema (Fossulorbis) Pilsbry c. periphery broadly rounded (type Drepanotrema anatinum (Orh. and the Argentine Republic. panuco Drepanotrema (Fossulorbis) sumichrasti (Crosse and Fischer) Remarks. narrow and deeply sunken. vary in the different species. Baker Drepanotrema (Fossulorbis) c. Isabela. including some names now considered synonyms. and F. the last section should be regarded as a separate genus until the type has been dissected and its anatomy made known. the diverticula .. collected by Dr. Drepanotrema (Fossidorbis) cultratum (Orbigny) Plate 78. p. (Pilsbry) Species Considered as Valid. Received from Dr. Baker castaneonitens (Pils. duenasi- Drepanotrema (Fossulorbis) H. Havana. Pilsbry (1934. W. Hoffman. Aguayo. Intha. very slightly concave. and Platytaphius Pilsbry. figs. Fossidorbis Pilsbry. For the above anatomical data.) Pilsbry. Five specimens dissected.) Pueblo Viejo. 1934 wide spiral on the right side. 17-19 c. as well as the number of lateral and marginal teeth. Puerto Rico. Puerto Rico. C. 1880 Spire on the right side. Drepanotrema hoffmani F. 59) divided the genus into three sections. By its prostate. Drepanotrema hoffniani F. It is jnirely an American genus and a number of species are known. The formulae. 14-16 Drepanotrema Drepanotrema Drepanotrema Drepanotrema and Van. Bolivia. This genus is found more or less abunfrom southern Texas southward through Mexico and Central America to Brazil. A. C. pistiae depressissi- anum mum c. Drepanotrema lucidmn (Pfeiffer). Drepanotrema (Fossulorbis) cimex(M. W. Drepanotrema proper. aruitiniun (Orbigny) 16-18 Drepanotrema lucidmn (Pfeiffer) Plate 78. melleum (Lutz) Subgenus Having a FOSSULORBIS Pilsbry. etc. collected by Dr. Hippeutis. Species Considered as Valid. Drepanotrema differs in its genitalia and radula from all other American groups of Planorbidae. 11-13 Drepanotrema Plate 79.118 The MoUuscan Family Planorhidae The radula teeth of Drepanotrema are peculiar in the possession of the small interstitial cusps between the larger cusps.) Drepanotrema chittyi Aguayo parapseide (Orbigny) ahenum H. In the writer's dantly opinion. B. three species have been examined: Drepanotrema anatinum (Orb. Hoffman. C. The other groups divide the genus Drepanotrema into two comprehensive subgenera. Five specimens dissected. figs. A. Cuba. figs. They somewhat resemble the multicuspid teeth of Segmentina. Subgenus DREPANOTREIVIA Fischer et Crosse. Baker Plate 78. It it also found in many of the West Indies Islands. Eight specimens dissected. Type Drepanotrema cultratum ((Drb. Baker. B. Geographical Distribution. )= Planorbis yzabalensis C. The radula of Drepanotrema is very uniform in the shape of the teeth. to which the genus is also allied by the presence of a fiagellum. figs.

and Hippeutis. Proc.51 mm. 86. and . Paraplanorbis condoni (Hanna). In view of the difficulty in this family of placing a group in its correct position in the classification by the shell alone. Genus Type by PARAPLANORBIS Hanna. is the only species ascribed to this originally monotypic genus. 212. in diameter. Lines of growth fine and even. 12. Diameter 2. composed of about four and one-half regularly increasing whorls. Oregon Pub.35 another reason for such procedure. all having the peculiar embracing whorls so characteristic of Segmentina. Paraplanorbis Pilsbry. Amer. Sci. mm. is altitude 0. Paraplanorbis H. and Subgenera — Recent and Fossil 119 club-^licii)ed aiitl placed in a ^^iiigle row on the sperm duct.'Vnna. p. Aperture roughly triangular but rounded on the outer lip. Species Considered as Valid. . also. condoni Hanna. as is usually the case in Drepanotrema and its allies. Pliocene period. umbilicus very narrow. Vicinity of Warner Lake. not expanded or thickened but a dejiression both above and below the periphery. 245. p. eastern Oregon. description of subgenus). Phil.. As section of Planorbis.. Non-Marine Moll. Although dei)arting from these groups in having the diverticula of the prostate placed directly on the sperm duct. As section or subgenus of Drepanotrema.67 nnn. 4. The presence of the fliagellum on the vergic sac places Drepanotrema near the Euroixnm and Asiatic groups Segmentina. it would seem that the interest of nomenclature is best served by considering Paraplanorbis a separate genus of Segmentininae related to Drepanotrema. 4. Univ.. No. 1922 Hanna Type Planorbis Type Planorbis Fossil. This fossil genus is related to Drepanotrema and is considered a section or subgenus of the genus by Pilsbry (1934. Shell (plate 81. The jirostate differs from the members of both Planorbinae and Segmentininae in having the diverticula placed directly on the sperm duct instead of on a separate prostate duct. (Par a planorbis) condoni Hanna. I. fig. genotype. 59. 1935. The form of the shell of typical Drepanotrema bears close resemblance to the edentate Oriental groups of Segmentininae. it is allied to the subfamily Segmentininae. the spire (left side) appears entirely different. original designation Planorbis condoni 1922. the other characteristics place the genus in the subfamily Segmentininae with little doubt. This feature alone marks the assemblage of genera as of subgeneric solidarity. Intha. planorboid. Hippeutis. apical cavity a cone of about 120 degrees. p. p. 1934. Its geographical location. Hanna's locality No.. While the wiiorls are of a clasping or embracing nature. size minute (Hanna. 5) Whorls rounded on the periphery as in Helisoma Swainson but aperture not expanded. 59). giving the shell a silky luster. Nat. Umbilicus deeply rimate 0. 1922. Remarks. so far removed from the present day distribution of Drepanotrema. the depression being narrow and the whorls deeply immersed. Type Planorbis condoni Hanna. N. Polypylis. Acad. The nature of the rachda also indicates relationship with these groups. Paraplanorbis Hexderson. Last whorl slightly angulated about the upper margin and descending slightly at the aperture. such as Helicorbis.' Geographical and Geological Distribution.Subjainilies. and Intha. Hanna's description of the species Planorbis condoni is as follows: 'Shell ultradextral. p. Genera.

p. Acad. and Van. and strongly angulate or keeled in the middle. Nat. and in fact. periphery conspicuously carinated on the left side. Pilsbry. Planorbis heteropleurus Proc. Phil.120 The Molluscan Family Planorbidae Genus Type by PLATYTAPHIUS Pilsbry and Pilsbry. that it would seem impossible for species of any group without intermediate habitats to become established so far apart. Platytaphius heteropleurus (Pilsbry and A'anatta I is the only recognized species. Proc. As section of Planorbis. p. Platytaphius Pils. titicacensis Clessin. Pliocene to Recent fauna. and Van. andecola. Pal. Platytaphius Pilsbry. which is shallowly vortex-shaped. Acad. rounded-pentagonal. 562). this Remarks. Sci. N. 268) have associated two extinct molluscan species from the Pliocene of Oregon with Platytaphius. Acad. 562. 1934. moderately corneous-white. Nat. 86. Cites two Shell (plate 79. it was made a section of the genus Drepanotrema. 10-12). 1896. the right margin produced forward (Pilsbry and Vanatta. the surface with fine. and Van. As section of Planorbis. Pilsbry says: 'This remarkable species is totally unlike P. This locality is so removed from Lake Titicaca. Sci. from Oregon. It is most like P. p. 1924 original designation Planorbis heteropleurus Pils. Nat. Animal. close growth lines. belong near Drepanotrema or it may belong elsewhere. 76. the whorls nearly flat. As section of Drcparwtrema. 1896. p. p. . 1934. Henderson and Rodeck 1934. all visible on both sides. in Peru (more than 5000 miles). montanus d'Orbigny. 51 this species was made the type of a section of Planorbis and in 1934 (p. In 1924 (p. Platytaphius fossil species Henderson. 1-3. Jour. 1896.. P. In a paper by Henderson and Rodeck (1934. Of medium size. Phil. and P. 51. on the right side. and Van. Whorls three and one-half. Peru. convex. Sci.' With this statement the present author is in complete agreement for the general form of the shell is so distinct from that of any other species of Drepanotrema that it does stand quite alone. so diverse in characters that no profitable comparison can be made' (1896. Adfar . p. At present unknown anatomically. W^hat its real affinities are can not be known until its anatomy has been investigated. Amer. figs. Y. VIII.4tt. Phil.. heteropleurus P. Lake Titicaca. heteropleurus Pils. Platytaphius Henderson and Rodeck. Teil 2. 1935. p. Type Pla7wrbis Thiele. The It ( may localities are sixteen miles southwest of Vale. Geographical Distribution. ultradextral. solid.. species already known from this Andean lake. Proc. p. andecola d'Orbigny. In describing peculiar mollusk. p. Geological Distribution. Aperture very oblique. earlier whorls rather deeply and about equally sunken on the two sides. ) . 254. 59. rather opaque. Type Planorbis heteropleurus Type Planorbis Type Planorbis Fossil. 1924..\.. 562). the last wider than the spire. 268. Handbuch. and V. Species Considered as Valid. heteropleurus Pils. There are fine punctations on the embryonic whorls. Last whorl slightly wider on the right than on the left side. but much flatter with differently lilaced keels. 59). figs. As genus. Yet the two species from Oregon correspond fairly well with the diagnosis of the genus Platytaphius. Non-Marine Moll. plate 26. 479. 268) the authors state that 'it is so distinct that it might be better to consider it a genus. Platytaphius 1931. p.

c and 9. narrow vagina by a very short duct about one-third the length of the diameter of the spermatheca. openly umbilicatc. 1937. recent or fossil. apex deep brown. smooth. 1937 Odhner 14. of circular form. figs. Height of aperture same as height of whorl. (fig. Band 29B. Acrorbis Pilsbry. figs. and Subgenera — Recent and Fossil 121 ditional si)ecies. (fig. There is a wide retractor muscle (rp) which appears to be attached about midway of the length of the vergic sac. The penial complex (fig. shell brown. a. c. The tentacles are narrow. tube-like verge and a diaphragm of fleshy ridge separating the preputium from the vergic sac. merging into the vergic sac without marked constriction between these sacs. Shell (plate 72. Acrorbis ACRORBIS fiir Odhner. Gray. Genus Type by 1937. Internally there is a long. globular. Male Organs 10). (translated from Odhner). 7nilleri Henderson and Rodeck. may be found somewhere between these extreme points. but swells to several times this diameter posteriorly where it joins the nidamental gland. The head is sharply pigmented between the eyes. The foot is o\'al with thin borders and without any furrows on the sides. 1-3). Neither description nor figure indicates the presence of a separate prostate duct. b. 9. dark and reddish above. The prostate is almost as long as the uterus and oviduct and has eight large diverticula in a single series. At present there seems no alternative but to include the Oregon fossils in Platytaphiiis. Above the vergic sac there is a wide flagellum less than half as long as the vergic sac. very thin because of the lack of lime in the mouth-building. b. Archiv. Ultradextral. Type Acrorbis petricola Odhner. a. 9) is elongated.. LI. original designation Acrorbis petricola Odhner. simple. 6). ]). but firm. figs. The vas deferens (v) is a long and narrow tube. No seminal vesicle is show^n in the figure or mentioned in the description. Zoologi. The eyes are on the inner side of the base of the tentacles. 1938. Genera. Animal broadly ANATOMICAL CHARACTERISTICS PLATE 72 GENITALIA. one on each side of a central tube (fig.5 mm. on the end obliciuely truncated. As genus. the upper surface of the mantle flecked with black.Subjmnilies. The uterus (u) is at first as narrow as the vagina. shell covered with a shining cuticle. 9. arched. p. about one-third of the entire length. The upper end of the flagellum is said to liave two cavities. The oviduct is not shown clearly but evidently narrows to the point of junction with the ovisperm duct. broadly ovate. Female Organs is and . figured on j:)late 37 of the publication cited. The head is darker gray and neck blackish. 5. lips simple. No. thin. Nautilus. since no diagnosis could be made that would include the one and exclude the other. Height 1. very small. underside more grayish. 6. when fully extended. whorls slowly increasing. 1. Nov. The left tentacle differs from the right tentacle in being bifid at the end. the preputium. The albumen gland is not figured. a). high. (plate 72. aperture very oblique. with elevated spire. diameter 2 mm. The spermatheca (b) is large and globular attached to the short. 10). 107 (review). the following whorls with very weak spiral lines. The two species of fossils from Oregon are Platytaphius vialheurcnHiH and P.

It lives rocks over which water trickles. these two genera belonging to another subfamily. which are smaller than the diverticula of the prostate. State of Santa Catharina. Acroi^bis is apparently a very distinct group of South American Alollusca. k) is shown as bifid is a narrow projection about twice as long as wide. 8). petricola Odhner. The kidney (fig. and a short entocone and ectocone about half as long as the mesocone. 6. The information concerning this genus has been drawn entirely from the original paper and figures bv Oclhner. Digestive System (fig. type of the only species recognized.. It is also compared with Neoplanorbis Pilsbry. The intermediate teeth have the cusps more nearly equal in size. the Helisomatinae. 5. consisting of but five diverticula. but that group is referable to the family Ancylidae. quite different from the . the globular spermatheca on a very short duct. c). genus. The jaw is said to be like that of Parapholyx iPompholycodea) The radula (fig. The buccal sac (fig. and a long intestine which makes a loop around the stomach and another around the liver. The ovotcstis (go) is few-lobcd. Species Considered as Valid. It is not related to Carinijex or Parapholyx {Pompholycodea) as indicated by Odhner. 107) that it is related to either Tropicorbis or Drepanotrema. His figures are reproduced on at the extremity. The center tooth has two wide cusps which do not reach the lower margin of the base of attachment. It plate 72. There is a long and narrow oesophagus. 7) is elongated. not Planorbidae. and probably bears the anal outlet as in other Planorbidae. in which there is a penial gland in the male comjilex. wide for the whole length with no differentiation between preputium and vergic sac. Acrorbis differs from Drepanotrema in the wide form of the fiagellum. in damp moss on is tlie Geographical Distribution. an ovate stomach with a short pyloris and a very short blind sac (caecum. Respiratory System. 122 The Molluscau Family Pknwrbidae Hermaphrodite Organs. entering the radula sac by a fine tube. The ovisperm duct (od) is almost as long as the uterus and oviduct combined and is a very narrow tube. and the helicoid form of the shell. wide at the posterior end and much elongated at the anterior end. extending forward in the rectum (r) to the outlet near the pseudobranch. The salivary glands (s) are short. 5. p. It scarcely appears to belong with Choanomphalus. There is an unusually long radula sac (t) which forms a narrow tube as long as the whole radula sac. The pseudobranch (figs. spade-shaped mesocone reaching the lower margin of the base of attachment. Brazil. the entocone and ectocone splitting into two smaller cusps. It is not stated whether the lobes of the ovotestis are in a single or a double row. 4) formula is not given but sixteen teeth show in one row in the figure which would make the formula 16-1-16. The marginals appear wide and low. Drepanotrema and the few diverticula of the prostate in a single series indicates the same relationship. The laterals are sciuarish with a long. Acrorbis. The presence of a fiagellum indicates a relationship with Remarks. Pilsbry has stated (1938. although the diverticula are more crowded than has been observed in species of Drepanotrema examined. the shape of the penial complex. n) is long and narrow without a superposed ridge.

fossil groups Vorticifex. Helisoma Fischer and Crosse.. FW. With the exception of Planorbarius. The in this Genus HELISO:VIA Swainson.ssix. These might have been present but overlooked because of tiic small size of the radula teeth. p. Helisoma Binney.'< bicarinatus Say. II.. The very long radula sac is also a characteristic feature of the anatomy. Y. p. As subgenus. pp. which appears to have characteristics admitting it to this group. 351. 106. With two penial retractor muscles: Carinifex and Parapholyx. 1865. but the ectocone and entocone are much shorter. 1884. A. Malac. 112. S. 33. the subfamily is wholly American and is perhaps the most important planorbid element in the North American molluscan fauna.. Terr. Tj^pe Planorbis bicarinatus Say. C. Type genus Helisoma Swainson. Type Planorbis bicarinatus Sowb. Conch. Genera. Helisoma Cle. bicarinatus S:iy. related to Drepanotrema. I. Mai. The groups referred to this subfamily are noted below. Lye. A penial gland is nearly always present with an external duct of greater or lesser length. Helisoma Tryon. Type Planorbis bicarinatus Sowb. therefore. Conch. p. Fluv. Moll. of three pieces. Univ. 1880. Syst.. Sh. 1870. Ill. II. It is widespread throughout the northern part of North America and the species are numerous. 1886. and Subgenem —-Recent and Fossil 123 overlapping whorls of Drcpanotrona. original designation Planorbis bicarinatus bicarinatits Say Planorbis anceps = Sowerby (= Planorbis Menke) Helisoma Sw. The jaws are sim})le. 1872. No type cited. With one penial retractor muscle: Helisoma and subgenera Seminolina. U.. Baker. 60. As subgenus. No type cited. Subfamily HELISOMATINAE F. the prostate exhibits a fanlike design. 337. The radula appears very different from that of Drepanotrema (see plate 68) because of the apparent absence of interstitial cusps between the larger cusps. p. et Guat. and FW. p. 198. As Helisoma Tryon. i).^nnsen. Without penial gland duct: Planorbarius. As subgenus of Planorbis. Prim. N. 516. In cross section. a superior and two lateral jaws. 1846. Helisoma Dall. Moll. Mon. Type Planorbis Tyjie Pl((uurbi. As subgenus. L. The radula is markedly different from that of Tropicorbis. Ann. As subgenus. 1928 Prostate diverticula compound or multiple. Mexico subgenus. S. H. but distinct by both shell and anatomy. IX. 188. appears to be another one of the peculiar groups of South American Planorbidae. Aa^orbis. 1840 Type by 1840. . Helisoma Herrm. X.. Gen.. consisting of a basal stem with branches grouping around a short prostate duct which is in intimate contact with the sperm duct. As subgenus. Picrosoma and Planorbella.Subfamilies. and Pompholopsis are included subfamily because of likeness of shell to Carinijex or Parapholyx. XVII. Ind. Treat. p.\ixsoN. and S. Perrinilla. As subgenus. The general shape of the mesocone in Odhncr's figure (4) is suggestive of the same cusp in Drepanotrema. N. The kidney is usually heavily ridged.

47. As genus. Proc. 1. HeUsoma F. 9) the prostate is fan-shaped. p. No. No type cited but includes Pierosoma. composed of many small follicles. 479. Baker. Fresh-water Moll. 1 mm. X. Soc. Trans. As genus. Type Planorbis bicarinatus Say. Zool. No. Fossil. London. HeUsoma Dall. aperture expanded. Chi. Plaiiorbis bicarinatus Say. anceps (Menke) Seminal vesicle (SV) short (about and widely rounded. and the mantle is spotted with dark mottling. sinistral. p. Amer. As genus. HeUsoma F. Proc. long. 6.. II. GENITALIA. 43. As subgenus. Baker.. Mus. carinatus Sowb. there are s]5ots of color near the eyes and between the tentacles. . London. Mich. Bull. especially over the kidney. Univ. in diametei'. Anterior to the seminal vesicle. Type Planorbis bicarinatus Sowb.). Cent. p. Baker. Ind.. Biol. 287. Type Planorbi^s bicarinatus Sowb. XXI. As genus. pp. Cat.5 mm. C.5 mm. p. Occ. flecked with whitish. p. pp. p. Acad. Moll. p. Type Planorbis carinatus Say.. Ser. spire and umbilicus funicular. Pars 22. Papers. As genus.. Rec. Sci. long) . 584. Soc. As subgenus. Acad. Genotype Planorbis bicarinatus Sowb. Type Planorbis anceps Menke Shell (plate 80. Univ. in height. In section (fig. Proc. No type cited. about 2. Type Planorbis bicarinatus Sowb. Yellowish or brownish. p. 1902. Univ. Utah. 11. Type As subgenus. I. Area.3 mm. 1928. It is composed of numerous diverticula forming a pavement-like pattern when viewed externally. The prostate (PRS) is ovate in shape. HeUsoma Germain. 1934. 210. fig.124 1899.. 381. for the space of 2. 1926. Genotype HeUsoma bicarinata (Sowb. As genus. of comparatively few whorls which rapidly increase in size. 1918. long and is of wide diameter. Type Planorbis As genus. Zool. 1912. p.. As genus. Type Planorbis bi- 1931. Wis. 1905. p. 81. HeUsoma Thiele. Miscel. HeUsoma Wenz. 12) The vas deferens (VD) 11). Baker. HeUsoma Chamberlin and Jones. HeUsoma F. Mai. 1-6). 64. Baker. carinated.. 1930.. Biol. 311. bica- HeUsoma Pilsbry. Mich.. As subgenus. rinatus Say. ANATOMICAL CHARACTERISTICS PLATE 1 23. 145. wide and less than 1 mm. No. Teil 2. C. Phil.. C. HeUsoma F. 201. Large. Type Planorbis Subgenus HELISOMA SS. XXII. As section of Planorbis. somewhat sausage-shaped. The shell is carried almost perpendicularlv. figs. composed of six to eight diverticula each of which is branched two to four times toward the distal end. The Molluscan Family Planorhidae HeUsoma Martens.. The tentacles are long and filiform and the velar area is notably developed. antrosus Conrad. HeUsoma Walker. HeUsoma Hannibal. p. Alaska Moll. The prostate is placed on a short prostate duct which unites with the sperm duct to form the vas deferens (plate 22. 157. Handbuch. Mus. 6. 84. Mus.. Moll. C. Nat. 1923. Animal.. Zool. As subgenus. Wis. Pub. HeUsoma (fig. HeUsoma H. 1931.. Arts and Sci. B. Male Organs mm. 1929. The sperm duct (SPD) is about 6 mm. 1921. p. outer lip thickened. 1522. the ovisperm duct is bordered by many small follicles which may represent a portion of the seminal vesicle (SO). 86. Planorbis bicarinatus Say. the tube about 0.

The floor of this cup is covered about 14 is still It with pavement-like folds. In the preputial space there is one large. somewhat smaller in diameter than the sperm duct.5 mm. The penial complex may assume a number of shapes. 6) shows the relationship of the muscular ring or diaphragm (D) to the gland duct. 5). Often it is seen as shown in fig. wide. At the terminal end of the verge the seminal opening is placed in the center of the organ with a small. vertical fold or pilaster (PL) and several smaller vertical ridges. solid band or it may be broken up into ten or twehT smaller muscular bands. an upper vergic space and a lower preputial space. long). 1 on plate 22 {Helisoma anceps latchfordi) it is shown well-extended. 8. The supporting muscles are attached to the roof of the mantle cavity. 1) this gland is seen to have the cupshaped. This group may form a wide. Attached to the wall of the preputium at the upper end of the large pilaster is a rounded. separated by a muscular ring or diaphragm (D) which is capable of great contraction and expansion (see fig. cylindrical form. fig. In fig. 11. Tlic penial comjilex (plate 23. cup-shaped penial gland (GL). depending uj^on the sexual condition of the animal. A section through the region at the junction of the two spaces (fig. There is no marked constriction between the preputium and the vergic sac. from which a narrow duct (DC) extends through the neck of the gland and emerges through the bodv wall of the preputium and becomes the external penial duct. plate 22. This penial duct tapers to half its diameter as it enters the junction of the prejiutium with the vergic sac. exhibiting an elongated. an upper portion with smooth or only slightly developed vertical fokls (OC) and a lower portion (LC) lined with heavy vertical folds (GF). bulbous termination divided into two parts. The enlarged epiphallus is also indicated. VD). and Subgenera is — Recent and Fossil 125 mm. There is a short (2 mm. There is a wide retractor muscle (RM) attached to the preputium just above the male opening. 3) is a short. In fig. .5 mm. the latter penetrating the thick wall of the lower portion of the vergic sac and opening into this cavity just above the diaphragm. fig. sharp appendage or papilla extending slightly below the end of the verge. The penial gland and the verge may sometimes be seen l^rotruding from the male ojiening and lying on the neck of the animal. 2) the penial comjilcx shows two cavities. Genera. long. In fig. plate 23.. long and 1 mm. Subfamilies. long. the penial gland pushed upward causing the com]ilex to assume a bilobed aspect. On the right side of the preputium there is a group of supporting muscles (S^I).) and about 2.7 mm. the supporting muscles (MS) plainly indicated at the upper end near the top of the vergic sac. 2. 3. external tube or duct (DC) on the outside of the preputium and adherent thereto. 1. V) is normally short and wide and tapers abruiitlv to a narrow point. In section (fig. long. narrow. The vergic sac (VS) is somewhat bullet-shaped. the end of the verge is shown with this feature enlarged to show the pointed pajiilla and the centrally located seminal opening. somewhat pyriform organ about 4 mm. 10 on plate 22. Internally (fig. fig. The preputium (PR) is very wide (1. the outer cup of the gland is turned back to expose more fully the folds of the inner cuji (Helisoma an reps percarinatian) Tlie A'orgc (plate 23. further diminished in diameter as it emerges from the muscular tissue to enter the jienial complex (plate 23. Near the summit of the vergic sac the vas deferens enlarges to form an epiphallus (EPI).

Near the albumen duct the lobules are much larger than in the body of the gland. It narrows notably as it nears the anal end of the rectum. which is rather wide and enlarged as it enters the long (1. long.wise from the ovisperm duct. A cross section of the kidney of Helisoma anceps latchfordi (fig. 4. There is a large and conspicuous superposed ridge extending from just below the pericardium to the urethral portion of the kidney and going beyond the kidney almost to the mantle margin. are remarkably uniform. 8). between the seminal vesicle and the ovotestis. the ovotestis is seen to be composed of about a dozen diverticula extending fan. Female Organs (plate 23.126 The Molluscan Family Planorhidae is The verge often extended into the preputial cavity as 22. The albumen gland (fig. 2. In section (fig. 4).) and narrow vagina. The ureter is less than 1 mm. an abnormal condition. The pseudobranch (plate 23. wide. sausage-shaped and slightly less in length than the duct. long and 0. as shown in fig. Respiratory and Renal Systems. The uterus (U) is about 5 mm. with a distinct ridge passing down the center from the rectal region.3 mm. 11). widening upward to the nidamental gland (NG). long and 2 mm. The pericardium is nearly 2 mm. is seen to form a long fold which extends over the kidney causing it to appear rounded in a ventral view of the kidney.5 mm. the (fig. tapering to about 1 mm. There is a large carrefour (CD into which the small. 11). high. as far as they have been examined. A depression crosses the gland on the under side marking the space occupied by the intestine. narrow duct of the albumen gland enters (DA). long and 1 mm. long. The kidney (plate 44. 3. long and is sharply reflexed parallel to the long axis of the kidney. 7). wide. wide at the lower part. 1 marked by the arrow. in diameter and is composed of small follicles. this being often divided into several branches toward the columella muscle (plate 22. The genitalia of the races of Helisoma anceps. Michigan. mm. 1) is 7 mm. this ridge. figs. fig. taken at about the middle of the kidnev.5 mm. In one specimen (fig. in both regions. about 2. The spermatheca (S) and its duct spermatheca elongated. The ovotestis (OT) consists of of club-shaped diverticula having a pavement-like appearance from the surface of the gland. long and 1 mm. and 4 on pLate are 5 fig. in diameter. rounded below. In specimens of Helisoma anceps Hermaphrodite Organs many rows percarinatum from Douglas Lake. This ridge in the mantle is present in all species of the family. fig. P) squarish in shape. but about 1 mm. 1. than between the seminal vesicle and the oviduct. The free portion of the ovisperm duct (SO) is somewhat shorter. In cross section (fig. pad-like. shown in figs. 10). The oviduct (OD) is about 4 mm. at the i)oint in fig. A small ridge to the left of the lumen is shown in the figure. which is about 2 mm. wide at the upper (pericardial) part. wide and about 1 mm. The lumen is regularly rounded and the two blood vessels appear on each side at the lower part of the section. 4) there were two muscles. 7) is squarish in form. This ridge extends backward on the rectum about a cjuarter of a whorl. there was some variation in the form of the retractor muscle. The rectum is very wide and there is an irregular crest or ridge extending down the center (R). The pneumostome (PS) is not notably large. The pseudobranch is 2 mm. The larger portion of the ovisperm duct is occupied by the seminal vesicle. shows a less extensive ridge (RK) is . 3. 2.

Baker. fig. is figured on plate 53. The superior jaw is finely striated on its The intestine backward in anterior face. The extreme outer marginals are vestigial. wdth one large tooth (the mesocone) in the center (see 14). The radula has the variable formula 22-1-22 to 30-1-30. There is one wide. 3 2 27-1-27 to 27-1-27 1-5 6-7 Plate 53. The ectocone of the fourth tooth is split into four small cusps. Plate 53. loops around the stomach. intermediate teeth. The mesocone becomes somewhat smaller but is not modified. The table below indicates the extent of this variation. low. fig. fig. Wisconsin 25-1-25 to 27-1-27 1-6 Cleveland. 4 Douglas Lake. The outer marginal teeth appear like a saw. fig..Subfamilies. lateral teeth. fig. fig. as is shown in several sj^ecies of Helisoina. The entocone is broken up into two or three small cusps and several small cusps appear on the outer edge of the tooth above the ectocone. Oneida L. passes over the oesophagus. Conneotiout 22-1-22 to 25-1-25 1-7 ChautaiKiua Lake. HcUsoinn anceps sayi F. In the marginal teeth the entocone is broken up into fine serrations numbering from five to ten cusps. The columns are all as follows. A radula from specimens from Maple River. turns a long loop around the liver and then runs forward to the l^seudobranch in the form of the rectum. the mesocone the longest and the ectocone the shortest. The buccal sac is short-ovate. this feature increasing toward the outer margin of the membrane where the teeth are ^'ery long and narrow (14). Genera. 1 fig. and Subgenera — Recent and Fossil 127 but is otherwise similar to typical anceps. reading from left to right: Locality. arched superior jaw (about 1 mm. ribbon-like lateral jaws somewhat longer than the superior jaw. Michigan. 1 on plate 48. The cusps are spade-shaped. 1). with 150 to 193 rows of teeth. . Wisconsin Helisonta anceps percarinatuni (Walker). Michigan 23-1-23 to 25-1-25 1-5 I'nionville. Helisoma anceps (Menke). Bayfield. 6-7 6-7 8-9 7-9 7-S Plate 53. The stomach is bulbous and tapers into the pyloris. The salivary glands are narrow and about twice as long as the buccal sac. The jaw (plate 49. The marginal teeth (8-14) are much narrower. The center tooth is squarish with two short. 23-1-23 to 24-1-24 1-5 Maple River. The lateral teeth (1-5) have a squarish reflection which extends well below the lower margin of the base of attachment. principally in the numerical formulae. Michigan 27-1-27 to 30-1-30 1-8 9-10 Plate 54. The inter- mediate teeth (6-7) are narrower than the lateral teeth. Plate 53. 1 There are 193 rows of teeth. 1. They are joined behind. C. The general form is like that of HeUsomn trivolvis shown in fig. wide cusps. the base of attachment very short. The radulae of anceps and its races vary considerably. New York There are 150 to 160 rows of teeth. and figure. but also in the number and position of the cusps. The ectocone is distinct in the 8th tooth with three small cusps above it. plate. The form of the section may vary when taken from different parts of the kidney. formula. wide) and two narrow. From the 10th tooth this area changes to small serrations five to six in number. New York 26-1-26 to 27-1-27 1-6 Moo^ie Ear Creek. Digestive System. the number increasing in the outer teeth (compare 8 and 14).

C. the first and second laterals of the 100th row had the entocone split into two smaller cusps (plate 54. the entocone was split into two small cusps (plate 53. fig. Michigan. A specimen of latchfordi. 3 1-6 7-9 Plate 54. antrosus Conrad. lower row). All of the laterals in this row had the entocone broken up into two small cusps (plate 53. 60). 4). C. 3. fig. lower row). Ontario There are 160 rows of teeth. The 8th and 9th teeth of this row had the intermediate in unusual form. the same tendency to form abnormal teeth is observed. New York. 2). fig. This row shows the shape of the teeth on the left side of the membrane (plate 53. Ontario There are 170 rows of teeth. Abnormal teeth arc connnon in radulae of Heluoma anceps. 1-S 9 Plate 54. the typical subgenus of the genus Helisoma. the fifth lateral of the ninetieth row had a split ectocone. had the central tooth with an extra cusp on each side of two smaller central cusps (plate 54. 1-6 7-9 Plate 54. both from the same locality and from different places. Baker. there is complete agreement in all radulac examined. in the sixtieth row. Baker Helisoma anceps latchfordi (Pilsbry) Helisoma anceps portagensis (F. Species Considered as Valid. C. Baker) Helisoma anceps cahni F. 27-1-27 to 30-1-30 Meach Lake. fig. lower right row). 4. fig.a anceps royalense (Walker) Helisoma anceps ruski F. Michigan. 2 Helisoma anceps cahni F. contains one sj^ecies with many races and a second species with two recognizable races. In a specimen from Maple River. The species of typical Helisoma inhabit North America from British America (Hudsons Bay) southward to Louisiana and Alabama and westward to the Rocky INIountains. C. C. fig.128 The MoUuscan Family Planorbidae Helisoma anccps royalense (Walker). 1. 4 For the above data more than forty siiecimens have been examined. Helisoma anceps anceps (Menke) 1830 (=^ Planorhis bicarinatus Say 1819 (preoccupied). Helisoma anceps latchfordi (Pilsbry). P. 27-1-27 to 27-1-27 Big Muskallonge Lake. fig. In a radula of percarinahmi from Douglas Lake. Baker Helisoma anceps percarinatum (Walker) Helisom. 1834) Helisoma anceps unicarinatum (Halde- man) Helisoma anccps striatum (F. the numbers of lateral and intermediate teeth likewise vary. had the center tooth of the seventy-fifth row with three small cusps on the left side and one small cusp on the right side of a central cusp. Geographical Distribution. fig. C. This. A specimen of saiji from Bayfield. Connecticut. C. In a specimen from Unionville. In a specimen from Chautauqua Lake. In general form of teetli. All of the teeth of this race examined had two very small denticulations on the edge of the reflection above the ectocones (plate 54. the fourth lateral of the fifteenth row had the ectocone broken up into four small cusps (plate 53. Baker) Helisoma anceps aroostookense (Pilsbry) Helisoma eucosmium (Bartsch) Helisoma eucosmium vaughani (Bartsch) Helisoma anceps sayi F. In the races of anceps. 4. 1). 25-1-25 to 27-1-27 Bamiji Lake. In the first lateral of the same row. fig. The formulae vary and along with this variation. Baker Helisoma anceps shellense F. the center tooth of the 118th row was broken up into three small cusps and one large cusp. fig. Wisconsin. Quebec There are 190 rows of teeth. Baker Helisoma anceps jordanense (Winslow) .

p. as has been made by some authors. p. Proc. Teil 2. Binney based his opinion evidently on the lower fig. 188). Typical Hclisotua is characterized by a short and wide. and generallyFrom Seminolina. Genera. not of Signoret. Helisoma differs from Pierosoma gland duct. p. 4 of the plate. 31. Pliocene to Recent fauna. Adams. 479. 1934. There is a cup-shaped penial gland with a short. Sowerby 's from 1824. 311). In 1931 (p. 1931. There is little question about the upper figure being intended for bicarinatus. Baker elevated the group to generic rank. The citation of Planorbis bicarinatus Say. Helisoma has been known in molluscan literain its shorter penial different penial complex. 356. Example Phym scalaris Jay p. The prostate and ovotestis are made up of many small diverticula radiating from a duct in fanlike i)attern when seen in cross section. The' shell of Helisoma is also quite different from Pierosoma and Seminolina in its general bicarinate form with the spire and umbilicus both deeply concave. 84) cited the type as Planorbis bicarinatus (Say) Sowerby. Binney (1865. and Subgenera — Recent and Fossil 129 Geological Distribution. Handbiich. it differs by its shorter duct. 124) thought it might be Planorbis campanulatus Say. 358. p. A careful study of Sowerby's description and his two figures on plate clxxvii (fig. Baker. 139). Type Helisoma . In 1926 (p. the genus was diagnosed by Baker and the distinctions of the genitalia described and figured. and A. also be neuter (H. Conrad's Planorbis antrosus 1834 has been used in place of bicarinatus. which resembles Helisoma in its short i)enial gland duct. 584). The Planorbis bicarinatiis of Sowerby. but both figures and description point to bicarinatus. 81) cite the name. has nothing to do with Say's species and Sowerby was probably not aware that there was another bicarinatus. basing the distinction on the peculiar genital complex of the male organ. 201) and in 1928 (I. Anatom3^ As subgenus of Helisoma. 1886. Amcria Dall. 112). Authors generally have misquoted the type of the genus. (Jay). Binney (1865. and Dall (1905. 1879 (Hescalaris Thnmsonia Thiele. Seminolina Pilsbry. X^at. onlv characteristics of the shell for its distinction. Phil. p. IX. original designation Helisoma scalare (JajO ]>. p. (non H.><te. Acad. is also incorrect. wider and shorter vergic sac. somewhat bulbous penial complex without a notable constriction between the preputium and the vergic sac. p. and might be mistaken for cam'panulatus. 86. of course. while with little doubt the same as Planorbis bicarinatus Say. Sci. but ]\Ienke's name precedes this by four years and must be accepted. it is a Greek noun in the neuter gender and all specific names must. ture for many years. Say's species dates from 1817. Lye. Type Planorbis p. Miocene?. Y. X. rn ipt era). Unfortunately this appropriate name can not be used. Remarks. Ann.. 1934 Type by 1870. 1930. Mil. narrow. As section of Planorbis. Tryon (1870. as it is preoccupied for a Paris Basin fossil. p. Dall (1905. 4) leaves little doubt about the unity of his species with that of Say. 1855). The radular teeth also differ from Seminolina in having the cusps sharply aculeate instead of rounded. H. Thomsnnia Axcey. which is incorrect.. almost straight penial gland duct.Subfa/nilies. N. Regarding the name Helisoma. Le Xaturali. using. however. Subgenus SEAIINOLINA Pilsbry. B. As a subgeneric name. which is not clear. although the type is correctly cited on page 81. scalare (Jay).

sinistral. fig. with which the sperm duct connects posteriorly and the vas deferens anteriorly (see fig. The i^enial complex (fig. The ovisperm duct bears many small vesicles along its side for over 1 mm. varying in width. The cavity of the pre- is partly filled by an elongate-ovate (irregularly paralleliform) penial gland attached by a short neck of less diameter to the body wall putium . With large velar area and long tentacles as in Helisoma. 9) the preputium was pushed upward sac appeared as though attached to the lower part of the this condition. Color variable. The vas deferens (VD) is about 10 mm. wide at the center and about 0. There is one large retractor muscle (RM) which is split into four branches at the preputium and in some specimens (fig. 9).5 mm. split into three branches at the upper part of the preputium.3 mm. purplish on top of head. each with two branches. In some specilike rows. and the vergic preputium. wide at the male opening. 14). in length composed of many diverticula arranged in fan- A section of the prostate near the middle shows sixteen rows of club-shaped diverticula (fig. but 1 mm. in duryi and varieties dark olivaceous in old specimens. ovate-pyriform in shape. with every gradation between these forms. without the thread-like striae of Pierosoma.5 mm. 14). The lower sac or prei)utium (PR) is pyriform. The muscles are attached to the upper part of the preputium. wide near the upper part of the sac. Mantle yellowish or grayish over lung and kidney. In one specimen there were five narrow.5 mm. The prostate diverticula discharge into a separate duct. the prostate duct. Internally (fig. Animal (plate 70. with few black or brown spots. long. vertical ridges or pilasters 11). below the seminal vesicle. wide anteriorly and posteriorly of this area. long. Seminal vesicle (SV) about 2.5 mm. Male Organs (fig. The seminal vesicle is composed of many small glands. in length. 9) there may be one wide muscle of three branches with a narrow. The vergic sac is of about the same length as the preputium. In the preputium This condition is observed in specimens in breeding stage. usually glossy. 3 (adult). separate supporting muscles. It is slightly enlarged (epiphallus) as it enters the vergic sac. Large. the internal organs yellowish or pinkish. usually widely or deeply umbilicated.130 The Molluscan Family Planorbidae Shell (plate 80. Immature specimens are more yellowish with few or no dots. figs. the penial gland duct extends from the top of and enters the vergic sac below the level of the duct's exit. In scalare olivaceous. surface smooth. 1. Sperm duct nearly 3 mm. fiecked with white. ANATOMICAL CHARACTERISTICS PLATE 28 GENITALIA. The duct of the penial gland (DC) is rather short (3. slight. 10) is about 3. long. fig. of large diameter. 6). . physa-shaped or planorboid. Bottom of foot dark horn or purplish. over 2 mm.5 mm. 2 (young) ). and is about half the diameter of the sperm duct. 7-12). wide at the upper part and less than 0. The supporting muscles (SM) usually form a wide band. long) and tapers in diameter toward the end entering the vergic sac (fig. all placed near the lower end of the preputium (fig. Prostate (PRS) oblong. 12). fiecked with fine light dots. The young animal is hyaline and transparent. the preputial sac contains several (four or more) (PL). Tentacles lead color. separate muscle below. narrow. mens examined (fig. about 1 mm.

11.5 mm. 18). 17. The penial complex is usually short and wide but varies somewhat in this respect (see figs. the short gland duct. The albumen gland (AL) is more irregular in shape.5 mm. the vergic sac. long) and sac-like. The spermatheca (S) is bulbous. long). The free portion of the ovisperm duct is also longer. The vas deferens (VD) is a long and narrow tube (over 20 mm. plate 28). 18). two-cupped penial gland and the lai'ge. wide and about 1 mm. GL. Genera. The vagina narrows to about half its diameter at 1 mm. The uterus (U) is about 3 mm. rather wide and increases notably in diameter at the lower end where it enters the short vagina. 17. The genitalia of Helisoma scalare examined agree perfectly with the ( I figures published by Pilsbry (1934. The peculiarities are the wide and pyriform penial complex. long. The prostate duct is longer and the space between the entrance of the sperm duct and of the vas deferens is several times wider. fig. one. has several large glandular folds (GF) and the other. long) and is more elongate-ovate in shape. 1. b. In Helisoma duryi normale Pilsbry. long the female opening (VG). sacculated seminal vesicle. The verge (V) is pyriform or reversed conic 0. 7. 9. Female Organs (plate 28. There is a narrow ridge or diaphragm which separates the vergic sac space from the preputial cavity. The sperm canal has a central outlet.9 mm. this gLand is seen to be composed of two cup-like cavities. long and is as wide as the diameter of the spermatheca. the genitalia are much elongated. The free portion of the ovisperm duct is very short (about 0. long). 14). Normally (figs. The gland duct enters the vergic space above this dial^hragm as shown in fig. long) and the gland sacs are larger. 15). The larger folds extend transversely across the cavity as shown in fig. Its duct is longer (5. c. the preputial sac is as long as.7 mm. 8.. AL) Hermaphrodite Organs. In section (fig.5 mm.) both anterior and posterior of the seminal vesicle. the outer cup.5 mm. Subfamilies. The sperm duct (SPD) is a long and narrow tube (7. The small cup is at the extreme end or tip of the gland. DC. very wide. wide at the upper part and tapering to a point. d). 11. The albumen gland (AL) is somewhat longer than wide and is composed of many small. has several smaller folds attached to the wall of the cup (OC). fig. as shown in fig. The seminal vesicle (SV) is much shorter in comparison (2 mm. The oviduct (OD) is about as long as the uterus. The prostate (PRS) is more elongated (5. Its duct (SPD) is 2 mm. due to the more widely-coiled shell (fig. The ovotestis occupies the first and a part of the second whorls and is composed of ten or more rows of club-shaped diverticula of unequal length (fig. 12. gland-like vesicles (fig. The gland duct enters the smaller cavity. the inner cup. DG). long. The nidamental gland (NG) is short (about 1 mm. The spermatheca (S) is much longer and larger (3. and bears the nidamental gland about midway of its length. long) and contains a greater number of diverticula. the elongated. and is always . 14. The several varieties under the name of Helisoma duryi (Wetherby) show some differences from scalare in their anatomy (plate 33). and Subgenera — Recent and Fossil 131 of the prepiitium through which the penial ckict extents (GL.) and much narrower. 7). or only a trifle longer than. 13. The uterus and nidamental gland are longer and narrower and the oviduct (OD) is narrow and much elongated.

d. The gland of a half-grown specimen is siiown in fig. The stomach and intestine are as in Helisoma. 1. high. normale (fig. 1. fig. the superior jaw with vertical striations on the inner face. eudiscus. long and is bent backward against the lower part of the kidney. 25) shows a rather long.5 mm. 4) of . In fig. the preputial sac has been pushed upward as in normale. narrow crest or ridge. Figure 6 shows the wider penial complex in an immature specimen with a shell 6. the jienial gland is short and wide. in diameter. 8) and wide (Pi. to the right of the anal opening (A). There is the same division into an inner and an outer cup as in scalare. In Helisoma duryi. there is considerable variation. duryi. scalare. Figure 8 shows an abnormal form of penial complex and fig. The penial gland in the duryi complex is of about the same shape as that organ in scalare. The kidney of Seminolina is more depressed than in Helisoma and the lumen is comparatively larger. while in fig. i. 19. A cross section of the kidney near the middle (fig. d. p. In fig. as are also the buccal sac and salivary glands. 4 and a section in fig. 10). 16. the gland duct is swollen in a bilobed form. long) and comparatively narrower (3 mm. The ureter is directetl diagonally backward. the penial gland was extended from the male ojiening and lay outside on the neck of the animal (fig. fig. As Pilsbry remarks on page 36. 12 shows another in which the part of the preputial sac containing the i)enial gland has been pushed upward. fig. the anatomical differences are sufficient to separate scalare from duryi and its races. 34. In fig. In fig. the penial complex is very wide. fig. The two veins are placed near the lower border of the lumen. There is usually more of a constriction between the preputium and the vcrgic sac than in scalare. In Helisoma duriji (Wetherby). a longitudinal section shows the same features. A cross section of the kidney near the middle ( a hirge cremilated ridge on the rectum iR) which runs out on the jiseudobranch in a curve. the external form is shown in fig. fig. long.5 nnu. duryi normale) as in Helisoma. the kidney is longer (17 mm. The small ridge in the mantle is shown to the left in both figures. The figures of the duryi complex agree with those by Pilsbry 1934. Figure 5 shows the penial gland pushed upward in the lu-eputial sac. H. In Heliso))ia duryi eudiscus Pilsbry. f. There is a large pneumostome. long). nortnale (fig. In H. It is contractile and its size varies greatly in preserved specimens. In H. the crest or ridge bent more to the left but of about the same form. normale). H. d. k). a small ovate lumen and two small veins placed just below the middle of the lumen. h. 13. wide). is shoi't . The pseudohranch (plate 28. In H. The penial gland duct is short. The lumen is more elongated and about two and one-half times as wide as high. fig.) than in scalare. The pericardium is al)out 1 mm. 2 it is very narrow. g. j.^c(dare is rather short (4 mm. The ureter is about 2 mm. Digestive System. The kidney (i)late 44. 14. 26) shows it to be more compressed than that of scalare. long and about 1 nun. ( Respiratory and Renal Systems. The pseudobranch is 1. as in scalare. 5). d. 3. it is shown just below the female opening. 9. Jaw (plate 49. 7. There. wider at the upper part near the ])cricardium (1 nnn. this feature is still further emphasized. In two specimens examined.132 The Molluscan Family Planorbidae wide. 4. 16.

In a specimen from Paines Prairie (fig. From Paines Prairie. on canal west of Miami. The mesocone is roundly blunt. aculeate cusps. the entocone as in the laterals. Shay. Florida Bridge No. duryi normale. A. The number of intermediate teeth is smaller in normale than in scalare. d. The marginal teeth (12-21) are very long and narrow and claw-like. p. Louis. the entocone and ectocone broadly aculeate. Boynton. Palm Beach County. near Gainsville.Subfamilies. 2). In another specimen from the same locality. several membranes gave 28-1-28 and 29-1-29 with 165 rows of teeth. Florida State Museum Santa Fe River. Florida Canal west of Boynton. St. In scalare. The outer ( . the two broadly aculeate cusps of which reach the lower margin of the base of attachment. 40 miles west of Miami. which is tricuspid. Genera. For the anatomical data herein presented thirty-five specimens have been examined from the following localities. The figures on plate 64 agree with those published bv Pilsbrv (1934. Two rows are figured. van Hyning. of normale from the Tamiami Trail is 28-1-28 to 31-1-31 with 175 rows of teeth. H. On the eleventh tooth. F. the 55th and the 140th. Collected by T. These specimens vary toward the race intercalare. High Springs. 32. In specimens from High Springs several radulae gave 27-1-27 as the formula. The two cusps of the center tooth do not reach the lower margin of the base of attachment. van Hyning . la). McGinty. Missouri Paines Prairie. but narrower. 1. The marginal teeth (13-29) are similar to those of scalare differing only in being somewhat narrower. The entocone becomes very wide and is split into from four to seven small cusps. and Suhgeiiera — Recent and Fossil 133 Radula (plate 64). the teeth are more like those of scalare. all in Florida: Helisoma scalare Lake Butler. The formula teeth are small and vestigial. 3). the center tooth (figs. extends below the base of attachment. McGinty. The variation in formulae is from 27-1-27 to 31-1-31 for the complex of duryi and to 33-1-33 for scalare as examined by Pilsbry. E. 4). the teeth are all narrower than those of scalare. 34. p. fig. Andrews of the Johns Hopkins University Helisoma duryi normale Tamiami Trail. the entocone is split into two short. eudiscus from Silver Springs gave a formula The formula of 27-1-27 to 28-1-28 with 154 rows of teeth. Boynton. The laterals (1-11) and the intermediate tooth (12) are narrow but the cuspidation is the same as in scalare. The intermediate teeth (9-11) have the mesocone as in the lateral teeth. the teeth are somewhat smaller (fig. Collected by Dr. The mesocone is aculeate in all teeth but is always distinct. The ectoconic region on the outer margin of the tooth bears three very small cusps. 35) A number of membranes were examined but none gave the larger formula. from the Tamiami Trail (fig. Collected by Paul P. for scalare is 27-1-27 to 29-1-29 and there are 160 to 190 rows of teeth. The lateral teeth (1-8) are about as broad as high and the reflection. Collected by T. In H. The ectocone in all intermediate teeth is split into two sharp cusps. although the shell is much larger. Collected by W. Pilsbry gives 33-1-33 as the formula 1934. 89) is ahiiost twice as wide at the base as at the summit of the reflection. Collected by Paul P.

Moll. Species Considered as Valid. A. Includes mostly species of Pierosoma.. The physoid aspect of its type species. narrower. Helisoma (Seyninolina) scalare (Jay) Helisonia (Seminolina) duryi (Wethcrby) Helisoma (Seminolina) duryi seminole Pilsbry Hrlisoina (Seminolina) duryi intercalare (Pilsbry) Helisoma (Seminolina) duryi eudiscus Pilsbry Helisoma (Seminolina) duryi normalc Pilsbry Helisoma (Seminolina) preglabratum (Marshall) For an extended account 1934. Subgenus Tjqie 1865. de Conch. H. Sh. Includes . duryi intercalare Canal west of Boynton.. this group is found only Bradford County. FW. Amer. 198. 1899. These include normale. Biol. is perhaps more variable than any other species found in America. F. but in the variations of the races of another species. is a characteristic group of Helisoma. Paludina scalaris Jay. of the species of this subgenus see Pilsbry Geological Distribution. eudiscus. intercalare. and duryi. Collected by T.. Seminolina is a notable group of the subfamily Helisomatinae and one of the most variable genera as regards species. Helisoma Fischer (non Swainson). As far as known. its extremes being from typical Physashaped to flatly discoidal shell. as normale. pp..134 The Molluscan Family Planorbidae H . The elongation of the spire always produces a physoid aspect. intercalare. S. p. 381. led the older conchologists to include it in Physa and in the genus Ameria of the family Bulinidae. PIEROSO^IA Ball. and duryi. Helisoma duryi (Wetherby). The duct of the penial gland is about twice as long as in Helisoma and the penial gland is more elongated. High Springs. In its anatomy. Helisoma Remarks. were completely obliterated. U. differing from typical Helisoma in the presence of a physoid shell. van Hyning Silver Springs. Collected by T. trivolvis Say. Moll. one with in the peninsuhi of Florida north to several races. species now referred to Pierosoma Type Planorbis p. Three species are recognized. Cent. Mon. The Helisoma duryi complex includes several races more or less heavily infested with parasitic worms. As section of Planorbis Helisoma Martens (non Swainson). 112. Collected by W. It is particularly abundant in the Everglade region of southern Florida. 1905 by original designation Planorbis trivolvis Say Helisoma Binney (non Swainson). Land and FW. As subgenus 1883. N. 509. Helisoma Tryon (non Swainson). II. it differs from typical Helisoma in the rounded mesocone of the lateral teeth wiiich are distinctly aculeate in Helisoma. Shay Santa Fe River. van Hyning Geographical Distribution. duryi eudiscus Marion County. Man. The races of duryi blend into each other and often three forms will occur in the same lot. SeniinoUna scalare was the least affected. not only in one distinct species. Pliocene to Recent fauna. The largest species. especially the genitalia and liver. No type cited. Includes mostly species belonging to Pierosoma 1872. Univ. p. 31-43. pp. and less cup-shaped than in Helisoma. IVIany specimens were so badly infested that most of the organs. 188.

p. the preputium rapidly narrowing as it receives the vergic sac. the foot nar- rowly pointed behind. svibgenus. As section 1934. 6.5 mm. sculpture of coarse growth lines and conspicuous spiral lines. 11) is usually about 5 mm. tentacles long and filiform. in some species reddish or yellowish. As section of subgenus Helisoma 1921. I. of few whorls. Sperm duct (SPD) long and narrow (about 4 mm. Acad. rounded before. the outer lip thickened within. as in pilsbryi. Pierosoma Germain. Pierosoma F. Genitalia and radula. . Tjq^e Planorbis trivolvis Say. pp. Soc. 13-15) Large. Baker. vis (Say. . p. Fresh-water Moll. wide.) As Fossil. the head and foot are usually lead color. Genotype H. figs. 479. XXII. 586. 11. As subgenus Pierosoma Dall. Zool.5 mm. The penial complex (fig. Handbuch. The vergic sac GENITALIA. .Suhfamilies.. long. 49. Alaska Moll. 1931. p. 135 As Zool. Type Planorbis trivolvis Say. Wis. never glossy as in Seminolina.. — Recent and Fossil trivolvis Say. Color usually dark horn. Proc. Pub. Pierosoma Wenz. H. As subgenus Pierosoma Thiele. fig.. surface usually dull. C. composed of fourteen rows of diverticula emptying into a short prostate duct which connects with the sperm duct and from which the long (15 mm. As subgenus Pierosoma F.5 24 (Planorbis trivolvis Say) . 7. The preputium (PR) is very wide (1. 1931. trivolvis Body long and somewhat narrow. of Planorbis Sci. Miscel. Type Planorbis 1923. Trans.. p. long) Prostate (PRS) about 4.5 mm. p. Baker. trivolvis macrostomum. 0. C. Mich. Pars 22. Wis. viscera reddish through the shell. Cat. (Pierosoma) trivol- 1926. Rec. sinistral. Acad.. 1522. 13) shows the fourteen rows of diverticula to be of unequal length and the whole section to be fan-shaped. Proc. even flaring in some specimens. Type Planorbis trivolvis Say. Genera. As subgenus Shell (plate 80. A cross section of the prostate (fig. long. especially on the front portion (plate 70. Baker. Mus. foot also somewhat reddish. Arts and Science. Type Planorbis trivolvis Say. the body dark horn color flecked wath white spots. London. and Subgenera 1905.. 6. 329. Teil 2.) and cylindrical in shape. 30. XXI. There is only a very slight enlargement (epiphallus) of the vas deferens as it enters the vergic sac. Type Planorbis trivolvis Say.. or without spots. pp. It occupies almost half of the length of the ovisperm duct. widelj^ umbilicate below spire of the adult shell usually flattened and sunk below the level of the body whorl aperture notably expanding. H. velar area well-developed. No. 86. C. p. 202. Jallax). 1918. The vergic sac (VS) is of the same length as the preputium but is very narrow. mantle spotted. . As subgenus Pierosoma F. fig. long and 1 mm. p. the early whorls usually flattened and carinate above. 1928. Nat. 85. Animal (plate 70. 15) wide) composed of many small vesicles.) vas deferens (VD) extends and passes to the vergic sac. . the tentacles are very long and slender. Univ. eyes at base of tentacles as in other Planorbidae. section under subgenus Helisoma P/emso ma Walker. Genitalia and radula. 6) . Phil. Pierosoma Pilsbry. ANATOMICAL CHARACTERISTICS PLATE (3. the body flecked with streaks or spots. Ind. as in trivolvis. Type Planorbis trivolvis Say. In the young. fig. the eyes black. 81. Mus. Seminal vesicle (SV) elongated (fig. 8. Male Organs mm. the last whorl large and capacious.

somewhat diagranunatic. which is wide and bulbous. A vertical section of the gland (fig. The si)ermathecal duct (SD) is 3 mm. a develoi)ment of the Pierosoma group from the HeUso))ia group. The seminal canal has a central exit beyond which a small. There is usually one wide band of sui)porting nuiscle (SM). Another form of gland is shown in fig. There is a large. and the entrance of the penial ghmd (hict.136 The Molluscan Family Planorhidae increases to double its diameter near the end where the vas deferens enters. The oviduct (OD) is short and wide (about 3. The bottom of the cup contains many small folds placed longitudinally. There is a large carrefour (fig. causing the diaphragm to be a narrow nmscular ring or a comjiarativcly thin membrane which nearly closes the opening between the preputium and vergic sac. 5). The duct is shown in fig. The walls of the cu]) (BW) are very thick. The uterus (U) is about 7 mm. 8). elongate-pyriform i)enial complex with a short penial gland duct (fig. . . The relationshi]) of the verge and the penial gland duct is also shown in this section. Figure 10. shows the relationship between the diaphragm. The gland duct (DC) enters the bottom folds of the cup. the preputial cavity (PR) contains a number of small vertical folds or pilasters. the preputium extending ujnvard until it is higher than the vergic sac (fig. bound down by many small muscles covering the preputial sac. long and 0. 7) is sciuarish and is composed of many small follicles. Ij. which is contracted near its attachment to the body wall of the i)reputium to form a narrow. The vergic sac contains the elongated verge (V) which is enlarged at the upper end and tajicrs downward to a point. The penial complex often assumes a bilobatc lorm. wide) and somewhat coiled. Internally (fig. long) and extends from the upper part of the preputium to a point just above the diaphragm (see fig. long. A vertical section of such a condition (fig. The duct (DGj passes through the wall of the j^reputium and extends to the vergic sac where it enters just above the diaphragm (DV). This condition is suggestive of typical Helisoma. to 9 mm. The duct of the penial gland (DC) is very There long and narrow (8 mm. which is a trifle too high in the figure. 5 partly uncoiled.). long. the verge. is a single. narrow. 6. Normally the duct lies coiled on the preputium just over the jienial gland and adherent to the preputium. narrow at the vagina and widening to double this diameter near the prostate where it unites with the nidamental gland (NG). triangular papilla extends. it would seem.5 mm. 10. 15. which has a wide penial complex and a short duct. 3). Female Organs (fig. as shown in fig. cup-shaped penial gland (GL) about 1 mm. The albumen gland (fig. and indicates. 4j shows the circular cup to be of considerable depth (about 1 mm. the prei)utium and vergic sac form a single wide. and somewhat enlarged at the lower end where it enters the short and wide vagina (YG). In young specimens of three and one-half whorls. 14. CF) into which the duct of the albumen gland enters (DA). the inner walls lined with many vertical folds (GF). 15) The spermatheca fS) is elongate-pyriform and about 2 nun. 2) shows the penial gland to be placed at the sununit of the preputial extension. 1. long.5 nun. In side view. it is seen to be cleft to accommodate the passage of the intestine on its lower surface (fig. IN). 8). in diameter. roundeil neck. wide retractor muscle (RM) which may be split into several branches (figs. There is a large muscular ring or diaphragm (1)1 which is capable of considerable extension and contraction.

This duct is similar in mature specimens of the different races and species. one whorl of composed being very short (about 1 mm. 1-6). In H. In fig. In fig. In fig. the shape of the entire verge of this specimen is shown. but there is some variation in the details of the sexual organs. This feature is characteristic of all members of the group Pierosoma. Characteristics of the verge are shown in figs. 13. 7. 4. four of which branch twice. Figure 3 shows the manner in which the penial gland duct is coiled on the outside of the preputium (DC). as would be expected. Figure 9 shows the entire verge which has been sectioned in two places to show^ the location of the sperm canal near the center of the verge. near the female opening. In fig. 11. pseudotrivolvis { lentum. In fig. The small papilla at the end of the verge is well shown. The large race of trivolvis (macrostomum) differs little from the typical form. It is peculiarly shaped at the distal end. and one is branched four times. In fig. Genera. 7-13. The genitalia of a race of trivolvis [lentum) from the type locality. unbranched.). the penial gland has been extended from the male opening. the penial complex is shown in its typical position w^ien the gland is exserted from the male opening. It is 15). 15 a condition in which the preputium has been mm. 9).) is provided with small vesicles along the side which are probably to be considered as parts of the seminal vesicle. New Orleans.). Figures 14 and 16 on plate 29 show the normal condition of the penial complex and fig. the verge. the penial complex is vertically sectioned showing a wider verge and a somewhat differently shai)ed penial gland. The chief characteristic is the large. from the front and from the side. there are six cliverticula entering the ovisperm duct. The cup-shaped gland (GL) is normal. 5. No figures other than those by the waiter (1923. the penial gland is shown in vertical section. the penial gland has been protruded from the male opening and is lying on the neck of the animal. that portion near the ovotestis (fig. Figure 8 shows the general form of the penial complex. In fig. and the great length of the penial gland duct is indicated. The waist-like constriction just above the middle of the verge shows where this organ was compressed when it was protruded through the muscular ring or diaphragm. In a cross section near the anterior end (fig. 2. one is simple.Subfamilies. also exserted. The form of the penial gland. Figure 12 shows the gland exserted with the verge beside it. 1931) have been published. and is shown in its typical cup-shaped condition. Figure 1 shows the penial complex a trifle wider than in typical trivolvis. the cup being somewhat diagonal. The free portion of the ovisperm duct (SO) is short. there is considerable variation. The branching is not uniform throughout the ovotestis. the preputial sac has been pushed upward placing the vergic sac on the side of the preputium. are shown in figs. and somewhat pyriform preputium and the very long penial gland duct. . The genitalia of the races of Helisoma trivolvis vary in minor details which are figured on plate 29. wide. The end near the oviduct is longer (1. Hermaphrodite Organs tlie slieU. 10.5 A portion of the duct beyond the seminal vesicle (for about 1. 9 and 10. Louisiana. t. As this gland is capable of some muscular contraction and expansion this variation in shape is of no taxonomic importance. and Subgenera — Recent and Fossil 137 The ovotestis (OT) occupies about of about thirteen rows of diverticula which radiate from the ovisperm duct. The shape and central position of the sperm canal outlet is shown in fig.5 mm. = greatly pushed upward. 6. figs.

Helisoma chautauquense F. fig. 6. 8. and consists of fifteen diverticula. 6. and the northern part of the United States and southern Canada. The prostate is shorter. the genitalia are similar in general form but show some minor peculiarities plate 26. The penial complex is wider than in trivolvis and is similar to some of the races.) and much larger. in THE GENITALIA OF SPECIES ALLIED TO TRIVOLVIS Species East of the Rocky Mouxt. A larger species. shorter and wider vergic sac. C. Variations in shape and position of the vergic sac are shown in figures 1. 2. more pyriform preputium. The genitalia are somewhat shorter than in Helisoma corpulentum.7 mm. The cross section of the prostate of truncatian differs slightly from that of trivolvis. a small race of trivolvis. The duct of the spermatheca is somewhat longer than in trivolvis. however. While regarded as a race of pilsbryi there are some features of the genitalia which indicate that is a very distinct race (plate 25. The penial gland (fig. but there are some differences.138 The MoUuscan Family Plauorbidae In H. the whole penial complex is wider. the cup is much shallower than in trivolvis (fig. 12). There are about eleven main diverticula. differs from the trivolvis complex in the wider. The genitalia (fig. Helisoma truncatum (]\Iiles). pyriform preputiiim and a small. fig. 15. In the immature stage of this species. 14. closely related species. Baker (plate 29. comparatively. and 17. The prostate (PRS) is much shorter (1. In the penial complex (fig. 7. The penial gland. 10). trivolvis fallax. 2). The seminal vesicle (SV) is of greater diameter and less length. 5). especially the vergic sac. a specimen was examined which the penial complex had a very large.\ixs A figs. living in Wisconsin. The ovotestis diverticula are very long.) and wider.) but the duct is long.7 mm. and the ovisperm duct is of slightly greater length (4 mm. The great variation in the form of the retractor muscle is well shown on plate 29. does not differ from that of trivolvis. 22). Michigan. 5. 7. 17). several of which are liranciied at the end. which are on a short prostate duct placed at the right edge of the group of diverticula (fig. lentum for example (figs. C. 19). Helisoma pilsbnji (F. but the prostate (PRS) is twice as long (5. is figured on plate 25. The spermatheca (S) is narrower and shorter (1. 1. In the penial gland.). 9) does not differ from that of ( trivolvis. and especially the much shorter penial gland duct (DC) which recalls the Seminolina group (fig. and the gland duct is very short. 16. 3. The penial gland of this specimen was quite normal (fig. 19-22). 4) are similar in general form to those of trivolvis. The seminal vesicle (SV) and ovisperm duct (SO) are similar to these organs in pilsbryi.3 mm. C. the whole male organ resembling that of typical Helisoma (fig. and 7. pilsbryi infracarinatum F. There . especially in figs. 18). Baker). It varies from a narrow band of muscle to a wide band with several branches. 13). cylindrical vergic sac (fig. A larger form than pilsbryi is known as H. Baker which is common in southern Canada. 13). the preputium is twice as long as the vergic sac and both are wider than in tiivolvis. The seminal vesicle is shorter and wider. In a rather distinct species.

The penial gland is rather shallow (fig. The genitalia greatly resemble those of H. 8). is similar in its genitalia to Helisoma corpulentum. as in other immature forms of Pierosoma. long. Figures 16. . has the genitalia much as in the typical form. In fig. In fig. high (fig. (Pilsbry) living in Greenfield pond. the spermatheca is more nearly like that of infracarinatum. wide). 8. 13. Figure 14 shows one form of the penial complex when the j^reputium is pushed upward. the latter nearer maturitv. The gland duct is very long mm. long and is placed at the end of a very long duct (5 mm. The ovotestis (OT) occupies a trifle over one whorl and consists of about eighteen rows of long diverticula (fig. Figure 4 shows a specimen in which the vergic sac is shortened and the upper part of the preputium has been pushed upward. Two forms assumed bv the verge are shown in figs. shows no essential differences in its genitalia. The form of the penial complex in immature individuals is shown in figs. 3).5 mm.) and is is about as long as the preputium. resembles the northern pilsbryi slender. A large Pierosoma. 5. It (24 . The prostate is 5 mm.5 mm. multicostatum F. progressively from 16 to 18. the shells being of the same size and age. Baker. 0. and Subgenera is — Recent and Fossil 139 also a greater number is of diverticula. long).5 mm. pilsbri/i iniracarinatum. Figiu'e 12 also shows an immature condition. pyriform. long. 17.5 mm. 1). a related species. 1. 15) but this could be from contraction. Baker. club-shaped diverticula The spermatheca A shown in section in fig. The penial complex of an adult individual is shown in fig. wide. and 1. the penial complex of a fully mature individual is shown. Helisoma ivhiteavesi F. vermilionense F. a penial complex is shown with the preputium pushed upward. C. In corpulentum (fig. 16 and 17 but much longer in fig. the seminal vesicle (SV) is long and wide (4 mm. near Wilmington. the preputium and vergic sac being about equal in length. Baker. The vergic sac differs from that of the corpulentum group in being very long (5. Helisoma corpulentum (Say) and its races inhabit the northern part of the United States and southern Canada. C. In the penial complex (fig. 13. Compare fig. 19. 17 and 18.). A race of corpulentum.SuhjamUies. 6 being from a younger specimen. The immature penial complex (fig. apparently rare in Canada. The penial gland is twice as long cross section of the prostate seventeen rows of long.) and the duct longer (3. Figure 11 is from an immature individual in which the vergic sac has been lowered to the side of the preputium. H.7 mm. the preputium is normally very wide. The gland duct is partly uncoiled.). 19 with fig. with the vergic sac small and narrow and about half as long as the preputium. fig. the preputium pushed upward and the long penial gland duct uncoiled (it is 14 mm. Minnesota. The principal features of the genitalia are figured on plate 31. In cross section. 10) is very wide with a short duct. The normal form of the penial complex is shown in fig. near the lower end shows (fig. C. Figures 9 and 10 show the form of the penial complex in immature individuals. 12). 6 and 7. In form. North Carolina. 9. and 18 show the penial complex of immature individuals. Another race found only in Lake Vermilion. (3 mm. The penial duct is short in figs. this is seen to be composed of twenty long diverticula arranged fan-wise. c. Genera. 2). a common characteristic in Pierosoma. 18. Helisoma magnificum.). The spermatheca (S) is 2 mm.

principally bordering the gland. shorter penial gland duct. long and more than 1 mm. The penial gland duct is not very long (3 to 4 mm. long in fully adult specimens (figs. One of the most abundant and widely distributed species is Helisoma subcrenatuni (Carpenter). long and is composed of twenty. the penial gland (fig. the penial duct and complex is of a young specimen. fig. The spermathecal duct is very long (12 mm. Internally. and the presence of the aculeate vesicles in the seminal vesicle. The uterus (U) is narrow anteriorly and widens posteriorly into the nidamental gland (NG). which is a large organ 1 mm. Figure 15 pictures the whole genitalia. 22. 12). long. the complex shows a large gland and a cylindrical verge. 13). The prostate is 7 mm. In adult or old specimens. plexatmn (Ingersoll). The vergic sac is small and less than half the length of the . and the duct is 5 mm. the organs spread out. The penial complex (fig. The penial gland duct is very long. These differences are shown in figs.5 mm.four rows of diverticula. The organs of the female system are long and narrow. however. The sperm duct (SPD) is short (2 mm. 10 and 15). long. 10). long or nearly three times the length of the spermatheca. is different in the absence of the digitate vesicles and is more like the seminal vesicle in pilsbryi. The general characteristics of plexatum are like subcrenatum. wide. A Species of the Rocky IMountains and Westward Several species and races of Pierosoma occur in the western part of the United States and Canada. cross section of the penial gland is shown in fig. In vertical section (fig. 17 and 18 (compare with figs. when adult. both nearly filling their respective cavities (fig. The genitalia of subcrenatum differ from those of the trivolvis group in the greater number of prostate diverticula. It is much like that of trivolvis.) and is normally coiled on the preputium. long and is notable for the number of rather long vesicles. A relative of subcrenatum. The vergic sac is slightly shorter than the jireputium. long. 19). the genitalia of which are figured on plate 30 (specimen not quite mature). In section. the penial complex shows a large penial gland (GL). It resembles the spermatheca of trivolvis. wide. somewhat sausage-shaped verge (V). The spermatheca (S) is about 1 mm. long and the duct is long and narrow and about 2 mm. The oviduct (OD) is a narrow tube about 2 mm. The seminal vesicle (SV). 11) is similar to that organ in trivolvis. a rather thick diaphragm (D). The preputium is widely pyriform while the vergic sac is narrow and cylindrical. long. The spermatheca (S) is very narrow. The seminal vesicle is 2 mm. which is much wider than the ovisperm duct. and a thick. the penial complex is very wide and the duct is very short (fig. 15. 20) has a pyriform preputium and a cylindrical vergic sac. over 2 mm. there is more difference between the preputium and vergic sac and the duct is always longer. In fig. The prostate is about 2 mm. The prostate is 5 mm.140 The Molluscau Family Planorbidae and corpulentum. long and more than 1 mm. The penial complex is about 3. There is a short prostate duct at each end of the prostate which joins the sperm duct beneath the prostate. H. wide and high. 21). 4 (genitalia).) and narrow. In immature individuals. long. long) and the vas deferens is narrow and about 7 mm. arranged fan-wise when seen in section (fig. 17. There are thirteen rows of diverticula. is figured on plate 30. and the spermatheca is 3 mm.

4) indi- preputium cates the relationship between these organs. The ]icnial complex (fig. Specimens collected in Tooele County. in length and the duct is 3 mm. 6). 17). In vertical section (fig. The presence of the two small and narrow retractor muscles is noteworthy. wide neck. the preputium. These are much wider and less in number than in subcrenatum. long and the ovisperm duct about 3 mm. fig. There is one retractor muscle (sometimes split into two smaller muscles). 4) is round. with relatively few large vertical folds and a number of pavement-like folds in the bottom of the cup. is Helisoma horni (Tryon) from Paul Lake. the penial complex shows the penial gland to be compressed as in subcrenatum (fig. The prostate is about 3 mm. long) and bears sixteen rows of long and narrow diverticula (fig. widening as it enters the narrow vagina. In vertical section. 3) and not globular as in trivoivis. in length. Subfamilies. Oregon. in length and connects with the long and narrow vagina by a duct slightly more than 1 mm. being more cylindrical and less pearshaped and the vergic sac is relatively longer. The spermatheca (S) is bulbous. however. The vergic sac is short. 7. and Subgenera — Recent and Fossil 141 (fig. The preputium is pushed upward. 1) has a large. 3) resembling trivoivis in this respect (compare figs. and one band of supporting muscle. . long. 3). 3). wide and has thirteen rows of large. 1) has a long vergic sac enlarged at the outer end. The penial complex of a form of subcrenatum from Pass Lake. The prostate is short (about 2 mm. long. Puget Sound. about 1 mm. J. The penial gland duct is longer than in subcrenatum (figs. attached to the lower part of the preputium. and almost half the length of the preputium. The genitalia are different from both subcrenatum and ple. by Mr. The seminal vesicle is similar to that of plexatum. Utah. The seminal vesicle is about 1 mm. Fidalgo Island. oviduct and nidamental gland (fig. The shape of the spermatheca is like that of horni but the duct is shorter. The spermatheca is bulbous. 8). 5). fig. as in ple. The retractor muscles are as in subcrenatum. The verge (V) is large and long ovate in form when at rest (fig. The penial gland (fig. and attached to the wall of the preputium by a short.). 5). 2. The penial gland is shown in section in fig. The latter bears short lateral vesicles for a large part of its length. Henderson belong to this species and the anatomy of this small planorbid is now available. is figured on plate 32 (fig. On plate . 1). The gland duct is about as long as in plexatum. the principal organs of the genitalia are shown. 2. British Columbia. pyriform preputium with the lower part lengthened. A section through the sperm duct. The penial gland duct is very long (5 mm. long. 1 and 17).vatum. but believed to be of specific rank. and 1 mm. 5. 8. 2) is like that organ in trivoivis. It is similar in form to the penial complex of subcrenatum figured on plate 30 (fig. to which the gland duct is connected (DC). about 0. 28. Washington. The ]ienial gland (fig. the cup is circular. Genera. and 9. Another form related to subcrenatum. rather short. The penial complex (fig. 1. IMany years ago Tryon described a small Planorbis from Pueblo Valley. The shapes assumed during extension into the vergic sac cavity are shown in figs. calling it Planorbis oregonensis (now Helisoma oregonense) This species has been one of the rarities of western Planorbidae.7 mm. swollen at the end.ratum (plate 26.. club-shaped diverticula (fig.

narrow. long and 0. The sperm duct is 6 mm. The ovotestis is made up of very large. It is about 9 mm. 6. The preputium is pyriform and very wide at the summit. The spcrmatheca (S) is of unusual length (3. widelyspaced vesicles on the lateral margin. by many small. These are probably part of the seminal vesicle system. Baker. It is 3 mm. The sperm duct (SPD) is 3 mm.). The prostate is short 1.). long. 1) and is coiled in much the same manner.3 mm. The Prospect Park specimens are very large. narrow vagina (VG). has been named H. club-shaped diverticula in eleven rows. There is one large retractor muscle and one or two supporting muscles. The spermatheca (S) is pyriform. the prostate in section). the nidamental gland (NG) is composed of several large diverticula.). wide) There is a short prostate duct branch at each end of the sperm duct beneath the prostate.). long. 1. The prostate (PRS) is verv long (6. Its duct is relatively short (5 mm. In section (fig. It swells in the center and tapers at each end. The genitalia of calij orniense are peculiar in the great length and size of the seminal vesicle.) and about one-fourth as wide as long (1.) which enlarges at its point of entrance into the short.5 mm. long. The ovotestis (OT) consists of large. is club-shaped and is placed at the end of a very long duct 10 mm. California. long. There is a wide. 2. wide for two-thirds of its length and tapers to about half this diameter near the point of contact with the prostate duct.5 mm. club-shaped diverticula (fig. The genitalia of this species are shown in fig. very muscular diaphragm.142 The Molluscau Family Plauorbidae A Helisoma abundantly distributed in California.) as in the Shandon specimens. The penial complex (fig. In the section. the vergic sac placed at the side. long. 5) is short and cup-shaped with many vertical folds. the prostate shows fourteen rows of diverticula of varying length. 12).5 mm.) and has twelve rows of long.5 mm. long and a cylindrical vergic sac 1.5 mm. Figure 2 shows the penial complex with the preputium pushed upward. 3) has a large. long) and wide 1 mm. The genitalia are shown on plate 32. The verge (fig. and in the sessile penial gland. in length and is not as pyriform in shape as in the other Californian sjiecimens examined. rounded vesicles. ( .) two-thirds of which bears small. fig. The vas deferens is a long. clubshaped diverticula of varying length. long. with a long.5 mm. tenue calij orniense by F. The seminal vesicle is shaped like that organ in calif orniense (see fig. The ovisperm duct is 5.2 mm. The vas deferens VD) is a long. on one side only. and for two-thirds of its length is bordered. long (when extended) and 1 mm. long). unbranched. near Redlands. wide and is composed of many small vesicles. enlarged as it enters the long and narrow vagina (VG). The penial gland (fig. narrow duct about 2.5 mm. narrow tube (25 mm. C. The gland duct is over twice as long (14 mm.5 mm. One of the largest of west American planorbids is Helisoma traskii (Lea). narrow tube 11 mm. The ovisperm duct (SO) is very long (16 mm. thirteen rows being counted in a ( ( ( . 1. The verge is 3 mm. It is made up of many small vesicles. long and is very narrow. The seminal vesicle (SV) is wide and long (5 mm. 17 on plate 32. and believed to be a race of the Mexican Helisoma tenue (Phil. pyriform preputium 2. in the very short prostate. which probably accounts for the great length of the duct. V) is wide above but tapers to a narrow point below. Figure 7 shows the penial complex of a specimen from Prospect Park.

wall of the preputium by a thick neck or base. parallel to the length of the kidney. 13). 9. fig. 2) is elongated. long and is directed backward. The kidney of Helisoma trivolvis (plate 44. 3. The penial complex is like that of californiense figured on plate 32. plate 25. The albumen gland AL) is squarish and is made up of small follicles. long and 2 mm. plate 26. fig. coiled in the vergic sac. fig. the material had been preserved in formalin and the anatomy could not be worked out in a satisfactory manner. long) and is a narrow tube.Subfamilies. 10. 16). 11). fig. There is a conspicuous crest or ridge extending down the center of the kidney from just below the pericardium to within 1 mm. Other details are not available from these specimens. One specimen of fraskii had a bifid tentacle (fig. truncatum. The shape of the pseudobranch does not vary to any degree in the different species and races of Pierosoma. subcrcnatinn. plate 32. 11. living characteristically in Klamath Lake. Another large species found in California is Helisoma occidentale (Cooper). Unfortunately. 14) has a pyriform i)rei)utium 4 mm. Sectioned vertically (fig. 17. 16. 11. californiense. The greater number of specimens examined had the penial complex as shown in fig. There is a large pneumostome (PS). 12) is very wide. The base of the cup has many folds to which the penial gland duct is attached (DC). The anal opening is to the left of the ridge. The shorter penial gland duct allies traskii with californiense and separates it from plexatum. united at their attachment near the upper part of the preputium. somewhat bilobed. fig. The pseudobranch of Helisoma trivolvis (plate 24. and Subgenera — Recent and Fossil 143 section at the proximal end of the ovotestis (see fig. 4. The retractor muscle (RM) is wide and is divided into two main branches. long (PR. Respiratory and Renal Systems. with a narrow crest or ridge extending down the rectum (R) and over the pseudobranch at about the center. The penial complex (fig. fig. Genera. jilate 32. the preputium pushed upward and the vergic sac attached to the side about halfway between the extremities of the penial complex. infracarinatum. The penial gland was about as in traskii (fig. Internally (fig. the preputium has a large penial gland (GL) which is attached to the. Several of these are figured on the following plates: pilsbryi. excepting in the general shape which in many cases is broadly rounded below and not bilobed (see plate 25. fig. fig. and in the generally long and narrow uterus and vas deferens. however. fig. 12. long and a sausage-shaped vergic sac 3 mm. It is different from californiense in the size of the prostate and in the size and length of the spermathecal duct. of the lower . is shown in fig. The genitalia of Helisonia trnskii are peculiar in the large prostate. traskii. VS). 11). Oregon. The ureter is 1 mm. the elongated spermatheca on an unusually long duct. The penial gland duct (DC) is of medium length (9 mm. 13) the penial gland is seen to have a large cup bordered by large vertical folds (GF).5 mm. 15. The vergic sac. The verge. plate 30. OTj. fig. wide at the upper part near the pericardium. plate 25. The supl)orting muscle (SAD is rather narrow and is branched several times in its attachment to the upper jiart of the preputium on the opj^osite side from ( the retractor muscle. is more elongated at the distal end. 3.

1. 19 (note difference in two races). and runs forward to the columella muscle. fig. . 3) is somewhat shortened and bent toward the pericardium. In fig. a section near the lower end shows a large. In macrostomum. 11). near middle. 13. calif orniense. The ridge is broad at the upper part and then narrowed for the greater part of its length. are: pilsbryi. In the race lentum (plate 44. long. fig. figs. at B in fig. fig. multicostatum. near middle. plexatuni. near middle. . much-elongated lumen with the veins at the lower lateral corners (fig. The size of the organs differs in the various species. near middle. near middle. section near lower end. 21 and 22 and note difference in ridges). Digestive System. The intestine is 33 mm. fig. pointed ridge. near lower end. Those figured fig. fig. the stomach is seen from below showing the position and form of the blind sac or caecum (BS) which is attached to the pyloris just below the opening of the liA^er (OLV). near middle. fig. section near middle. 9) shows the ridge to be rather short and narrow and placed at about the middle of the kidney. (compare figs. is larger in diameter than the intestine.2) is cliaracteristic of that of all species of Pierosoma. horni. 6. A cross section is shown in fig. The species and races of Pierosoma show considerable variation in the form of the lumen. 2. near middle. fig. It is about 10 mm. The ridge is approximately like that feature in trivolvis. and is particularly characterized by the presence of a frilled crest or ridge (RD) which extends the whole length of the rectum and ends over the pseudobranch. with the veins at the middle on each side. truncatum. The digestive system of Helisoma trivoh'is (plate 48. fig. oregonense. the largest race of trivolvis. and especially in the form and position of the ridge. A cross section made through the upper part (fig. increasing several times in diameter as it passes to the crop portion of the stomach. The intestine (IN) makes a complete turn over the oesophagus. fig. 4). The kidney of calif orniense is figured on plate 45 (fig. subcrenatimi.144 The Molluscan Family Planorhidae end of the organ. traskii. near middle. at A in fig. 16. figs. The stomach is composed of a short crop (CR) a bilobed gizzard (GZ) and an elongated pyloris (PY). The oesophagus (OE) is a narrow tube as it leaves the buccal sac. which is found in all species of Helisoma and its subgenera. section near lower end. 22. ovate lumen with the veins at the lower corners. long from the end of the pyloris to the rectum. 10). long. The whole kidney is much larger and much thicker with a relatively small ridge. The rectum is 10 mm. infracarinatum. 24. and a high. 20. fig. This blind sac is present in all species of Pierosoma. fig. . performs another loop. near middle. 15. fig. 17. A cross section of the kidney made at about the middle portion (plate 44. fig. Several of these are figured on plate 44 and need no special comment. 18. fig. The lumen is rounded and the veins at each end are large and placed at the center of the margin of the lumen. 14. 4) shows the roundly-ovate lumen occupying the greater part of the kidney. occidentale. 12. 2 and 9. 23. near middle. 21 on plate 44. fig. It is relatively short and variable in width. 21. A section made near the lower end of the kidney (fig. depressum. 4) shows a very wide and flattened lumen with the veins at each end. extends backward around the liver. 15. Compare with fig. Compare this with the kidney of trivolvis. 5. magnificum. The kidney of corp^dentum (fig. a section made near the lower end shows a small. 6. corpulentum.

In chantaiiquerise (plate 55. 3. sharp cusps and the ectocone region developing two or three cusps above the ectocone on the outer margin of the tooth. 6. that the more elongated aspect may be due to distortion in the animal when killed. fig. The radula of the type species. There is some variation from the type described above. 3 mm. 2. The radula of Pierosoma is similar to that of Seniinolina in the bluntly rounded mesocone of the lateral teeth. fig. plexatum. corpulentum. long) on each side. fig. and subcrenatum (plate 70. some of the lateral teeth developing two cusps in place of the one ectocone. the entocone breaking up into three small. 9. fig. and Subgenera — Recent and Fossil 145 high. There is not much variation in the form of the buccal sac in Pierosoma. 12. and with slight denticulations. In winslowi (plate 56. The jaw in Pierosoma is always of three parts. the whole sac is a trifle more elongated. 5). 1). Helisoma trivolvis. plate 50. wide and low. their posterior ends connected. 2. the entocone and ectocone sharp and spade-shaped. RL) which has to do with the formation of the radula membrane. tenue californiense. There are two strong buccal retractor muscles (BR). outer margin of the tooth.5 mm. of the races of trivolvis differ little from the type species. corpulentum. The face of the superior jaw is marked by fine striations which form a more or less denticulate border to the jaw when seen with high magnification. several of these arc shown. The salivary glands form a loop. 7. fig. is shown on plate 55. The jaw of Helisoma trivolvis is shown on plate 49. fig. The salivary glands (SG) are two in number. macrostomum. There are two long and narrow lateral jaws one on each side bordering the lips of the animal. sometimes somewhat arcuated. The lateral teeth (1-10) have a quadrate reflection which is tricuspid. There is reason to believe. The jaws of some other species are figured as follows: truncatum. 12). 2. plate 49. oregonense. the lateral teeth (1-11) are narrower than in trivolvis. pilsbryi. 15. 13. and this is characteristic of all members of the subgenus. long. The center tooth has two spade-shaped cusps which do not reach the lower margin of the base of attachment. fig. claw-like. magnificujji. fig. occidentale. fig. At the back end of all sacs there is a rounded or elongated radula sac (plate 70. Radula. 3.Subfamilies. In some species. 6. fig. very narrow. the The radulae . 5. 1. fig. fig. fig. The superior jaw is oblong. with the entocone broken up into four to five small cusps and the ectocone into four or more small cusps on the upper. which enter the buccal cavity near the oesophagus by a long. This varies in form in some of the species. The extreme outer marginal teeth are small. 7) and pseudotrivolvis (=lentum) figs. as pilsbryi. fig. narrow duct (1 mm. especially in the fore part. 16. The intermediate teeth (11-13) have the reflection somewhat narrower and more elongated than the lateral teeth. 10. is shown on plate 56. The radula of the large race. traskii. fig. 4. 2 mm. as shown in fig. fig. On plate 55. fig. the mesocone rounded. 8. infracarinatum. 13. californiense. The buccal sac (BP) is a pear-shaped organ. and usually seen in the form of a question mark. horni. Marginal teeth (14-18) still narrower. lentum (figs. long and 1. 1. subcrenatum. fig. however. fig. Genera. The oesophagus leaves the oral cavity near the upper posterior por- tion of the pouch.

fig. Idaho. 4. the teeth are arranged in a similar manner to those of trivolvis (plate 57. 2. In whiteavesi (plate 58. 5) carinatum (plate 56. fig. were normal. Another form from Lost Lake. In the eighty-second row of another sjiecimen. figs. 6). There are eight subquadrate lateral teeth and four intermediate teeth. 21-26). fig. this feature was well marked The (% 4). 2. fig. 5-10). the early laterals are quadrate (fig. 2. the intermediate teeth are narrow and the entocone and ectocone are not broken into smaller cusps (10-14). the teeth are similar in form to those of the corpulentum group. 3) are similar in form. figs. large traskii from Kern Lake. In pilsbryi figs. Colorado. 4. fig. somewhat obliquely formed mesocone on the lateral teeth. lateral teeth are also narrower. similar to those of trivolvis (plate 59. fig. 4). The marginal teeth are normal as to cuspidation. the ectocone is placed high up on the margin. 3). fig. Other individuals were normal (fig. the center tooth of the eighty-eighth row had seven small cusps and the fifth lateral had two sharp entoconic cusps (plate 60. and some of the marginals. fig. however. 4-6). The marginal teeth are also narrow and the cuspidation is different from that in trivolvis and other species of Pierosoma (fig. long and narrow. near Gypsum. 2). Washington. The marginal teeth (11-22) arc wide and the denticulation is very irregular. plate 57. Helisoma plexatum has narrower lateral teeth than trivolvis (plate 59. Washington. Puget Sound. 7). and the extra cusps of the entocone appear later the series (plate 57. figs. 2). with teeth m quadrate laterals (plate 60. . the last intermediate tooth and the first marginal tooth were peculiar in this respect (fig. horni. had very wide. A lateral and two intermediate teeth of a specimen from Smartweed Lake. 13. An allied species. Specimens from Skagit County. 2). 27-41). 5). In specimens from another locality. which are not as quadrate as in subcrenatum (1-9). California. many of the laterals. the intermediate teeth are very 1. small interstitial cusps appearing among the larger cusps in some teeth (as 11. The lateral teeth (1-8) are longer than wide and are unmodified tricuspid.146 The Molluscan Family Planorhidae (plate 56. In one specimen from this locality. 14. 2) the laterals are narrower (as in chautauquense and winslowi) In several laterals of truncatum. 3) and injramore quadrate. A form of subcrenatum from Pass Lake. fig. is peculiar in several respects (plate 60. has quadrate teeth. 1. there was observed to be considerable splitting of the entocone and ectocone cusps (see fig. In another row (eightieth). The marginal teeth (15-32) are less cusjiidate than similar teeth in subcrenatum. the two large cusps extend a considerable distance below the lower margin of the base of attachment. In the center tooth. In the western group of Pierosoma. The races vermilionense and multicostatum (plate 58. 6 on plate 58. in one specimen had the mesocone broken and wavy and otherwise modified (plate 58. low lateral teeth (plate 60. 1). In the large magnificum. In corpulentum. Colorado. fig. 1). fig. 1). A nearly complete row (the ninetieth) is shown in fig. Puget Sound. Typical subcrenatu)n from Cottonwood Pass. The outer marginals (32) are small and more or less vestigial (see plate 60. has a large. 5). 17). fig. but the marginals show some variation. 4. are shown in fig. the laterals are . is shown on plate 61. 5. In truncatum (plate 57. The intermediate teeth (9-10) have the entocone and ectocone split into small cusps. In specimens of plexatum from Rexburg. similar to those of trii'olvis.

SubfaffiUies, Genera,

and Subgenera

— Recent and Fossil


In another large species of Pierosoma from Klamatli Lake, Oregon (occidcntalc, plate 61, fig. 1), the teeth are more uniform with but little splitting of the cusps. The radula differences between the two species occidentale and traskii are very marked, as shown on plates 60 and 61. In specimens of tenue calif orniense from Shandon, California (plate 63, fig. 2) the center tooth has two wide, spade-shaped cusps- extending below the base of attachment. The laterals (1-11) are at first wider than high, the cusps rather short, but toward the marginal teeth they become narrower. The entocone and the ectocone tend to split in some teeth (as 1, 3, 6, 7). There is one narrow intermediate tooth (12). The marginal teeth (13-19) are narrow, the entocones 4-6 cuspidate. In the 100th row of

the same membrane, the entocone and ectocone of the lateral teeth were largely modified by splitting (see fig. 2). In a radula from a specimen living in the San Bernardino Mountains, there were eleven laterals, one intermediate tooth and nine or more marginals. No splitting was observed in this radula but in another membrane all of the laterals were abnormal (plate 59, fig. 4) in the sixtieth row, small cusps appearing irregularly on the entoconic region. The entocone of the first lateral was broken up into four small cusps. The intermediate and marginal teeth were narrow and many of them abnormal in the splitting of the entocone (as 14, 16, 17). The mesocone in all teeth was more or less irregular. In fig. 6, a very abnoniial eleventh tooth from another specimen is figured. The relatively small species oregonense (plate 61, fig. 3) has narrow laterals (1-8) and intermediate teeth (9-10). The marginals are normal for the group. The radula formulae of the different species vary more or less and the individual radulae may differ in number of teeth in a row. For comparison all of the species of Pierosoma which have been examined, together with all of the localities represented, are shown in the following table. This also indicates the material studied and the source from which the material was obtained. ]\Iore specimens were dissected for the genitalia than for the


From one
23-1-23 23-1-23 23-1-23 23-1-23 23-1-23 23-1-23

to seven specimens of each
to to to to to to

were examined.

26-1-26 24-1-24 24-1-24 25-1-25 24-1-24 24-1-24

Canandaigua L., New York Green Lake, Wisconsin
Devils Lake, Wisconsin Murphy Creek, Wisconsin

trivolvis trivolvis

Sturgeon Bay, Wisconsin


Joseph, Illinois


pseudotrivolvis pseudotrivolvis pseudotrivolvis pseudotrivolvis pseudotrivolvis


macrostotnum viacrostomum

macrostomum macrostnmum
chautnuquense truncatum winslowi

24-1-24 22-1-22 23-1-23 24-1-24 23-1-23 26-1-26 26-1-26 27-1-27 27-1-27 31-1-31 37-1-37 24-1-24 20-1-20 27-1-27

to 25-1-25 to 24-1-24 to 24-1-24 to to to to to to to to to

Peoria, Illinois

Reelfoot L., Tennessee

Mammoth Spgs.,


Ada Lake, Oklahoma
25-1-25 28-1-28 28-1-2S 30-1-30 32-1-32 39-1-39 26-1-26 21-1-21 29-1-29 Trinity R., Texas New Orleans, Louisiana Near Boston, Massachusetts
Bayfield, Wisconsin

Moose Ear Creek, Wisconsin Saskatoon, Canada
L. Grauveau,



New York

Winnel)ago L., Wi-sconsin Arbor Vitae L., Wisconsin

pilsbryi pilsbryi filsbryi itifnicdriiitdum injrdciin nittam

The Molluscan Family Plaiiorbidae
42-1-42 27-1-27 27-1-27 28-1-28 28-1-28 32-1-32 31-1-31 25-1-25 30-1-30 32-1-32 36-1-36 25-1-25 32-1-32 40-1-40
to to to to

43-1-43 29-1-29 29-1-29 29-1-29


infracarinatum infracarinatum corpulentum corpulentum corpulentum multicostatum
viulticoi^tntum rcrmilidiicnse

to 37- 1-37 to 38- 1-38 to27- 1-27 to37- 1-37 to 35- 1-35 to 41- 1-41 to 28- 1-28 to 35- 1-35 to45- 1-45
to to to to to to to


subcrenatum subcrenatum

38-1-38 30-1-30 30-1-30 25-1-25 36-1-36 25-1-25 29-1-29

41-1-41 37-1-37 31-1-31 27-1-27 42-1-42 27-1-27 32-1-32

Wilmington, North Carolina Chetek L., Wisconsin Prairie L., Wisconsin Chetek L., Wisconsin Moose Ear Creek, Wisconsin Basswood R., Ontario Bimaji L., Ontario As above (immature) Mille Lacs, Ontario Rideau R., Ontario Lake La Croix, Ontario As above (immature) Knife Lake, Minnesota Kahnipiminanikok L., Minnesota Mille Lacs swamp, Ontario As above (immature) Vermilion L., Minnesota As above (immature) Mille Lacs, Ontario Gypsum, Colorado Pass Lake, Washington
Lost Lake, Washington
Ponchlla L., Washington

180 180-185 205 185 185 140-163 140-151 140-153 140-155 171-177 165-190 109-117 140-150 163-170
145-150 152 120-123 119 139-154 123-130 150-160

P. F. F. F. F.


C. Baker C. Baker C. Baker C. Baker A. R. Cahn A. R. Cahn A. R. Cahn A. R. Cahn A. LaRocque A. R. Cahn A. R. Cahn A. R. Cahn A. R. Cahn

A. R. Cahn A. R. Cahn F. C. Baker F. C. Baker A. R. Cahn
J. J.

Henderson Henderson Henderson


28-1-28 to 29-1-29 28-1-28 to 31-1-31





subcrenatum plexatum plexatum plexatum plexatum plexatum
horni oregonense
occidentale traskii californiense


30-1-30 26-1-26 25-1-25 26-1-26 27-1-27 27-1-27 32-1-32 20-1-20 31-1-31 28-1-28 24-1-24 26-1-26 24-1-24

to to to to to to to to to to to to

32-1-32 30-1-30 30-1-30 30-1-30 28-1-28 31-1-31
21-1-21 32-1-32 30-1-30 25-1-25 27-1-27 27-1-27

McMurrav, Washington Columbine L., Colorado Smartweed L., Colorado Grand Mesa, Colorado Gocseberrv Creek, Utah Rexburg, Idaho

150-170 140-173 155-170 155 150-165 145-155 Paul Lake, British Columl)ia 110-114 111-117 Tooele Co., Utah 150-160 Klamath L., Oregon 161-182 Kern Lake, California 155-160 Shandon, California 150 San Juan R., California 140-157 Near Redlands, California

J.Henderson J. Henderson J.Henderson J. Henderson J. Henderson J. Henderson


J.Henderson J.Henderson
S. Siegfus

Hanna & Church D. Hanna
S. S.



Showing Range of Formulae

Different Species







20 to 21 truncatum

28 to 32 subcrenatum var.
occidentale traskii




lentum subcrenatum plexatum

infracarinatum corpulentum multicostatum


This table shows that, while there is wide variation in the formulae of the radulae in Pierosoma, the range within the species is fairly constant. The difference in many membranes is due to the presence of several extra marginal teeth on some radulae which do not appear on others. It is to be observed, also, that the immature animal usually has a smaller formula than the same species when adult. This is noted in the table under several species {infracarinatum, corpulentum, vermilionense, and multicostatum). Magnificum has the largest formula, 42-1-42 to 43-1-43.

Geographical Distribution. The subgenus Pierosoma is distributed from Ungava, Labrador, Mackenzie, and Alaska southward to Georgia, Louisiana, Texas, and California, the whole of Mexico, and southward, on the west coast of South America, to Peru. In the West Indies, it is known

Subfamilies, Genera, and Subgenera

— Recent and Fossil


from Cuba, Jamaica, Haiti, and other islands. The metropohs of the group in North America, principally in southern Canada and northern United States, where the largest number of species and races is found. Southward the number of species rapidly decreases, only two species being known from South America and four from the West Indies. These appear to be species which have migrated southward in past geological time, possibly by the

aid of migrating birds. In the recent fauna, no member of the Helisoma group is found outside of North and South America. In Pleistocene deposits, however, a form of Helisoma allied to trivolvis has been reported by Lindholm (1933) from the Kolyma River in northeast Siberia beyond the Arctic Circle. It would appear that this planorbid is not referable to trivolvis but to some form like horni or subcrenatum. The latter species is common in northwestern British America and also occurs in Alaska. The Siberian form should probably be allied with subcrenatum. Lindholm called it Helisoma trivolvis koly7nense but it might be better to designate it simply as Helisoma kolymense Lindholm until more is known about its affinities. As suggested by Pilsbry (1933, p. 137), it might represent a migration by the way of the

old land bridge between Alaska and Siberia, used mammals during Pliocene and Pleistocene times.
to the

by both man and other

Species Considered as Valid. The large number of species attributable genus Helisoma, subgenus Pierosoma, are listed below. The names Helisoma (Pierosoma) are understood to be prefixed before each specific


in the


(Gould) binncyi (Tryon)

plexatmn (Ingersoll) salvini (Clessin)



caribaeum caribaeum (d'Orbigny) caribaeum guatemalense (Clessin) chnutauquensc F. C. Baker
contrcrasi (Pilsbry)

subcrenatum subcrenatum (Carpenter) subcrenatum disjectum (Cooper)
tenue tenue tenue tenue tenue tenue
tenue (Philippi) boucardi (Fischer and Crosse) calij orniense F. C. Baker chapalense (Pilsbry) exaggeratum (Martens) pertenue F. C. Baker (= applanatus Martens preoccupied) tenue sinuosum (Bonnet) tenue strebeUanum (Fisch. and Crosse) traskii (Lea) trivolvis trivolvis (Say)
trivolvis fallax (Haldeman) trivolvis lentum (Say) trivolvis macrostomiim (Whiteaves) trivolvis turgiduni (Jeffreys)

corpulentum corpidentum (Say) corpulentum vermilionense F. C. Baker
costaricense (Preston)

equatorium (Cousin) eyerda??ii Clench and Aguayo
jovenle (Menke) horni (Tryon)

{^ affine

C. B.


kolymense Lindholm
viagnificum (Pilsbry)

mullicostatum multicostatum F. C. Baker multicostatum whiteavesi F. C. Baker occidentale occidentale (Cooper) occidentale depressum F. C. Baker oregonense (Tryon)


intertextum (Sowb.)


peruvianum (Bred.)
pilsbryi pilsbryi (F. C. Baker) pilsbryi injracarinatum F. C. Baker

truncatum (Miles) tumens (Carpenter) winslowi (F. C. Baker)


Remarks. Pierosoma is a very distinct group of Helisoma, distinguished from the subgenera Helisoma and Seminolina by peculiarities of genitalia and radula. The duct of the penial gland is always longer in adult animals
than in the other groups mentioned. Some of the special anatomical features have been described and figured in the preceding pages. The shell is large

and distinctly sinistral, especially in immature specimens. Pierosoma dominant planorbid group in North America north of ]\Iexico.




The Molluscan Family Planorbidae
Type by


Haldeman, 1842
campanulatus Say

original designation Plnnorbis

1842. Planorbella 1861.

Haldeman, Mon. FW. Univ.

1865. 1869.

campanulatus Sa}' (plate 1, figs. Adula H. Adams, Proc. Zool. Soc, p. 145. Type Planorbis midtivolvls Case. Non Adula H. and A. Adams, 1857 Planorbella Binxey, L. FW. Sh. N. A., II, p. 109. Type Planorbis campanulatus Say. As subgenus Adula Binney, op. cit., p. 111. Type Planorbis midtivolvis Case. As subgenus Ancaeus H. Adams, Proc. Zool. Soc., p. 275. Non Ancacus Fauvel, 1863. In place
of Adula,

Moll., Physidae, p. 14. 7-11, 1844)

Type Planorbis

preoccupied Dall, Ann. Lye. Nat. Hist. N. Y., IX, p. 351. Tj'pe Planorbis campanulatus Say. As subgenus 1870. Adula Dall, op. cit., p. 351. Type Planorbis multivolvis Case. As subgenus 1870. Planorbella Tryon, Contr. Hald. Mon., pp. 188, 196. Type Planorbis campanulatus Sa.y. As subgenus
1870. Planorbella

Adula Tryon, op. genus


pp. 188, 197.

Type Planorbis

multivolvis Case.

As subSay.

1883. Planorbella Fischer, Man. de Conch., p. 509. As section of Planorbis 1883.

Type Planorbis campanulatus
multivolvis Case.

Adula Fischer, Man. de Conch.,
section of Planorbis

p. 509.

Type Planorbis


1884. Planorbella

1884. 1886.

Tryon, S. and S. Conch., Ill, p. 106. Type Planorbis campanulatus As subgenus Adula Tryon, op cit., p. 106. Type Planorbis multivolvis Case. As subgenus. Adula Clessin, Conch. Cab., XVII, p. 33. Tyjie Planorbis multivolvis Case. As

1886. Planorbella

Clessin, op.




Type Planorbis campanulatus Say. As
Type Planorbis campanu-

1902. Planorbella F. C. Baker, Moll. Chi. Area, II, p. 294. latus Say. As subgenus 1905. I'lanorbella

Dall, Alaska Moll., pp.

81, 85.

Type Planorbis campanulatus Say.


As section of subgenus Helisoma Planorbella Hannibal, Proc. Mai. campamdatus Say. As subgenus


London, X,


Type Planorbis

1918. Planorbella W.\lker, Miscel.

Pub. Mus.

Univ. Mich., No.

pp. 12, 94.


Type Planorbis campanulatus Say. As subgenus Planorbella Germ.\in, Rec. Ind. Mus., XXI, pp. panulatus Sa3\ As subgenus


Type Phniorbis camp. 202.

1926. Planorbella F. C.

Baker, Trans. Wis. Acad. Sci. Arts, norbis campanulatus Say. As subgenus


Type Pla-

Hypsogyra Lindholm, Trav. Com. I'Etude Lac Baikal, for Adula H. Adams, 1861, preoccupied



New name

1928. Planorbella

F. C. Baker, Fresh-water Moll. Wis., I, p. 345. Type Planorbis campanulatus Say. Radula and genitalia. As subgenus 1931. Planorbella F. C. Baker, Proc. Zool. Soc, p. 586. Type Planorbis campanulatus Say. Genitalia and radula. As subgenus 1931. Adula F. C. Baker, op. cit., p. 566. Type Planorbis nudtivolvis Case. As syn-


of Planorbella

Hypsogyra Thiele, Hantlbuch, Teil 2, p. As section of Planorbis Planorbella Thiele, Handbuch, Teil 2, Sa.y. As section of Planorbis

Type Planorbis

multivolvis Case.



Type Planorbis campanulatus

Shell. Sinistral, of medium size, resembling Pierosoma in general form but the last whorl notably restricted behind the aj^erture which is campan-

(plate 80,

Genera, and Suhgenera










19-21, multivolvis


Animal (plate 70, fig. 1, campanulatum) With short, wide foot, oval in shape, otherwise like Pierosoma. Color dark reddish brown or blackish, bottom of foot very dark reddish with flecks of white, tentacles dark colored with indistinct white rings. There is some variation in the shades of color. The animal is usually very slow in movement.

27 {Hclisotna

campanulatum (Say)


GENITALIA. Male Organs


The seminal

vesicle (SV)



and wide (about 2 mm. long and 0.5 mm. wide) and the vesicles are very large and comparatively few in number. They are mostly attached laterally. A few short \'esicles extend down the ovisperm duct for about 1 mm. The sperm duct (SPD) is about 4 mm. long and 0.3 mm. wide. The prostate (PRS) is long (4.5 mm.) with a short prostate duct at each end which connects with the sperm duct and the vas deferens. There are fourteen rows of diverticula of varying length, the median ones the largest, the outer ones the smallest. Some of the diverticula arc branched toward the end, the section figured (fig. 12) showing four such branches. The section also shows the fan-like pattern of the diverticula. The vas deferens (VD) is a long and narrow duct (16 mm. long). The penial complex (fig. 11) is much like that of Helisoma trivolvis.

The i^reputium is distinctly pear-shaped, 1.5 mm. long, the vergic sac (VS) smaller, shorter (1.5 mm. long) and club-shaped, and very narrow at its
junction with the preputium. The penial gland duct (DC) is very long (5.5 mm.) and narrows considerably at each end where it enters the preputium and the vergic sac. Normally, the long penial gland duct is coiled on the preputium between this organ and the vergic sac, as shown in fig. 9, DC. Usually, there is one more or less branched retractor muscle (R^l) but in some specimens two retractors were observed (fig. 9). There are
several (1-3) narrow supporting muscles (figs. 5-8, 10, 11) or there may be one wide band with several smaller muscles. In fig. 9, two small muscles are shown below the retractor muscles. These are attached to the wall of the mantle cavity and not to the columella muscle. Internally (fig. 10), the preputium has several vertical folds or pilasters and a muscular, cup-shaped penial gland (GL) which is attached by a

small neck to the wall of the upper third of the preputium. In section (fig. 1), the gland is cup-shaped, one side of the cup being thicker than the other as shown in the figure (BAV). There are many vertical, thickwalled folds (OF) and a number of small, flat, pavement-like folds in the bottom of the cup from which the gland duct (DC) proceeds to the exterior of the preputium. The space between the preputium and the vergic sac is almost closed by a thick muscular ring or diaphragm (fig. 3, D) which is dependent between the walls of the preputium. The penial gland duct enters the vergic sac immediately above this diaphragm (DY). The verge (fig. 10, V) is club-shaped, tapering rapidly toward the end. In figs. 5 to 7, the penial complex of H. campanulatum s))iithi is shown indicating how the preputium becomes pushed upward and the vergic sac


The MoUuscan Family Planorbidae

appears to descend to the middle of the preputium. In fig. 8, the penial complex of H. campanulatum canadense is shown with the preputium notably pushed upward.
(fig. 15). The spermatheca (S) is very long (2.5 mm.), and tapers to the long (5.5 mm.), narrow spermathecal duct (SD). The vagina (VG) is 1 mm. long and is very narrow. The uterus (U) is long (7.5 mm.) and narrow. The nidamental gland (NG) is about 4 mm. long, very wide, and sac-like. The free oviduct (OD) is about 2 mm. long and is about twice as wide as the sperm duct. There is a large, elongated carrefour (CF). The albumen gland (fig. 2) is ovately rounded and is composed of many small follicles. The space for the passage of the intes-

Female Organs





(fig. 2,


Hermaphrodite Organs (fig. 15). The ovotestis (OT) consists of eight or more rows of club-shaiied diverticula, some of which may branch two or three times, as shown in the section fig. 4 (FOV), this branching rendering the counting of rows difficult. The figure shows fourteen rows on the surface and but eight entering the ovisperm duct. The free portion of
the ovisperm duct (SO) is 3 mm. long near the oviduct but only 1 mm. long near the ovotestis. The only previous description and figures of the genitalia of Helisoma camqiamdatum are by F. C. Baker 1931, p. 585, plate iv) where the penial complex is shown (plate 27. figs. 10-11 of this work).

Respiratory and Renal Systems. The pseudobranch (plate 27,



leaf-shaped, broadly ovate, with a ridge down the center which is a continuation of the ridge on the rectum (R). The pneumostome is large. The kidney (plate 45) is similar to that of Pierosoma as represented by Helisoma trivolvis. Sections through the kidney of H. campamdatum irisconsinense are shown in figs. 15, 16, and 17. In fig. 15, the section through the upper part shows a large rounded lumen with a large vein on each side near the middle. The ridge does not begin until lower down on the kidne3^ Figure 16 shows a section through the middle of the kidney. The lumen is larger than in fig. 15, the veins are at the lower part on each side, and the ridge is just beginning to show. In fig. 17, taken through the lower part of the kidney, the lumen is smaller, the veins relatively larger, and the ridge forms a high crest. All varieties of campamdatum show essentially this form of kidney.

Digestive System. The general digestive system does not differ from that of Helisoma trivolvis. The buccal sac is much elongated, differing in this respect from many of the species of Pierosoma. The jaw (plate 49). The superior jaw (of smithi, fig. 11) is elongated and low, somewhat arched, its face covered with vertical striations as shown in fig. 20. The lateral jaws are about as in Pierosoma. The radula (plate 62). Typical campamdatum (fig. 1) has a w'ide, bicuspid center tooth, the cusps not reaching the lower margin of the base of attachment. The lateral teeth (1-7) are squarish, tricuspid, with the mesocone bluntly rounded, the entocone and ectocone aculeate. The intermediate teeth (8-9) have smaller, sharper mesocones and the entocone and ectocone are modified by the addition of smaller cusps. The marginal teeth (10-13) are long and narrow, somewhat claw-like, with the entocone wide and armed with from four to eight small cusps. The mesocone is small




The MoUuscan Family Plauorbidae
in place of the

preoccupied names Adula and Ancaeus. Should anatomical investigation, to have characteristics different from Planorhella, it should bear Lindholm's name. It is very rare and the animal is wholly unknown. It seems, from the features of the shell, perfectly safe to merge the group with Planorhella (see plate 80,
this species prove, after


Type by
1863. Carinifcx


G. Binney, 1863

original designation Planorbis newberryi
of Shells.

Binney, Smith. Check-List

Type Planorbis newberryi Lea. Type Planorbis

As genus Megasystropha Lea, Proc. Acad. Nat.
newberryi Lea. As genus

Sci. Phil., p. 5 (footnote).

1865. Carinijex




Binney, L. and FW. Sh. N. A., II, p. 74. Type Carnijex newberryi As genus Carinifex, Amer. Jour. Conch., I, p. 50, plate vii, figs. 6-7 Megasystropha Tryon, Con. Hald. Mon., p. 84 Carinifex Tryon, Con. Hald. Mon., pp. 187, 214, Type Carinifex newberryi (Lea). As genus Carinifex Dall, Ann. N. Y. Lye. Xat. Hist., IX, pp. 344, 353. Type Planorbis newberryi Lea. As genus

1883. Carinifex Fischer, Man. de Conch., p. 508. As subgenus of Choanomphahis 1884. Carinifex

Type Carinifex newberryi


1923. 1923. 1924. 1924. 1925.

Tryon, S. and S. Conch., Ill, p. 105. Type Carinifex newberryi (Lea). As genus Carinifex Hannibal, Proc. Mai. Soc. London, X, p. 162. Type Carinifex newberryi (Lea). As subgenus of Pompholyx Carinifex Walker, Miscel. Pub. Mus. Zool. Univ. Mich., Xo. 6, pp. 15, 106. Type Planorbis newberryi Lea. As genus Carinifex Germain, Rec. Ind. Mus., XXI, p. 188. Type Carinifex newberryi (Lea). As genus Carinifex Wenz, Fossil. Cat., Pars 22, p. 1671. Genotype Carinifex newberryi (Lea). As genus Carinifex Dall. Prof. Pajiers U. S. Geol. Surv., Xo. 132 (G) ]). 112. As synonym of Megasystropha Lea Megasystropha Dall, op. cit., p. 112. Type Planorbis newberryi Lea. As genus Megasystropha Lea, Smith. Miscel. Coll., LXXIII, No. 3, Dec. 16. Opinion 87, Int. Com. on Zool. X'^omenclature
Defending use of

1926. Carinifex Pilsbry, Science, LXIV, p. 248, Sept. 10. instead of Megasystropha Lea


1929. Carinifex J. 1929.



Henderson, Univ. Col. Studies, XVII, No. 2, p. 143. Type Carinifex newberryi (Lea). As genus Carinifex Chamberlin and Jones, Bull. Univ. Utah, XIX, No. 4, p. 155. Biol. Series I, No. 1. Genotype Planorbis newberryi Lea. As genus. Carinifex Thiele, Handbuch, Teil 2, p. 480. Type Carinifex newberryi (Lea). As genus Carinifex Pilsbry, Proc. Acad. Nat. Sci. Phil., 86, p. 48. Type Planorbis newberryi Lea. As genus. Figures genitalia of Carinifex ponsonbyi and C. newberryi subrotunda Carinifex J. Henderson, Non-Marine Moll. N. A., p. 255. Discussion of genus and listing of fossil species C. malheurensis Henderson and Rodeck

Shell (plate 78, figs. 23-25). Medium to large, ultradextral, body whorl angulated; spire depressed or elevated, the whorls terraced and angular; base funicular; last whorl broad at the periphery, rapidly attenuated below; aperture triangular, outer lip thin; inner lip with slight callus.

Subfamilies, Genera, and Subgenera

— Recent and Fossil



anatomically the animal
shorter (see plate 34,

the spire of Carinife.v becomes elongated the shell is dextral although is sinistral like that of Hclisomn.
in general form,

Animal. Resembling Helisoma
fig. 1).

but the tentacles are

by many short
34 {Carinijcx ponsorihyi E. A. Smith)

Seminal vesicle (SV) represented ovisperm duct for a length of 4.5 mm. Sperm duct (SPD) long and narrow (2.5 mm. long), diminishing to less than half its diameter as it joins the vas deferens, which is a tube of fine

GENITALIA. Male Organs

vesicles bordering the

long. The prostate (PRS) is elongated (3.5 mm. through the prostate and uterus (fig. 7) shows that the ]:)rostate is peculiar in having the diverticula (14 in this section) occurring laterally from a main pouch into which they empty and which, in turn, empties into the sperm duct. There appears to be no special prostate duct at each end, as in Helisoma. A section through the upper end of the prostate (fig. 8) shows only 11 diverticula but the arrangement is the same. Penial complex (fig. 2). The preputium (PR) is elongated and sac-like, 4.5 mm. long and about 1 mm. wide at its greatest diameter. The vergic sac (VS) is short (1.5 mm. in length) and there is no marked constriction between the two sacs, as in Pierosoma. The musculature is different from that feature in Helisoma. There are two sets of retractor muscles (RM), one set, composed of two main branches with several small branches, is attached to the vergic sac and the upper part of the preputium; the other set, composed of two or more branching divisions, is attached to the upper part of the vergic sac, with one branch connecting with a branch of the first set. The vas deferens passes under this branch. There are three sets of supporting muscles (S]\I) attached to the preputium. These are divided several times in their attachment to tlie preputium and to the roof of the mantle cavity. Several small muscles appear on the base of the preputium below the first set of retractor muscles. Another specimen of ponsonbyi (plate 35, fig. 1) had a variation of this musculature, there being but one main retractor in the first set, and several narrow muscles, much branched, in the second set. The supporting muscles (SINI) were also somewhat different. All specimens have a very short (about 0.5 mm. long) penial gland duct on the outside of the penial complex between the vergic sac and the preputium (DC). This duct does not vary in any of the specimens examined.

diameter, 14-15




Internally (plate 34, fig. 3) there are two fleshy pilasters in the preputium (PL). These are shown in section in fig. 12. The penial gland (01^) is an elongated, somewhat pyriform sac, narrowed at its junction with the body wall of the preputium and enlarged at the cup-shaped cavity. A section of the gland (fig. 10) shows many vertical glandular folds on the wall of the cup and a number of pavement-like folds in the bottom of the cup (IC). The gland duct (DC) extends into the cup and connects with the vertical and pavement-like folds. The verge (V) is short and thick. There is a muscular ridge or diaphragm (D) separating the cavity of the vergic sac from the preputial cavity. Female Organs. The spermatheca (S) is about 1 mm. long, long-ovate in shape and has a narrow duct (SD) almost 2 mm. in length, which enters


The MoUuscan Family Planorbidae

the short vagina (VG) without marked swelling. Another form of spermatheca and duct, from another specimen, is shown in fig. 11. The uterus (U) is tough and leathery, long and wide (6 mm. long and 1 mm. wide), and is covered with wart-like prominences. The nidamental gland (NG) is wide and somewhat globular, covered with small follicles. The oviduct (OD) is much narrower than the uterus and is about 3 mm. long and 0.3 mm. wide. There is a large, swollen carrefour (OF). The relationship of the organs near the junction with the ovisperm duct is shown in fig. 6. The albumen gland (fig. 5) is about twice as long as wide and lies crosswise of the body. The space for the passage of the intestine is shown near the center of the figure.

Hermaphrodite Organs (fig. 4). The ovotestis (OT) is composed of large club-shai)ed diverticula. A section of this organ near the front end is shown in fig. 9. Six rows of diverticula of varying length are to be seen. The sections vary in different parts of the ovotestis. There may be only six unbranched diverticula of large size, or there may be six or seven main
branches, each branch subdivided two or three times to form a surface aspect of fourteen, fifteen, or eighteen diverticula. The free ovisperm duct (SO) is very short and appears only between the oviduct and the seminal

In another species, Carinifcx jacksoneiisis (plate 35, fig. 5), there is in ponsonbyi. The seminal vesicle (SV) forms a grape-like bunching just beyond the ovotestis (about 1 mm. long and 0.7 mm. wide) with lateral vesicles on the ovisperm duct as in ponsonbyi, and for the succeeding 2 mm. with very small vesicles which ap])ear as fine crenulations on one side only. The prostate (PRS) is shorter (only 3 mm. long) and is connected with the sperm duct in the same manner as in ponsonbyi. The diverticula are of the same form and condition as in ponsonbyi. The spermatheca and its duct are also of about the same form (see fig. 9). In figs. 6 and 7, the features near the junction of the ovisperm duct are shown. There is a large carrefour (CF). The albumen gland (fig. 8) is relatively shorter than in ponsonbyi. The penial gland is of the same form as in ponsonbyi (fig. 3) and has the same short gland duct. The retractor muscles (RM), however, are much more complicated in form and position. They are much smaller, narrower, and more numerous. The supjiorting muscles (SM) are small and numerous and are attached to both vergic sac and preputium. Small muscles on the retractor muscle side of the preputium and on its lower half, are attached to the wall of the mantle cavity. These are believed to be supporting muscles. The attachment of the retractor muscles to the vergic

some deviation from the structures as seen


which is the reverse of fig. 3. 50) figured the genitalia of Carinife.v ponsonbyi differing in some respects from the drawings on plate 34. The musculature is nearly the same but is not shown in as great detail as in the figures on plate 34. The spermatheca is more nearly spherical in Pilsbry's figure. The seminal vesicle is shown as a mass of small vesicles, much as seen in Carinifex jacksonensis on plate 35 (fig. 5). The penial complex and penial gland in C. newberryi siibrotunda are figured. It is probable that the sex organs vary somewhat in form, especially those that are evertible. Also, the condition of the preserved material will make a great difference, espeis


in fig. 2,

Pilsbry (1934,

Suhjamilies, Genera, and Subgenera

— Recent and Fossil


cially in those organs like the nidamental gland, the uterus and the oviduct, which may become swollen and thus alter the true form of these organs. Three species of Carinijex have been dissected by Pilsbry and the writer, ponsonbyi, jacksonensis, and newberryi subrotunda, and all show the homogeneity of the group and its distinctness from Helisoma. Respiratory and Renal Systems. The pseudobranch (plate 34. fig. 1) of ponsonbyi is a rounded extension from the rectal region, in form like that in Helisoma, including the rectal ridge which extends over the pseudobranch. In jacksonensis (plate 35, fig. 4), the pseudobranch is narrower. In the specimen figured, the pseudobranch was folded in such a manner as to make the ridge appear on the left margin of the organ. The kidney (plate 45, fig. 2) is long and narrow (15 mm. long, 1 to 1.2 mm. wide) and the ureter is reflexed at right angles to the kidney. A section of the kidney near the middle (fig. 12) shows it to be much flattened, with a bottle-shaped lumen and roundly elongated, and flattened veins. There is no superposed ridge like that in Helisoma. The ridge on the mantle to the left of the kidney is seen in the section to be flattened and wide. Digestive System. The digestive system is similar to that of Pierosoma. The stomach region is shown on plate 48, fig. 3. The intestine is looped about the stomach and liver. The buccal sac is similar to that organ in Helisoma. The jaw is peculiar. There is but one jaw which is horseshoe-shaped, narrow, composed of many small, vertical plates which produce a saw-like edge. In ponsonbyi (plate 49, fig. 19), there are spiral striae on the jaw

plates. In jacksonensis this striation

is not visible (plate 49, fig. 18). The attached to a very large cartilage which has the relative shape and size shown in fig. 17. In fig. 19, the jaw is somewhat distorted. The jaw of Carinijex bears some resemblance to the fragmented jaw of the Planorbinae, but the general form is different.

Radula. The radula of Carinijex ponsonbyi (plate 67, fig. 8) has a square, bicuspid center tooth, the spade-shaped cusps extending below the lower border of the base of attachment. The lateral teeth (1-6) are tricuspid, all cusps sharply spade-shaped, including the mesocone. In some rows, single cusps may become bifid, as the ectocone in the fifth lateral tooth in fig. 8. The intermediate teeth (7-8) differ only in the splitting of the ectocone into two sharp cusps. The marginal teeth (9-27) are long and narrow, the entocone remaining unchanged at first (9-10) but the ectocone undergoing additional splitting. On the eleventh tooth the entocone begins to split, developing three small, sharp cus})s. In the outer part of the membrane, the marginal teeth become narrower and the entocone splits into seven very small, sharp cusps and the ectoconic area becomes simply serrated along the outer margin. The mesocone remains unchanged throughout the entire row. In Carinijex jacksonensis (plate 65, fig. 4), the teeth have practically the same shape, but are smaller. There are ten lateral teeth and two intermediate teeth. In the splitting of the marginal teeth, the inner cusp of the entocone is larger than the four cusps on the inner side of the mesocone. This feature persists throughout the radula. Tlic radula formulae of the two species are as follows: 32-1-32 210-217 ponsonbyi J. Henderson Klamath Lake, Oregon 25-1-2.5 215-221 jacksoijensis Jackson Lake, AVyoming J, Henderson


The Molluscan Family Planorhidae

The data for the specimens examined are as follows: ponsonbiji, from lower end of Upper Klamath Lake, Klamath Falls, Oregon, collected by Mr. and Mrs. J. Henderson, August 7, 1931; ten specimens dissected. Jacksonensis, from Jackson Lake, at Moran, Wyoming, in shallow water just above dam at very low water stage, collected by Mr. and Mrs. Henderson, August 20, 1931; eight specimens dissected. Geographical Distribution. Carinife.v is a purely American group found only in western America from California eastward to Wyoming. Pilsbry believes that all of the described species and races of Carinifex are subspecies of a single stock comparable to the subspecies of Helisoma anceps (Menke). There are, indeed, comparable characteristics between these two groups, although the anceps group is much more widely distributed, both in latitude and area, and is subject to more diverse cnvii'onmental influences than is Carinife.v.
Species Considered as Valid. A careful study of all sj^ecics, many specimens from the type lots, leads the writer to the following conclusion concerning the specific range of the genus. There are doubtless anatomical characteristics which separate some of the s))ecies as has been noted
in ponsoiibiji

and jacksonensis.
Carinifex occidentalis Hanna Carinifex ponsonbyi E. A. Smith Carinifex jacksonensis J. Henderson

Carinifex Carinifex Carinifex Carinifex

newberryi newherryi (Lea) newberryi minor Cooper newberryi malleata Pilsbry neivberryi subrotunda Pilsbry

Geological Distribution. Pliocene to Recent fauna. The genus extend as far back as the Miocene period.


Remarks. Carinifex has been thought to be closely allied to Helixoma and Pilsbry states (1934, p. 48) that there is very little structural difference between the two groujis. There are, however, characteristics of both shell and anatomy which, in the judgment of the writer, are sufficient to mark Carinifex as of generic rank. The ultradextral shell bears some resemblance to certain forms of typical Helisoma but the anatomy, especially
the genitalia, exhibits features of importance. The external penial gland duct is shorter than in any form of Helisoma and is of different form. The prostate, when seen in section, shows the diverticula to be formed in a totally different manner from those of Helisoma. The musculature of the penial complex is far more comjilex with two distinct retractor muscles. The jaw is also of entirely different shape and nature. The radula teeth have the mesocone as it is in typical Helisoma. The kidney is wdthout a

superposed ridge.

By a ruling of the Liternational Commission on Zoological Nomenclature (Opinion 87), the name Carinifex of W. G. Binney has been ruled out of systematic nomenclature because of its publication in what is termed a '])roof sheet' and therefore not usable as a nomenclatorial reference. Proof sheets are obviously not to be quoted as systematic references to publication, but the example chosen to sustain this ruling was most unfortunate, since the publication in which the name Carinifex appeared (Smithsonian Miscellaneous Collections), is an octavo pamphlet of twelve leaves, printed on one side (the left) for corrections. This pamphlet was widely distributed among conchologists and is even now available among secondhand book dealers. The contention that this pamphlet is merely a


Subfamilies, Genera, and Subgenera
'proof sheet' seems

— Recent and Fossil


unwarranted and the writer fully agrees with Pilsbry that Carinifex was sufficiently 'published' by Binney and that this name should be used in place of Megasystropha Lea, which is ruled by the Commission to be used instead of Carinifex. Dall (1870) says (in footnote under Carinifex) 'Megasystropha Lea. Neither genus was characterized by the author, and the former is in general use and has priority.' In 1924, however, Dall reversed this opinion (Prof. Papers, p. 112) and used Megasystropha as the name for this group, contending that the earlier 'proof sheets' of 1863 were simply proofs and should not be quoted. However, in a general review of the literature of this genus, it is evident that the name Carinifex has been almost universally used, as noted in the bibliography of the genus on a previous page of this volume. Henderson (1929, p. 143) well states the situation in the following words. 'It must not be forgotten that the purpose of rules


nomenclature is to insure, so far as possible, stability of names and certainty in their use in the designation of species,- etc., wdiich purpose would be defeated rather than accomplished by this proposed substitution.' The name Carinifex is, therefore, adopted in this work.

Type by
1870. Vortifex


Meek, 1870
Carinifex (Vortifex)

original designation Carinifex tnjoni
Phil., p. 59.


Meek, Proc. Acad. Nat. Sci. Meek, new species. Proposed


as subgenus of Carinifex


1870. Vorticifex Dall, Ann. N. Y. Lye. Nat. Hist., IX, p. 353. originally misspelled. Type Carinifex tryoni Meek.

Correction of Vortifex,

1871. Vorticifex Meek, Tryon, Amer. Jour. Conch., VI, p. 285, April, 1871. Review of Meek's paper. Vortifex corrected to Vorticifex and Carinifex tryoni stated

to be the type
1877. Vorticifex

Meek, U.


Geol. Surv. 40th Parallel, IV, p. 187. Subgenus described

and Carinifex binneyi and tryoni both placed therein
1883. Vorticifex Fischer, Man. de Conch., p. 508. subgenus of Carinifex 1884. Vorticifex


Carinifex tryoni

Meek. As





Carinifex binneyi Meek.
1888. Vorticifex C.^ll,

As subgenus

Conch., Ill, p. 105. Type wrongly stated to be of Carinifex
p. 148.



Genus formally described and Carinifex


Meek wrongly

designated as type

1922. Vorticifex Hann.\, Univ.

Oregon Pub.,



p. 7.

Type Carinifex


Meek. As genus
1924. Paradines

Dall, U. S. G. S., Prof. Papers, No. 132, p. 112. Type by original designation Carinifex binneyi Meek. As section of genus Megasystropha

1935. Vorticifex

Henderson, Fossil Non-Marine Moll. N. Meek. As genus

A., p. 255.



Shell (plate 76, fig. 12 tryoni; fig. 13, binneyi). Large, ultradextral, of few rapidly enlarging, rounded whorls, the body whorl very large, rounded, and sculptured with distinct, rib-like striae parallel with the lines of growth; right side flatly convex; left side with deep 'umbilical' dej^'ession almost completely obscuring the inner whorls; aperture large, rounded. 'The chief distinguishing features of the genus are (1), lack of carinae, either at periphery, suture or around umbilicus; (2), much rounder body whorl than Carinifex; and (3), much less triangular aperture than Carinifex' (Hanna, 1922, p. 7)



The Molluscau Family Planorhidae

Geological Horizon and Distribution. Eocene, Triickee Lake beds, Nevadci; Payette Lake beds, Idaho; Fossil Hill, Kawsoh Mts., Nevada; Pliocene, Cache Valley, Mendon, Collinston, Utah; Tulare formation, Kettleman Hills, California; Warner Lake beds, eastern Oregon. A single species, Vorticifex stearnsii (White) has been reported from the INlorrison formation in Wyoming, belonging in the upper Jurassic or Lower Cretaceous period.
Species Considered as Valid. The following species and races have been referred to this genus.
Vorticifex tryoni (Meek) Vorticifex tryoni concava (Meek) Vorticifex tryoni ventricosa (Meek)

Vorticifex binneyi (Meek) Vorticifex stearnsii (White)

Remarks. Vorticifex appears to be a rather distinct genus related to both Carinifex and Parapholyx. Tlie wide body-whorl with its distinct costae is ciuite characteristic. The name Paradines Dall (1924, p. 112) founded upon the larger species Carinifex binneyi IVIeek does not appear more than specifically distinct from the smaller Carinifex tryoni Meek. Both species seem, from the published figures, to be generically closely related and to be distinct from the genus Carinifex. Several species referred to "Vorticifex appear to belong elsewhere. Vorticifex sanctaeclarae Hannibal and V. sanctaeclarae marshalli Arnold seem rather to belong in Carinifex as originally described rather than in Vorticifex as listed by Henderson (1935, p. 257). They were placed in Carinifex by Pilsbry (1934, p. 567). Tlie Vorticifex laxus of Chamberlin and Berry (Nautilus, XLVH, p. 26, 1933) does not appear to fit into the genus Vorticifex, but might be included in the genus Carinifex. It appears to be an internal cast and the external characteristics would not show clearly in such a condition.
Type by
1912. Perrinilla 1923.


Hannibal, 1912

original designation

Helisoma cordillerana Hannibal


Hannibal, Proc. Mai. Soc. London, X, p. 159. Type Helisoma cordillerana Hannibal. As subgenus of Helisoma Perrinilla Wenz, Fossil. Cat., Pars 22, p. 1518. Genotype Planorhina (Perrinilla) cordillerana (Hannibal). As subgenus of Planorhina Haldeman Perrinilla Pilsbry, Proc. Acad. Nat. Sci. Phil., 86, p. 48. Type Helisoma cordillerana Hannibal. Presumably considered as genus Perrinilla J. Henderson, Fossil Non-Marine Moll, of N. A., p. 246. Type Helisoma cordillerana Hannibal

Shell (plate 79, figs. 31-32). 'Similar to Planorbella, except that it is totally ultra-sinistral; spire-jiit deep and narrow, umbilicus but slightly

excavated, superior and inferior peripheries subangular' (Hannibal for subgenus). 'Shell of considerable size, whorls large, somewhat compressed, and strongly ultrasinistral, the umbilicus barely concave, superior and inferior periplicries subangular in young stages, becoming rounded in adult, growth striae strong, spiral striae occasionally preserved, aperture expanded somewhat in adult; habitat apparently lacustrine. Diameter 22, alt. 9, diameter of aperture 12 mm.' (Hannibal, p. 161, description of Helisoma cordillerana)

Subjaniilies, Genera,

and Subgenera

— Recent and Fossil


Geological Horizon and Distribution. Eocene period. Truckee Lake Nevada, several localities. Miocene. Contra Costa Lake beds, California, Planorbis pablocinus J. G. Cooper, 1894, is believed by Hannibal to be congeneric.

Species Considered as Valid. Perrinilla cordillerana (Hannibal), the genotype, is the only recognized species of the genus.

Remarks. PerriniUa appears to be more closely related to Carinifex than to Helisoma, as suggested by Pilsbry (1934, p. 49), who says Perrinilla Hannibal, 1912 .... seems to stand near the form of the Carinifex phylum
to be expected in the Eocene, Carinifex.



maj' possibly belontj in the ancestral line of


best disposition of the group in planorbid classification appears to be as a distinct genus near Carinifex.

1856. 1857.


Hanna, 1922


original designation




Lea, Proc. Acad. Nat. Sci. Phil., VIII, p. 80. Type efjusa Lea. Not Pompholyx Goose, 1851, Rotifera. As genus



Pompholyx Le.\, Journ. de Conch., VI, p. 208. As genus Pompholyx W. G. Binxey, L. and FW. Sh. N. A., 11, p. 73. Type Pompholyx efjusa Lea. As genus Pompholyx Dall, Proc. Cal. Acad. Sci., Ill, pp. 265, 268. De.scription of subfamily Pompholiginae


Pompholyx J. E. Gr.\y, Ann. Mag. Nat. Hist., (4) II, p. 387. As genus Pompholyx Dall, Ann. N. Y. Lye. Nat. Hist., IX, pp. 334, 344, 353. Type Pompholyx efjusa Lea. As genus. Anatomy imperfectly described Pompholyx Tryon, Con. Hald. Mon., pp. 83, 174. Type Pompholyx efjusa Lea.
As genus

1884. 1886. 1912.

Pompholyx Fischer, Man.

de. Conch., 507.

Type Pompholyx

efjusa Lea.


Pompholyx Tryox,
As genus





Ill, p. 105.


efjusa Lea. efjusa Lea.

Pompholyx Clessix, Conch.
As genus


p. 226.

Type Pompholyx

1918. 1922.

1927. 1929. 1931.
1934. 1935.


Proc. Mai. Soc. London, X, p. 162. Type Pompholyx As genus Pompholyx Walker, Miscel. Pub. Mus. Zool., Univ. Mich., No. 6, pp. 14, 105. Type Pompholyx efjusa Lea. As genvis Pompholyx Germaix, Rec. Ind. Mus., XXI, p. 185. Type Pompholyx efjusa Lea. As genus Parapholyx Hanxa, Univ. Oregon Pub., I, No. 12, p. 5. New name for Pompholyx Lea, 1856, non Goose, 1851 Pompholycodea Lixdholm, Proc. Commission for study of Lake Baikal, II, p. 180. New name for Pompholyx Lea. not Goose Parapholyx J. Hexdersox, Univ. Col. Studies, XVII, No. 2, p. 145. Type Pompholyx efjusa Lea. As genus Pompholycodea Thiele, Handbuch, Teil 2, p. 480. Type Pompholyx efjusa Lea. As genus Parapholyx Pil.sbry. Proc. Acad. Nat. Sci. Phil., 86. pp. 46, 51. Type Pompholyx efjusa Lea. As genus Parapholyx J. Hexder.sox, Fossil Non-Marine Moll. N. A., p. 254. As genus Parapholyx F. C. Baker, Nautilus, L, pp. 30-31. Notes on anatomy
efjusa Lea.

Pompholyx Haxxibal,


The Molluscan Family Planorbidae
Of medium size, ultradextral, globose, the above the body whorl which is ventricose
aperture wide, greatly expanded; outer

Shell (plate 78, figs. 20-22). spire short and but little raised

and embraces most of the
lip thin, acute,


reflexed in adult or old specimens; inner lip more or less thickened; base convex or conical, imperforate or with but a small chink.


Animal (plate 36, fig. 1). Foot short, about half as broad as long. Velum as in Helisoma. Tentacles short, cylindrical, blunt at the end, not tapering. Eyes sessile at the inner base of the tentacles. The eyes are not conspicuous in some preserved specimens. The body is darkly pigmented,
the pigmented areas extending to the head and sometimes forming streaks on the tentacles. The posterior portion of the body is light gray. The ovotestis area is yellowish.

36 {P(imi>hohjx cffuM, klamnthcnsis F. C. Baker)
(fig. 3). Seminal vesicle (SV) about 4 small vesicles projecting from the otherwise smooth surface. The sju^rm duct (SPD) is about 3 mm. long and of large diameter (0.4 mm. L The prostate (PRS) is long (3.5 mm.) and wide and is composed of many small diverticula. In section (fig. 4), these diverticula are seen to be composed of about a dozen irregular follicles linked together like sausages. From the outside of the prostate, these diverticula appear to be covered with small tubercles irregularly placed. These follicles enter the sperm duct at a common center, shown at in the figure. The vas deferens is about half the diameter of the sperm duct and is 11 mm.

GENITALIA. Male Organs
long, swollen, with





The penial complex (fig. 7) is sac-like and is nearly 4 mm. in length. The preputium is 1 mm. wide and occupies most of the length of the penial complex. The vergic sac (VS) is 1 mm. long and 0.7 mm. wide. There
only a slight constriction between the prejiutium and the vergic sac. penial gland duct is relatively short (about 1.5 mm. long) and rather thick. It extends from the i)reputium near its upper third and enters the vergic sac at the constricted area. The portion of the duct entering the vergic sac and the preputium is very small and narrow, the duct swelling greatly in diameter on the surface of the preputium. There are two sets of retractor muscles, each set consisting of two main branches which subdivide several times at the penial sac insertions and at the connections in the columella muscle. All insertions are on the preputium, none on the vergic sac. There are several narrow supporting muscles (SAI) attached to the preputium. Internally (fig. 9) the preputium has two large, fleshy folds or pilasters (PL). There is a mucii elongated, cylindrical penial gland (1.7 mm. long) which resembles a pipe-stem, with a cup-shaped or bowl-shaped gland (about 0.5 mm. in diameter) placed at the lower end (GL). The duct (DC and X) enters the cylindrical portion of the gland and extends through this stem-like portion, then enters the bowl-like cup of the gland where it is enlarged into a bulb-like body (fig. 11, X). The cup-shaped gland resembles the bowl of a pipe. In section (fig. 11), the cup shows a shallow area at the top and the sides are lined with wide, thick folds (GF). The bottom of the bowl-like cup is covered with small follicles which are



Suhfamilies, Genera, and Subgenera

— Recent and Fossil


placed at the upper end of the rounded enlargement of the penial gland The right pilaster is attached to the gland at its upper end, curving toward the gland beneath the gland duct. There is a thick ring or diaphragm (D) which separates the cavity of the preputium from the vergic cavity. This diaphragm has a circular canal into which the upper end of the penial gland duct, which is greatly diminished in diameter, enters. This feature is shown in fig. 10. The verge (V) is narrowly pyriform, tapering to a point at its lower end. The sperm canal extends through the center of the verge and has a central outlet (fig. 121. A large nerve and blood vessel supply the penial complex (fig. 8).

Female Organs (fig. 3). The spermatheca (S) is large and egg-shaped, almost globular. It is connected with the short, narrow vagina (VG) by a narrow duct shorter than the length of the spermatheca (SD). The uterus (U) is narrow as it passes upward from the vagina but increases abruptly in diameter to three times the width of that region, combining with the nidamental gland. This region is concealed by the prostate in fig. 3. The oviduct (OD) contracts in diameter to form a sac-like organ of small diameter. There is a large carrefour (CF). The albumen gland (AD is 4 mm. long and is longer than wide. The duct of this gland is long (0.5 mm.).
Hermaphrodite Organs. The ovotestis (OT) is long and sac-like and composed of several club-shaped diverticula. A section near the middle of the ovotestis (fig. 5) shows eight rather long diverticula, the longest follicles being in the center. A section near the posterior end (fig. 6) shows six diverticula in less fan-like pattern. Sections from the middle to the anterior end show eight diverticula all in fan-shaped pattern, but sections made toward the posterior end show but six diverticula which are more or less irregularly arranged. The ovisperm duct is very short in the portion between the ovotestis and the seminal vesicle. The free portion near the oviduct is somewhat longer. The seminal vesicle (SV) occupies about twois

thirds of the length of the ovisperm duct. The genitalia of Parapholijx effusa diagonaUs Henderson, also dissected, do not differ from the figures and descriptions herein presented. In 1870, Dall dissected Pompholyx soUda and published descrii-)tions and several figures of the genitalia. These are not in sufficient detail and embrace some errors, such as the presence of a distinct testicle on the male system. This might have been the albumen gland. As far as can be seen,

anatomy of solida is similar to that of the species personally examined and herein figured. Respiratory and Renal Systems. The pseudobranch (fig. 2) is a triangular appendage extending from the rectal region. There is a distinct ridge extending over the rectum in a broad curve, ending above the anus. The kidney (plate 46, fig. 1) is short and wide (about 3 mm. long and 1 mm. wide) the ureter is large and sharply refiexed. directed upward into the mantle cavity. The pericardium is also large (about 1 mm. long). A cross section of the kidney near the middle (fig. 2) shows a much flattened form with a wide lumen, the veins ovate and flattened, and placed at the lower lateral sides of the lumen. There is no ridge and the section rethe

sembles that of Carinifex. Digestive System. The stomach is like that of Carinifex and essentially as described by Dall for Pompholyx solida. The oesophagus is short


The Molluscan Family Planorhidae

enlarged just behind the buccal sac. The salivary glands form a The buccal sac is short, high and pyriform, as in Carinifex. The jaws are three in number, differing markedly from the single horseshoe-shaped jaw of Carinifex. The superior jaw is low and about three times as wide as high. The side jaws are as in Helisoma. The face of the


superior jaw is striated vertically. There is little difference between the jaws of klamathensis and diagonalis. Dall says there are no accessory plates (side jaws) but they occurred in all specimens personally examined. The jaw figured by Dall (his plate 2, fig. 11a) is different from the jaws found in the two races examined. The radula of P. effusa klniitathoi.'iis (plate 65, fig. 3) has a squarish center tooth, the bicuspid reflection not reaching the lower margin of the base of attachment. The lateral teeth (1-8) are scjuarish and tricusi)id, all cusps being sharp and spade-shaped. The intermediate teeth (9-11) are narrow, the ectocone splitting into two sharp cusps as long as, or nearly as long as, the mesocone. There are two small cusps on the outer border of the tooth, above the ectocone. The marginal teeth (13-15) are narrow, the entocone with four to five small cusps, the ectoconic region with three cusps. The outer marginal teeth (16-19) are very narrow and are simply serrated on the ectoconic side, there being no division into entocone, mesocone, and ectocone. Ball's figure of the radula of solida (his plate 2, fig. 9) shows the lateral teeth with a rounded mesocone and all cusps appear too blunt. The marginal teeth are too wide and the small cus])s are not shown.

Radula Formulae of Paraphnhjx

19-1-19 21-1-21 22-1-22



J. J.

diagojialis solida

Klamath Lake, Oregon
Crater Lake, Oregon

White Pine, Nevada

125 100-114 150

Henderson Henderson

Ball's paper

following material has been examined for the anatomical data contained in this work. P. effusa klamathensis from outlet of Upper Klamath Lake, Oregon, eight si)ecimens examined. P. effusa diagonalis from Crater Lake, Oregon, eight specimens examined. Both lots collected by



Geographical Distribution. Parapholyx is purely an American genus confined to the west coast of America from Washington southward to California and eastward to Nevada. Pompholyx leana H. and A. Adams was described from 'West Columbia,' but no specimen of this genus has been seen from British Columbia and it is very doubtful that the genus is found north of the United States. A number of species and races have been described and the same opinion expressed for the division of Carinifex may also ap]^ly to this genus.
Species Considered as Valid.
Parapholyx ejjusa Parapholyx ejjusa Parapholyx ejfusa J. Henderson Parapholyx ejfusa Baker
effusa (Lea)

coHata (Hemphill)

klamathensis F. C.

Parapholyx Parapholyx Parapholyx Parapholyx Parapholyx




mailliardi Hanna solida solida (Dall) solida optima (Pilsbry)

leana (H. and A. Adams) Species practically unknown

Geological Distribution. Parapholyx is certainly known from the Pliocene to the Recent fauna. It may occur in earlier deposits. One extinct

As genus . the four eyes. 814. N. expanding in the bottom of the })enial gland cup. Henderson (1929. Genus PO^^IPHOLOPSIS Type 1888. and Subgenera — Recent and Fossil 165 species is known.v packardi in eastern Oregon. Pompholopsis Call. INIus. p. Geol. Dall obtained specimens of a Fompholyx iFarapholyx) different specifically from the known species.. I. 14). 507). p. 1888 designation Pompholopsis tvhitei Call 1935. As genus Pompholopsis J. Type Pompholopsis ichitei Call. The penial gland duct is external as in Carinifex. S. In 1866. as in Carinifex. at the inner base of the tentacles. 147. p. 352) although the chief reason for its adoption. p. p. pipe-shaped body with a cup-like termination. 255. but it is somewhat longer and of greater diameter. one at the inner base of the tentacles and one at the end of the effusa. The prostate is not composed of distinct clubshaped diverticula. The penial gland is not a short. The shell is depressed and almost neritaeform. Fossil Xon-Marine Moll. 161) elevated it to family status as Pomplioligidae. Type Pompholopsis whitei Call. 155). Gray 1868. 143). The genus stands out as one of the most distinct among American planorbids. who made drawings of the animal. 387) questioned this statement. basing the separation tentacles. p. 1901). This course was indicated by Pilsbry in 1934 (p. showing its general relationship to Helisoma and Carinifex and stating that the old subfamily Pompholiginae was without anatomical distinction. Hannibal (1912. b}' original Call. A. but an elongated. the supposed eyes on the end of the tentacles being simply an aggregation of pigment.Suhfaniilies. In 1936 (Nautilus. J. with a long duct passing through the center of the longitudinal. p. Nat. 185). p. Only one pair of eyes is present. Dall has also used the name Pompholiginae for a section of the marine genus Divaricella (Proc. E. The jaw' is like that of Helisoma and is not fragmented like that of Carinifex. Dr. and Chamberlin and Jones (1929. XXIII. but of small gland-like bodies arranged like a string of sausages. from Warner Lake deposits Remarks. 3. Hexdersox. Dall still retained the subfamily Pompholiginae (1870. p. cup-shaped organ. The termination of the duct is swollen and bulb-like. Dall instituted a subfamily Pompholiginae for Fomphoh/x on the supposition that there were two pairs of eyes. p. An examination of the tentacles of this species at once cleared up the anomaly of the four eyes. Trvon (1884. including Fischer (1883. which he called solida. No. In 1870. sac-like 'stem' of the gland. fact of this peculiar condition was positively stated by Gabb.. Parapholy. 30). Amer. Hanna. L. The subfamilv has been used bv most writers on fresh-water Mollusca. Walker (1918. the writer indicated the general anatomical features of Farapholyx. The anatomy is peculiar in several respects. had been found to be invalid. p. Germain (1922. The study of the animals of Carinifex and Farapholyx shows that they have no general anatomical characteristics not shared by Helisoma and the subfamilv Pompholiginae must therefore be abandoned. U. 46). The Mr.. p. as is the case in other species of Planorbidae. p. William ( ]\I. Genera. 105). Parapholyx is a very distinct genus both in shell and anatomy.

Akad. 445. As genus Planorbis Moxtfort. p. Coretus (Gray 1847) Wenz. might add greatly to our knowledge of the relationship Genus Type by PLANORBARIUS Froriep. Type Planorbis corneus (Linn. pp. Ann. As genus 1840. II. Hist. IX. Miscel. Berlin. Coretus We. fig. 1917. CoreUis (Adanson. 81. Lye. V. N. Coretus Clessin.\y. Fauna Pal.' (Call. As genus Planorbis Soos. As genus p. Hungarici. Type Planorbis corncus As 1883. p. XVII.) Walker. 6. Cab..). 10. Type Planorbis corneus (Linn. apex obtuse. 9. bis As genus Muscheln Type (1st species) Planor- corncus (Linn.\rd. surface of the shell smooth. Type Planorbis corneus (Linn.). France. Treat. No. Zool. sjiire short. As subgenus 1905. Pompholopsis differs from Parapholyx in being deeply um- bilicated and in having a high spire and smaller. 1421. Spirodiscus Stein. Planorbis Dall. No type cited. XV. body-whorl very large. Pub.26 mm.) 1855. p. Proc. 270. and S. Type Planorbis corneus (Linn. Musei Nat. Ponipholopsis ivhitei Call is the only Remarks. 73. As subgenus Spirodiscus Westerlund. Tyjie Helix corncus Linn. Planorbarius Kennard and Woodw. Spirodiscus Fischer. Jugoslav. 37... Ann. 1810. Post-pliocene (Pleistocene). Soc.andon. Type Pla: 1850. As genus 1847. ZooL. Y. Analyt. As subgenus of Planorbis 1921. Univ. Type Helix cornea Linn. Grovrth lines inconspicuous. p. 165. H.. California.. As subgenus of Planorbis 1923. 1806 original designation Helix cornea Linn. Tassajara Lake beds. deeply iimbilicated. Planorbis Tryon. of Planorbis 1886. (Linn. 151. diameter 8.).). 32. Cat. Schn. Planorbis Germain. Coretus Moquin-T.166 The Molluscan Family Planorhidae Shell (plate 76. Geological Horizon and Distribution. Conch. Reg. Planorbis 1924. 1884. Planorbis Swainson. sutures rather irregularly impressed. in Dumeril. Genotype Coretus corneus (Linn. Fluv. A scries of shells of this species. 1757) norbis Cornells (Linn.. As genus 1885. 120. As genus 1918. 337. whorls three to three and one-half. Conch. London. S. Type Helix cornea Linn.13 mm.). de Conch. p. 147). pp. Species Considered as Valid. section of Planorbis 509. . Rec. Man. Soc. pp. Length 7. p.. 1902. Mus. London. Rad. Ind. 1888. globose. peritreme continuous. p. Type Helix cornea Linn. Malac. p. XVI.. (Linn. 'Dextral. Geoffroy or Miiller. rather solid. 6.).). 106. Mai. Conch. 424. 1806. Planorharius Froriep. H. Mus. rounder aperture. N. aperture roundly ovate. 94. p. p. Gr. Moll.)... forming a heavy callus on the parietal wall of the body-whorl. Fossil.sterlund. Alaska Moll. p.). p. Type Planorbis corncus Type Planorbis corncus 180. species assigned to this genus. Proc. p. Pars 22.. 11). 351. Terr. As subgenus Planorbis Dall.. Type Planorbis corneus (Linn. (Linn. convex. Zool. XXI. Ill. Type Planorbis corneus (Linn. 64.). u. 140.) 1870. It is related to Parapholyx but appears sufficiently distinct to justify a genus. p. Syst. Type Planorbis corncus (Linn.. Non p. Mich.

Male Organs mm.. Baker. p. Moll. London. N. Proc. Brit. when feeding.). The heart may be observed beating rapidly. Color of living animal lead. Type Planorbis corneus (Linn. prostate in section <fig. in the same plane as the whorls.Subfamilies. The radula sac may be seen working backward and forward. the seven to nine main diverticula branching several times (usually lour) toward the outer part of the section.5 mm. lips sharp. year 58. Soc. tentacles bluish black. wide. The body shows pinkish through the shell. the figure being a cross section cutting through these organs. Syn. Fluv. of few gradually enlarging whorls. The . Specimens of the young with bifid tentacles are not uncommon. As genus 1931. not thickened. Tcil 2. Figure 4 also shows the relationship between the uterus. The seminal vesicle (SV) is about nun. p. The duct of the prostate is a continuation of the vas deferens and is about 1 mm. 67. p. Coretus (Gray 1847) H. Mus. Planorbis Subgenus Thiele. 22-24). Planorbis F. No type cited but Planorbis corneus (Linn. ANATOMICAL CHARACTERISTICS PLATE 4 20 {Planorbarius corneus (Linn. Genotype Helix cornea Linn. Handbuch. Type evidently Helix cornea Linn. As genus 1929. Type Planorbis corneus (Linn. Planorbis — Recent and Fossil Non-Marine Moll. and tapering to a point. 378. head and foot with border of white on edge. Nat. As genus 1931. 1774 figs. aperture slightly enlarged. 4) is radiate or fan-shaped in form. Helix cornea Linn. The vas deferens (VD) is a narrow tube about 12 mm. Genera.) (fig. long. the vela area large. 1757) Germain. Large. Where it leaves the ovisperm duct it is of small diameter. As subgenus of Planorbis 1931. 583. sometimes nearly black tinged with brown. 252. rather solid. Amer.. The young snail of two whorls. ) GENITALIA. discoidal. the prostate diverticula radiating from this duct. France. the nidamental gland. Zool. long. Heft 6. Sometimes the whole body is reddish. Planorbis Pilsbry and Bequaert. as in the adult animal. Barcelona. sinistral. Planorbiirius 1926. Museo Cien. 4. wide at the base. Ideogenotype Helix cornea Linn. long and 1 13). Bull. Trab.^As. LIII. long. The prostate diverticula may be noted to enter the prostate duct directly. fiir Mollusk. tentacles long. Shell (plate'80. C. is very translucent. 517. 167 Type Kennard and Woodward.) described. p. This organ is a rounded mass of small vesicles. tapering behind. p. equally visible above and below. XIII. type by Montfort 1927. but it gradually enlarges to twice this diameter near the prostate where it abrui^tly expands to several times the first diameter. recently hatched from the egg capsule. 116. 115. the organs plainly visible through the hyaline shell. wide foot. of Planorbis Miiller. Terr. rounded before. II. pp. H. eyes sessile at their inner bases. narrowing at the anterior end to the diameter of the ovisperm duct. Coretus (Andanson. long. It lies over the sperm duct with which it is connected. The prostate (PRS) is elongated and somewhat fan-shaped. and the prostate. like The young snails are distinctly physoid in form and glide about Physa. 479. and then suddenly diminishes again to almost its original diameter. and Subgenera 1926. A number of small vesicles continue down one side of the ovisperm duct. As genus. p. With short. Archiv. The sperm duct (SPD) is about 10 nnn..). Animal. As subgenus Lindholm.

There is a narrow canal or channel which extends down the center of the appendage for its entire length (C). The nidamcntal gland (NG) is a large saclike organ 3. about 1. 6. long and 2 mm. somewhat enlarged toward the lower end. There are two retractor muscles (RM). 11). ending in two lateral swellings which notably resemble the glans penis of the human genitalia. tapering to 0.5 mm. 5). the penial gland has the form shown in fig. wide. The verge (highly magnified) is shown in fig. in diameter (see fig. The vergic sac and a part of the vas deferens are attached to the surface of the preputium. The oviduct (OD) is over 4 mm.7 mm. There is a narrow vergic sac cavity VO) The channel which extends down the center of the penial gland appendage is seen to extend upward and into the vergic sac cavity (C). an upper one attached to the preimtium near the vergic sac by two branches. there is a peculiarly shaped penial gland (GL). It is shown somewhat restricted in fig. one branch on each side of the vergic sac. long) and very narrow and is attached to the vas deferens at about the apex of the preputial sac. The gland becomes a rosette surrounding the opening into the vergic sac (X in figure) and the gland appendage (GA) extends forward (or downward) in the preputial cavity. Hermaphrodite Organs (fig. The spermatheca (S) is large and pyriform in general shape (fig. It is about 8 mm. In some specimens. vertical pilasters (PL). 3. There is a conspicuous gland appendage (GA). The uterus (U) begins with the same diameter as the vagina but rapidly enlarges to about 1 mm. 168 The Molluscan Family Planorbidae The penial complex (fig. NG). in diameter. VIM). It is composed of many diverticula ( . There are four or five sui)porting nuiscles (SM) on the opposite side from the lower retractor muscle. It is short and narrow. 12. In section (fig. 4.5 mm. 8 (PA). The vergic sac (VS) is very small (about 1 mm. 12. a trifle more than 1 mm.5 mm. wide. 13 ». wide at the male opening. with a centrally located sperm canal and exit. When fully extended. RM.5 mm. long and 1 mm. 7. with smaller muscles joining the two larger muscles and a small muscle extending to the vas deferens (fig. and a lower muscle attached at about midway of the preputium. as shown in fig. Female Organs. The ovotestis (OT) is very long and occupies a little more than one whorl. which form a pavement-like pattern on the surface of the ovotestis. as shown in fig. the large preputial cavity has two heavy. the organ shows seven main diverticula each of which is . Internally (fig.5 mm. long) and is placed at the summit of the preputium.. long and is a large flattened tube about 0. wide. long. lying on that organ. There is a large carrefour ((5Fj. 9) is squarish in shape and is composed of many small follicles. This form of musculature was more common than the one shown in fig. The vagina (VG) is narrow. There is a small penial appendage beside the sperm canal outlet of the verge which is flattened and extends diagonally from the end of the verge. long and less than 0. The duct (SO) is 2. the two retractor muscles are placed one above the other attached to the preputium near the upper part. The verge (V) is very small (0. 13). In the upper part of the preputium. 1. 12) consists of a very large. long and about 2 mm. A cross section of the appendage indicating the ]X)sition of this canal is shown in fig.5 mm.5 mm. The albumen gland (fig. long and is a narrow tube. pyriform preputium (PR). 11 and drawn upward in the preputial sac. wide at the upper part. 2 (C).

Genera. 1912. In fig. Germain's figure of the genitalia of Planorbis corneus (1931. The rectum (R) is large. p.Subjamilies. fig. p. 1. 308) has copied these figures and has erroneously accei^ted the division into types of the male organ in Planorbis. Soos's figure (1917. 18) shows a rounded lumen (TK) with a vein on each side at the lower angle of the lumen (RA. p. fig. The pseudobranch (plate 20. plate 4) correctly figures the cylindrical appendage and gland but the appendage is erroneously called the penis. and Subgenera — Recent and Fossil 169 divided into four secondary branches. LacazeDuthier's figures are also accurate as regards general anatomy and nerve organization (1872. Baker 1928. the first type of the penis (penis complex) of corneus is an error of interpretation. which does not extend over the pseudobranch as in Helisoma. p. 13 on plate 20 in general gross anatomy. pneumostome (PS). the lobations being more numerous than in Helisoma corpulentum. F. The portion of the kidney bordering the pericardium is flattened or even a trifie concave. The free portion of the ovisperm duct (SO) is about 5 nnn. Taylor (1900. There is a is large fig. and curved (RD). Baker's paper appeared in 1931 (p. long between the vesicle and the ovotestis. 502) shows the penial complex incorrectly. elongated body extending from the anal region. which has a channel down the center. shows the gland of corneus which is called the penis (verge). There is a high. long between the seminal vesicle and the oviduct and only 1 mm. The kidney is in general 3) except that it is . tafel 26. a single ripe ovum is shown in the diverticulum. C. to the left in the section. which it ( ( probably is. plate 1). rounded crest or ridge. the lower part is rounded and lobate on both margins. Other authors. the lower part being designated as a dilation and the end of the groove is said to represent the stylet. Internally. The published figures of the external and internal aspects of the penial complex have been inaccurate. is wide. 3. plate 20). either in figuring or in the interpretation of parts of the organs. C. tafel 5. The small verge appears to be an enlargement of the vas deferens and has been completely overlooked. AP). 366) figures the terminal bulbous portion. That of Simroth (1912. L. copied by Simroth. longer and wider (14 mm. I. wide at the upper part). It is usually somewhat leaf-like but. 10) an oval. 3) is one of the best and agrees with our fig. like that of Helisoma corpulentum (plate 44. calling it the vibratile stimulating appendage. The Respiratory and Renal Systems. rounded. the penis of Buchner not being the true verge but the gland appendage. There is a large ridge extending down the center of the kidney. Buchner (1891. p. 579. sharp crest or ridge (RK) causing the section to be higher than wide. thought to be the sperm canal by Buchner. The ridge on the mantle. 37 lacks detail. fig. it may often be folded forming a rounded tube (P). high. The published figures of the genitalia of Planorbarius corneus vary considerably. have also used these penial subdivisions. as Annandale (1922). Moquin-Tandon (1855) on his plate 32. true nature of the gland and its appendage and the very small true verge have not been recognized and figured until F. and bears a simple. fig. The anal opening is above the pseudobranch (A). Baudelot (1863. 518) lacks details of structure and is less clear than that of Simroth. 3. fig. long and 4 mm. A cross section of the kidney near the lower end (plate 45. as in the figure.

there were odd teeth (72nd row. A. Roger P. fig. T. Van Hyning. Center tooth higher than wide. elongated. p. The mesocone remains large in all teeth with but little modification. p. The face of the jaw is heavily vertically striated. Poland. In some specimens from Germany. Puerto Rico. The outer marginals (39-42) are very narrow.170 The Molluscan Family Planorhidae Digestive System. Great Britain and Europe east to northern and western Asia. . Jankowski. marginal) in which the cusps of the endocone were uneven and the smaller ones might be assumed to be interstitial cusps. The stomach region is rather narrow and resembles that of Promenetus exacuous figured on plate 48 (figs. Lateral teeth (1-9) squarish. Roger P. tricuspid. the ectocone becoming tricusind. 14) is wide and low. Jankowski Germany 45-1-45 170-210 Dr. in this respect diff'ering from the salivary glands of Helisoma. 20. rounded radula sac. Radula (plate 66. The salivary glands are short and are composed of large branched follicles. Ojus. A. It has been introduced into New Jersey. F. Gray. convex. F. Some outer marginals (17. Germain (1931. 22) were of a like nature. The buccal sac is pyriform. Haas. the entocone breaking up into two to four sharp. pointed cusps. the two spadeshaped cusps not reaching the lower margin of the base of attachment. 5). fig. Florida. south to Portugal and Italy. the cusps becoming very small and forming a serrated outer margin of the teeth. All specimens studied have had tricuspid lateral teeth. Whether the larger species found in northern Africa are referable to Planorbarius can only be determined bv an antomical examination. Gray- Material from the following sources has been used for the above data: Branch of Wista River at IVIorysinek. 270) figures the laterals as bicuspid and in this he is followed by Simroth (1912. 5 km. and probably other places. Intermediate teeth (10-14) narrower than the lateral teeth. p. the ends attenuated and bent downward. sent by Air. Marginal teeth (15-38) narrow. the cusps very small. Air. as has been noted for every genus in the family as yet examined. much smaller than the other marginals. Haas Florida (aquarium specimen) 45-1-45 200-218 Mr. Copenhagen red snails. the ectocone becoming bicuspid. Geographical Distribution. Taylor (1900. collected by Mr. The side jaws are very narrow and as long as the width of the superior jaw. the entocone breaking up into five to six subequal cusps. Germany (locality unknown) from Dr. The digestive system is like that of the Helisomatinae in general. bred for aquaria. south of Warsaw. 2). There is a large blind sac or caecum. 4. The following radulae have been examined: Locality Source Formula Rows Poland. The superior jaw (plate 50. the cusps sharply pointed. near Warsaw 42-1-42 200-217 Mr. The intestine makes a coil about the stomach and another around the liver. with a short. the mesocone larger than either the entocone or ectocone. 12-15. 518) figures the radula of corneus as having interstitial cusps. 316). but such were not observed in the material personally examined. all in a straight row. Taylor also figures thirty-three teeth in a row while in specimens personally examined the number has been from forty-two to forty-five.

1810. Dall's conclusions were questioned by Kennard and Woodward (1924) who maintained that the type of the genus Planorbis is the Helix planorbis of Linn. Many authors have considered it the type of the genus Planorbis of Miiller.) Geological Distribution. dating it from Montfort. H. the genus Corneus = Planorbarius) begins in the late Eocene period. as has been observed in Helisoma. p. According to Wenz (1923. It is the only group of the subfamilv thus far known outside of America.) Planorbarius kabylianus (Let. Baker (1930) has also expressed the same opinion. extralimital. The large shell resembles that of Helisoma especially the subgenus Pierosonia. Germain includes in Planorbis (= Planorbarius) several species which belong in how many of a question. Remarks. in a later work (1931). Geoff roy 1767. vestigial true penis. Lindholm (1926).) Species Considered as Valid. by absolute tautonomy.) Planorbarius elophilus (Bgt. Dall (1905) considered it the type of Planorbis. name Coretus is in current use by many European conSome of these use the Coretus of Adanson 1757 (as Clessin 1886 and Germain 1931) but that author was non-binomial and preThe generic . The presence of the penial gland and the multiple character of the diverticula of both prostate and ovotestis indicate this relationship. 1481). The very large preputium and very small vergic sac are particularly noteworthy. 1931) includes one species with several varieties. Germain. The structure of the prostate and ovotestis is likewise distinctive. The strongly ridged kidney is also another evidence of this kinship. Germain other genera. In the catalogue of Planorbidae in the Indian Museum. who definitely established cornea as the tvpe of the genus. The two retractors of the penial complex are also diagnostic features. has been in some dispute. Baker 1931) and Thiele (1931). The systematic position of Planorbarius is without doubt in the subfamily Helisomatinae. and Subgenera Just — Recent and Fossil 171 species may be included in Planorbarius is somewhat (1921.Subfamilies. Other authors have ignored all details of the internal anatomy of the penial complex. The genus Planorbarius is at once distinguished from all other groui)s of Planorbidae by characteristics of the. cites the Helix planorbis of Linn. and the form of the penial gland with its elongated appendage and the very small verge are unique features of the genus. Genera. genitalia. The proper genus for the reception of the Helix cornea of Linn. B. Planorbarius corneus (Linn. as the tvpe of Planorbis Guettard 1756 and ( "( . This procedure has been followed bv Germain (1921). No specimens with everted gland or verge have been observed and one can only surmise that the gland is everted as an excitatory organ and probably the small vergic sac and verge are extended from the male opening. Almost all writers on European fresh-water Mollusca have mistaken the penial gland appendage for the true penis (verge) and have completely missed the small. Pilsbry and Beciuaert (1927). Since the members of this genus are Westerlund which appear to be referable Planorbarius metidjensis (Forbes) Planorbarius stenostoma (Bgt. principally Afroplanorbis. simply figuring the large pre]:)utium and the vas deferens as directly entering this organ. the species listed by to Planorbarius are cited here. chologists.

Part IV. As subgenus 1865. several (four to six) smaller diverticula. London. 1923. Genera of Rec.. Mon. p. 508) used the name Coretus. the Helix cornea of Linnaeus which. Z. Micromcnctus. preoccupied Planorbula H. No type cited. 1847. Proynenelus. and A. in this The fossil group Planobifex Pilsbry doubtless belongs subfamily. 2 (Oct. Planorbula Binney. This antedates INIontfort's use of corneus as a genotype by four years. Dumeril (Zool. and FW. 1840. Type Planorbis armigerus Say. Non Fitzinger. In 1847. Monotype Planorbis arynigerus Say. 1929) use Coretus Gray. II..S. New name for Discus Haldeman. The that of Coretus Gray and there appears to be as published by Froriep. non Beck. As subgenus Dentatus Gray. however. 181. on one side. Es sind dies Flussschnecken.-<c}(s PLANORBULA Haldeman. 1837.: 172 The MoUuscau Family Plauorbidae Linnean. Mon. becomes the monogenotype of the genus Planorbarius of Froriep. p.Z. the group might pass for anything among fresh-water pulmonates and the name is absolutely useless for purposes of nomenclature. Moll. he applied the name to a minute species about an eighth of an inch in diameter. Suppl. In 1806. Genus Type by 1840. Froriep used this name and supplied a type.. cit. Type Planorbis armatus Gray. As subgenus of Coretus H. Sh. The jaws are as in Helisomatinae. II. 14. main Type genus Planorbula Haldeman. Proc. Froriep says of Planorbarius in his German translation (p. p. a land genus. The few groups belonging in this subfamily With are noted below: short. without an operculum. inhabiting fresh water. There is a pcnial gland but no external duct. but Planorbis armigerus Say by assumption. with the type Helix cornea. p. A. 136. N. Limn. 4 of cover. The rarity of Froriep's translation in libraries has doubtless been the cause of its being omitted in most })rcvious discussions of planorbid nomenclature. as of Adanson. p.'VLDEman. Soc. Zool. Subfamily Prostate diverticula PLANORBULINAE Pilsbry. according to Opinion 46 of the International Code. 164) coined the name Planorbarius for a group of mollusks.. July. furthermore. Gray (P. Except for the name. supposed to be the same as Playwrbis armigerus Say.B.. Type Planorbis armigerus Say. there being a division from which branch. In a German edition of Dumeril's work. Plariorbula . should not be used for the large European snail jireviously known as Planorbis corneus. L. 1840 Say original designation Plajwrbis armigerus 1840. op. 265. p. p.. 1847. Physidae. to Part I. 165) Die Scheibenschnecken {Planorbarius) iinterscheiden sich (die Schaele ubemicksichtiget) \on den vorigen niir durch den mangelnden Schliessdeckel. The ovotestis diverticula are placed in pairs. p. case is no reason much better than why Planorbarius. Helix cornea. Without distinct duct but with open channel: Menetus. which is always placed inside of the preputium. As subgenus Planorbula Haldeman. As subgenus of Segmentina 1855. Haas. 1833. but definitely cited the Helix cornea of Linnaeus as the type. 1934 compound but not fan-like in section. Di. Haldeman. 1840). Adams. Part I. Analyt. which suggests Planorbis. 1842.. tube-like duct: Planorbula. Later authors (Wenz. Limn.

For group of Planorhis armigerus Say May 1902. Biol. Con. Phniorbiila Dall. As genus 1899. C. Proc. Fresh-water Moll. — Recent and Fossil No 173 Ann. Planorbula F. 35. p. As subgenus of Segmentina Planorbula Germain. 1934. with a small tubercular lamella below. As genus Planorbula F. 213 (non Fleming). but includes Planorbula of Haldeman. Type Planorbis armigerus Say.. As genus Planorbula Thiele. p. 59. p. Radula figured. ultradextral.\ldeman. Rec. 1930. Zool. p. 97. IX. As subgenus of 1905. Ind. Planorbula Fischer. XVII. Baker. p. Wis. p.. 8). Planorbula 1886. slowly increasing in diameter. p. 25-27). As genus Planorbula Lindholm.. p. Univ. VI. Morrison. N. Form of pro. 14. obliquely transverse. Baker.. Ill. Heft 6. Type Planorbis armigerus Say. As genus Planorbula F. Phil. 1912. Papers Mus. Proc. Bull. 1918. Notes correct date of publication. 43. 1928. Soc. 58 year. Say. Monotype Planorbis armigerus Say. Planorbula Walker. Acad. Sci. p. p. and Subgenera 1870. placed as follows: a large parietal lamella. Cab. Handbuch. 509. H. cit. Inst. jaw and radula figured. figs. Aperture with six lamellae situated well within.. Proc. 1884. Type Planorbula armigera (Say).^l.. lips more or less thickened by a varix. 131. Man. 166. Phil. Type Segmentina armigera (Say). p. Wis. Genera.. Genitalia. 1934. 54. As genus Planorbula H. Mich.. equal Tropicorbis in part. Occ. with flatly rounded whorls above and a wide depression below. at the left end of the parietal lamella.. fiir Mollusk. Oct. 352. p.. 107. 210... Doubts that Planorbula of America is found in Egypt Segmentina (part) Hannib. As subgenus of Segmentina Planorbula Pallary. 1927.. of few closely coiled whorls. de Conch. 258. Tryon. & S. Type Planorbis armigerus Say. Part II. N.. X^o. p. p. 1926. Hald. Mus. Type designated by Pilsbry Planorhis armigerus Say Planorbula H. As subgenus of Planorbis p. Alaska Moll. Mem. Mus. LIV. No. Planorhis armigerus Say. As subgenus of Segmentimi Segmentina Tryon. Thinks it should be separate genus Planorbula Pilsbry and Bequaert. Nat. Type Planorbis armigerus Say. Baker. loc. Mon.state and presence of penial gland indicated Planorbulina Pilsbry. As genus Planorbula Pilsbry. Y.. fig. Zool. Planorbulina M. Amer. XXII. the body whorl usually carinate in the middle. somewhat sigmoid. 203.. Miscel... 6. Moll. p.. XXI. four palatal lamellae (plate 76. a large lower palatal (3) which is . Small. Univ. Conch.. 1931. Sci. S. B. Conch. N. p. 400. 104. Typographical error of von Martens for Planorbula. 154.. LIII. p. 1923.\rtexs. Arts. Type Planorhis armigerus type cited but 1870. 86. Shell and radula. Type Planorhis armigerus Say. 1883. Moll.Subfamilies. Acad. Sci.. p. 1909. 54. Cent. Egypt. Trans. p. Type Planorbis armigerus Say. 1840 Subgenus Type by PLANORBULA SS.. I. p. London. H. a large slightly transverse basal (4). listed. pp. Mich. Amer. X'at. Papers Mus. Segmentina Planorbula Dall. Subgenus of Segmentina Planorbula Clessin. original designation Planorbis arjnigerus Say Shell (plate 79. As subgenus of Segmentina Planorbula Pilsbry and Ferriss. Acad. Baker. Monotype Planorbis armigerus Say. 1940. C. 300. Clerical error for Planorbula. A group mixture. Lye. Mai. 179. C. and Lett. 480. Type Planorbis armigerus Say. Chicago Area. As genus. 1926. 1906. Archiv. 65. Nautilus. 353. X. 2 Teil.

5 mm. ANATOMICAL CHARACTERISTICS PLATE 1 37 (Planorbula nnnigcra (Say) (fig. the foot with lighter edges. the sides of the body lighter. is almost black. There are about thirty rows of diverticula in the i:)rostate.5 mm. There is only a slight constriction between the preputium and the vergic sac. the walls of which have many vertical folds (fig. but a shallow channel. also pointed upward. The penial complex (fig. ) GENITALIA. Male Organs nnn. 9) has a strikingly jiyriform preputium. The prostate (PRS) is about 1. fleshy muscular ring or diaphragm between the ring and the body wall of the prei)utium . wide and composed length and less than 0. rounded fold or ridge across the upper part of the preputial sac to the right pilaster. connects the penial gland with a circular space bordering externally the high. five or six smaller and shorter diverticula. the black eyes are placed at the inner bases of the tentacles. the old set appearing to be absorbed before the new one is formed as the shell increases through growth. the preputium has two heavy vertical pilasters (PL) the left one sending a transverse. the top of the head is wine-colored or reddish. New Hampshire. the prostate seen to be composed of a long diverticulum to which are attached. It is sometimes carried flat. The vergic sac is cylindrical in form. The tentacles are long and tapering. There is one heavy retractor muscle (RM) attached to the summit of the preputium near the vergic sac. The preputium is less than 1 mm. GF). in height. at an angle of about forty-five degrees. In section (figs. several of which may be branched two to four times. in diameter at its greatest width. The velar area is very wide and extends beyond the foot when the animal is in motion. The seminal vesicle (SV) is about The (about 0. Only one set of lamellae occurs in each shell although the lamellae are found at a very early period in the age of the individual. 10). Internally (fig. 1. The mantle is blotched with dark gray dots. about 0. thick. 11). basin-shaped penial gland (GL). There is no external gland duct. Animal (plate 70. fig. 2). open above (DC).174 The Molluscan Family Planorbidae always directed upward. The penial complex of another six^cimen shows a more swollen preputium and a wider vergic sac (fig. The vas deferens is of smaller diameter than the sperm duct. The shell is carried tilted to the left. narrowing anteriorly to a round tube of small diameter. a blackish band borders the head and neck above the foot and velar area. thick.5 nnn. long and 0. The stomach region is pigmented. blackish in color with a light zone near the head. long and a trifle more than 0.5 mm. The living animal of Planorbula jenksii from Lake Wentworth.5 or follicles. There is a rounded. Three wide supporting muscles are attached to the upper and middle area of the preputium. diameter under is of a cluster of small glands sperm duct (SPD) is roundly enlarged at its distal end long and wide) tapering anteriorly to less than half this the prostate. 6. long. on one side only. the left side lowermost. 4). with a shallow cup. and is 6 nnn. The liver and ovotestis are yellowish. There is no external penial gland duct.3 mm. in diameter and as long as the preputium. and a suprapalatal (1) which is usually in a transverse position. in is mm. 11). This lamella may be vestigial or absent in some forms of Planorbula. for the most part about half this diameter. an upper palatal above and smaller than the lower palatal (2).

long and 0.8 mm. DA. The albumen gland (AL) is large. crop. 153) is incorrect in several particulars. The ureter is sharj^ly recurved and is directed upward into the mantle cavity. passes backward to the liver. especially in the form of the prostate and the seminal vesicle. narrow duct (2 mm. fig. and Subgenera — Recent and Fossil 175 (fig. and is folded for the passage of the intestine. (fig. and a longer. 356. AL). The pseudobranch (plate 37. near middle).r. There Respiratory and Renal Systems. The genitalia of Planorbula jenksii are in most respects like those of armigera. 11. the kidney is seen to be much flattened. The uterus (U) is about 2 mm. Genera. OT) occupies a little more tlian one whorl and is composed of long. fig. 3. Female Organs (SD) are about 1. 3) is long and narrow (5. D). 154) is more nearly correct. There is a conspicuous blind sac or caecum. long) between the seminal vesicle and the oviduct. except those by F. ending in the anus over the pseudobranch. where there is a large carrefour (CF) which receives the long duct from the albumen gland (fig. The nidamental gland is slightly wider than the uterus and is located beneath the prostate. long and is of the same diameter as the sperm duct. In section (fig. with the lumen twice as wide as high and with a vein at each end. The oviduct (OD) is over 2 mm. The buccal sac is of 5. The ridge on the pseudobranch is heavy and when bent over (as shown in the figure) causes the pseudobranch to appear folded. Baker (1928. is rounded and has a ridge extending down the middle which connects with a crest-like ridge on the rectum (R). but does not show the details of the penial gland. wide the spermathecal duct entering it rather high. long. The intestine makes a loop around the stomach. The pseudobranch of jenksii is like that of armigera. long. Both jenksii and armigera have the same type of digestive system. makes another loop and runs forward to the rectum. The spermatheca (S) and spermathecal duct long. 12).3 mm. and pyloris) are similar to those in Carinifex and Parapholy. 4). ranged center. long. 154). p.3 mm. 13. Digestive System.Subfamilies. club-shaped diverticula areach pair bending to right or left or placed in the a very short free ovisperm duct between the ovisperm duct and the seminal vesicle. 8). somewhat triangular in shape. The vagina Hermaphrodite Organs. The verge (V) is sac-like. The kidney (plate 45. There is no ridge on the kidney. No figures of the genitalia of Planorbula have been published previously. in diameter. The figure of the entire genitalia (1928. There is a large pneumostome (PS). The diagrammatic figure of a secion of the penial complex (1928. The digestive organs (narrow stomach. 4. 1 mm. 153. It is somewhat convoluted at the upper end. The pericardium is about 1 mm. fig. about 1 mm. in pairs (fig. figs. the duct and gland of equal length and the than twice the diameter of the duct. The sperm canal extends through the center of the verge and has a central outlet bordered by a small penial papilla (fig.5 mm. 7) is . A few vesicles extend down a part of the ovisperm duct from the seminal vesicle. long and is but little wider than the vagina. The ovotestis (fig. P) . fig. gland swollen to more (VG) is 0. The prostate has about twenty-five rows of diverticula while in armigera there are thirty rows. wide at the pericardium). about 1 mm. and tapers to a point at its lower end (fig. C.

Radula Data for Pkmorbula Species . The marginal teeth (12-19) are long and narrow with cusps at the lower end. 2) is slightly arched. The jaws of jenksii (fig. The radula of jenksii has the same form of teeth. The superior jaw of armigera (plate 50. 3) are the same as those of armigera. The side jaws are about as long as the width of the superior jaw and are shaped as in Parapholyx. equal cusps (14-16). vertically striated on the anterior face. the cusps sharp. spade-like cusps of the same shape. fig. The endocone at first is split into two cusps about as long as the mesocone (12-13) they then become split into four small. The salivary glands extend half their length behind the buccal sac and form a loop. 1).176 The MoUuscan Family Planorhidae the same form as that of Carinifex. Radula of armigera (plate 65. fig. about four times as wide as high and finely. The center tooth is slightly higher than wide with a bicuspid reflection which does not reach the lower margin of the base of attachment. The lateral teeth (1-8) are squarish with three long. . The intermediate teeth (9-11) have the ectocone split two or three times. The outer marginal teeth are shorter than the earlier marginals but have the same form (19). The ectoconic margin has three to four small cusps.

fig. therefore. 105. and the prostate with small. T3-pe Planorbis icheatlcyi Lea.. the lower palatal lamella within the aperture very long and curved upward as far as the upper palatal lamella. Species Considered as Valid. p. Zool. In 1905 . have considered Haldemanina an absolute synonym of Planorbula. Subgenus Type by 1905. The lower palatal lamella is about twice as long as in armigera and the upper part is bent upward almost at right angles to the transverse lower part. 8.. XXI. generally. 47). the whole lamella being bent like an Australian boomerang (see plate 76.^lker. of Animal. Type Holdemaniua Dall. Pub. 480. Xot at present known outside of the state Alabama. branched diverticula on one side of a main diverticulum. Alaska Planorbis wheatleyi Lea. armigera). tyi)e of this subgenus. fig. The animal is unknown anatomically. with but which it agrees in general in the form of prostate and penial gland. Mich. wheatleyi. HALDEMANINA Moll. Rec. as suggested by Pilsbry in 1934 (p. 6. Handbuch.. Suhjamilies. These genera and their relatives constitute a subfamily Planorbulinae. Genera. as this is found in several species of Planorbula. but the difference in the lower palatal lamella is persistent in all specimens of wheatleyi here examined and is not found in any other species or race of Planorbula. As synonym of Planorbula Shell (plate 79. Considered synonym of Planorbula Haldemanina Thiele. 181. p. is disposed to accept Haldemanina as a subgroup under Planorbida. and Subgenera — Recent and Fossil 177 while in Planorbida this lamelhi points iiiiward.. (p. so that this end is on a line with the upper palatal lamella. Teil 2. No particular characteristics different from Planorbis armigera appear in his diagnosis. The chief characteristics of the genus Planorbula are the ovate or rounded penial gland without an external duct but with a canal-like duct within the preputium. 1931.' Upon examining the lamellae of wheatleyi.. Remarks. p. Ind. 28-30). Synonym of Planorbula Haldemanina Germ. the only species ascribed to it. Planorbula (Haldemanina) wheatleyi is (Lea). Segmentina is the type genus of the subfamily Segmentininae.^in. As in Planorbula but with the aperture notably thickened. Type Pkmorbis wheatleyi Lea. The writer. Mus. Mus. except the statement 'lamellae complex. T3'pe Pla7iorbis wheatleyi Lea. p. Its nearest relative is Proinenetus. the complexity is at once seen. 97). 9. Authors. 1905 original designation Planorbis wheatleyi Lea 1918. The thickening of the lip has no special significance. As section of subgenus Planorbula Haldeman Haldemanina W. the pHcae are always present but in Tropicorbis they may be absorbed late in life and the aperture become edentulous (see remarks under Tropicorbis) Planorbula has also been subordinated to Segmentina as a subgenus it has no relationship to that genus. Miscel. Ball. the internal organs being of a totally different character. 97. In Planorbula. Xo. figs. Dall instituted a section Haldemanina to contain the Planorbis wheatleyi of Lea. Geographical Distribution. 1923. Univ.

Wis. The sperm duct (SPD) is about 1. The seminal vesicle (SV) is about nearly three times the diameter of the ovisperm duct and has small diverticula or vesicles. 1935 1928. long and about one third the diameter of the sperm duct. . Color blackish. Type Planorbis exacuous Say. the preputium simply narrowing as it joins the vergic sac. Baker. There is a wide projection at the end of the gland in which there is an open groove (DC) or channel connecting the gland with the muscular ring or diaphragm (D) and separating the upper from the lower cavity. B. C. outer lip thin. exacuous megas) in which the three supplementary diverticula are each branched three times at the extremity. The vas deferens enlarges only slightly as it enters the vergic sac (VD). projecting from one side of the organ.5 mm. long. The verge (V) is long and narrow. the pyriform preputium occupying half of the length. ) GENITALIA. the much narrower vergic sac the other half. The i)enial complex (plate 41. Genitalia of Planorbis exacuous Say Promenetus F. 2) is 3 mm. I. Baker. right side flat. the pilasters separated to show the ovate cup of the gland and the glandular folds lining the walls of the cup.. figs. long and is composed of thirteen multiple diverticula. 5) the prostate is seen to have four supplementary diverticula projecting from the side which lies next to the oviduct.3 mm. and A. is 10). Adams). 360. Foot very short and rounded. in diameter and appears sausage-shaped. 3) and protruding from the diaphragm into the preputial cavity. p. long and 0. carinated or rounded. fig. 3). which. aperture wider than high. usually filling the vergic sac (plate 41. A longitudinal section of the gland (plate 42. the preputium has two heavy pilasters (PL) to the upper part of the left one of which the penial gland is attached (GL). ANATOMICAL CHARACTERISTICS PLATE 41 {Promenetus exacuous (Say) (fig. Say Menetus F. Fresh-water Moll. Animal. As new group original designation Planorbis exacuous Shell (plate 79. External features similar to those of Planorbula. The vas deferens (VD) is about 7 mm. XLIX. C. a penial complex is shown in which the preputium is much swollen by the enlarged penial gland. Internally (fig. 11. 1935. 7) shows the arrangement of the vertical folds within the gland. itself. plate 42 (P. The penial complex of variety mcgas docs not differ from that of exacuous. pp. 19-21). many 2 long. In fig. In section (fig.\ker. 1. 3 on plate 42. 48. 362 (non H. fig. the penial gland is shown from above. left side with broad spire depression exhibiting all of the whorls. The gland appears sac-like as seen from the side and is bent crosswise in the cavity of the preputium. also exhibiting all of the whorls. In fig. There is no sharp constriction between the upper and lower sacs. Nautilus.5 mm. Male Organs mm. An unusual form of prostate is shown in fig.178 The Molluscan Family Planorbidae Genus Type by PROMENETUS F. about as long as the diameter of the seminal vesicle. There is one narrow retractor muscle (R]M) and a wide group of three to five sujiporting muscles (SAI) forming a lattice-like pattern. The prostate (PRS) is 1. fig. Ultradextral Avith a small number of whorls rapidly increasing in diameter. C. also diminishes greatly in diameter beneath the prostate. 357.

long) and is a small tube. A specimen from Winnebago Lake had the verge and penial. 7) is about 2 mm. 6) shows five suppletermination of the main divera narrow tube about 3 mm. Small vesicles project from the ovisperm duct for a long distance (over 1 mm. Promenetus innhilicatellus. 9). long. A related species. ovisperm duct.Subfcunilies. long. shows some differences and the anatomy of this species is shown on plate 43. 1) and is bordered on both sides by several more or less sharply pointed gland-like protuberances. and albumen gland duct is shown in fig. The vagina (VG) is wide and a trifle more than 1 mm. 4) is about 1. and Subgenera — Recent and Fossil 179 In fig. The a little ovisperm duct (free portion) extends about 1 mm. In this specimen. fig. 6). on each side of the seminal vesicle and is without glandular projections or swellings. The vergic sac is not much narrower than the preputium. bulging in the center.5 mm. Hermaphrodite Organs. (plate 41. long and half as wide. with the penial gland (GLj extended crosswise of the preputial cavity. 8) and one near the posterior end (fig.5 mm. 10. In section (fig. it is seen to be composed of twin. A circular ridge near the end of the verge was observed in this specimen which did not appear in others examined. the two parts together measuring about 2 mm. A section about midway of its length (fig. The duct of the albumen gland is quite long and folded in an S form. six including the is The vas deferens (YD) '*>A* liiflLIBRAR . The sperm duct (SPD) is short. plate 42). 4). The uterus and oviduct combined are about 4 mm. 12. plate 42. 13. The seminal vesicle is about twice the diameter of the ovisperm duct (fig. long and the preputium and the vergic sac are of about equal length. Genera. The nidamental gland (XG) is about 1. the gland was somewhat contracted but showed well the trough-like canal or channel connecting the gland with the circular canal near the diaphragm. club-shaped diverticula. fig. The jirostate (PRS) is about 1 mm. 91. the latter having a swelling or ridge at the upper part indicating the position of the diaphragm. with a wide duct (SD) about as long as the spermatheca. The albumen gland (plate 41. 4. the verge is shown in this condition. one near the middle of the ovotestis (fig. The spermatheca (S) is irregularly ovate. fig.) below the seminal vesicle. in length. long. fig. The ovotestis OT. both glands entering the ovisperm duct. plate 42. gland extended from the male opening and lying on the neck of the animal (plate 41. Sections of variety megas are shown on plate 42. the side ridges or bands of muscle connecting the two pilasters with this canal are well shown. long and is composed of twenty diverticula which appear very long and narrow when the prostate is in its natural position lying against the uterus. The end of the verge of variety megas is shown in fig. The uterus (U) is the same width as the vagina and narrows only at the upper part of the oviduct (OD). The penial complex (fig. 10). carrefour. 10) occupies more than one whorl (fig. and narrow (less than 1 mm. oviduct. formerly known as Gyraidiis umbilicatiis. plate 42 (of variety megas). The relationship of the sperm duct. indicating the centrally located opening of the sperm canal with a small jienial papilla placed just below and to one side of the opening. There is one narrow retractor muscle (R^I) attached to the upper part of the preputium and a wide band of supporting muscle (S]\I) attached to the middle portion of mentary diverticula or ticulum. Female Organs (plate 41. In a specimen of variety tnegas (fig.

British Columbia (plate 42. There is no decided ridge on any specimen examined and the ridge on the rectum is not like that in Helisomatinae. The retractor muscle is shown with an attachment to both the preputium and the vergic sac. there is a short spermatheca (S) with a wide duct twice as long as the spermatheca (SD). fig. The ureter is a long tube and turns at right angles to the long diameter of the kidney. the penial complex is like that of exacuous (fig. In section (fig. In a specimen of variety megas from Paul Lake. In these examples. The albumen gland is somewhat quadrangular in form and is composed of large vesicles (fig. 157 on p. (plate 46. The verge (V) is short and wide. The oviduct (OD) is longer than in exacuous. many of the diverticula were gravid (plate 43.180 The Molluscan Family Planorbidae the preputium. fig. The pericardium is less than 1 mm.5 mm. the pseudobranch is rounded in one specimen and folded in another (plate 43. Alberta (plate 42. and there is a smooth ridge extending over a part of the rectum and forming a thickened border to the pseudobranch. fig. The kidney of umbilicatellus is of the same shape and general nature as The kidney 0. with a central opening of the sperm canal bordered by a distinct penial papilla (fig. Wisconsin (plate 41. 1). 3) of exacuous is 3. fig. 7. fig. the ovotestis is composed of paired In the specimens examined. (3T. The penial complex of Menetus e. had a single retractor without two branches for the vergic sac and the preputium. p. The pseudobranch appears somewhat variable in form. ( Respiratory and Renal Systems. wide and rounded below. fig. It is somewhat folded in the middle. 10.vacuou-s is figured by Baker 1928. long. . There is an open. 2). In specimens of e. 9). with the anal opening conspicuously placed on one side within the area of the i:)seudobranch. 2). It borders the thickened mantle margin on the inside. of exacuous and umbilicatellus. 8). rather long and narrow. fig.5 mm. There is no superposed ridge. 357. The uterus (U) is wide and short and there is a swollen nidamental gland (NG). A cross section of the kidney near the middle (plate 46. In the female system (fig. diverticula. 1). The relationship of the oviduct. Of the hermaphrodite organs. the anal region is outside of the area of the pseudobranch. A small supporting muscle may be present on the preputium below the retractor muscle. Internally. 8). 1. long and about wide. 6). sperm duct and ovisperm duct is shown in fig. the variation being largely due to the amount of expansion of the organ. This feature was not observed in any specimens more recently examined and this figure must be considered abnormal. A portion of the ovisperm duct between the seminal vesicle and the ovotestis is without gland-like swellings. 5).vacuous from Winnebago Lake. 12). The ovisperm duct differs from that of exacuous by having a number of small swellings on one side of the duct for half the length of the portion between the seminal vesicle and the oviduct. In umbilicatellus (plate 42. 3). All material studied. the pseudobranch is rounded. it is large and leaf-like. 11). In an example from Wainwright Park. There is a large penial gland (GL) which may be folded back as shown in fig. 4) shows an ovate lumen with a small vein on each side. fig. 154). The vagina (V) is wide but shorter than in exacuous. The same criticism extends to fig. it is like that of exacuous. canallike duct (see fig. 362 of the work mentioned. fig. it is fully expanded. fig.

It has been recorded from ]Maine west to Washington and Oregon and from Hudson Bay and Alaska south to New JMexico and Alabama. Promenetus are as follows. wide. collected by Prof. Baker (1). with the face vertically of the group (plate 50. Geographical Distribution. Swales Dr. arched. . Rawin . and low. C. 4. collected by F. with a rounded radular sac. looped. the entocone splitting into three to four small cusps and the outer margin of the teeth having two to three small cusps above the ectocone. umbilicateUus: Mott Lake. Nipissing. Baker (2) Specimens of exacuous from Wainwright Park. C. as in other members form to that of exacuous. there is a sharp median projection on the lower edge of the superior jaw. Alberta Vermilion Lake. Baker Dr. cracuou-^: Winnebago Lake. C. tricuspid. the mesocone longer and larger than the entocone and ectocone. Subfamilies. Alberta Wainwright Park. Swales (8) North Star Lake. two of each having been examined: catellus (fig. collected by F. the entocone splitting into two long. Alberta. . ^Marginal teeth (11-15) long and narrow. D. Swales (4) exacuous megas: Paul Lake. spade-shaped cusps not reaching the lower border of the base of attachment. British Columbia. Baker (4) Lake Mott Lake. The lateral teeth (1-6) are squarish. Ontario. Alberta. The stomach is elongated with the gizzard longer than wide and with a rounded pyloris (plate 48. figs. Minnesota. Swales F. received from the Canadian National Museum (5) Wainwright Park. were heavily infested with cercariae. 15). C. the two wide. Vermilion Lake. The genus Promenetus has a wide distri- bution. In several specimens both of these organs were almost obliterated. . sharp cusps and one or two small cusps appearing on the outer margin of the tooth above the ectocone (9-10). The side jaws are as in other species of the subfamily. A single species {i7nus) is known from Bermuda. Figures in parenthesis indicate number of specimens studied. . The buccal sac is elongated. Digestive System. collected by F. Genera. Intermediate teeth begin on the seventh tooth. C. extends backward to the liver and then makes another loop and runs forward to the rectum. there is only a rounded bulge at this point. The salivary glands are narrow. Wisconsin. S. son (2). Minnesota F. There is a long blind sac or caecum (BS). All cusps are sharply pointed. Alberta. In exacuous. and as long as the buccal sac. 2) is similar in The radula data for to four specimens Collector Species Locality Rows 110-112 115 138-140 140 Winnebago Lake. Wainwright Park. The intestine (IN) makes a tight loop around the stomach. 5). Wisconsin Wainwright Park. The outer marginals are almost vestigial. and Subgenera — Recent and Fossil 181 is that of exacuous. fig. fig. Baker The material examined for the anatomical data described and figured the preceding pages is listed below. but the ureter bends backward at a sharper angle and directed upward into the mantle cavity. more abundant in the ovotestis and liver. The radula of exacuous has a squarish center tooth with expanded lower margins. collected by Dr. 8. The sixth lateral has a small cusp above the ectocone. In umbilicatellus (plate 50.. collected by Dr. The radula of umbilisuperior jaw is The striated. exacuous). Minnesota.

188. XVII. Ind. Mon..\ll in 1870. Type Planorbis opercularis Gould. The form of the penial complex is also more like that of Planorbula than like this organ in Menetus. 1885. ZooL. which is not Anisus. Menetus W. p. opercularis (Gould). As subgenus Me7ietus D. Planorbis opercularis Gould. Its gland resembles Planorbula in the lack of a distinct duct and in the opening of the gland which is lengthwise instead of at the end of a cylindrical sac-like gland. p. Col. 1870. Fossil. Menetus Tryon. 1869. 206. 82. 12. 156. As subgenus of Planorbis Menetus F. 1855 Type designated by D. 1886. or von Martens. Bixney. Wis. Sci. II. 203. A. T3'pe Planorbis opercularis Gould. Univ. No. As genus Menetus D. Genotype Hippeutis (Menetus) of the . XXI. Remarks.. Five species. Type designation. exacuous {== exacutus) As subgenus of Planorbis . pp. Fresh-water Moll. exacutus.. p. 360. p. Trans. Rec. The discovery that the species long known as Gyraulus umbilicatellus is a Menetus closely related to exaciious (Baker. differs so radically from Menetus in the form of the penial gland that it is deemed necessary to raise the group to generic rank. p.. pp. Westerlund. Miscel. as in typical Menetus and its subgenus Micromenetus.. Haid. P. Arts and Lett. Type Planorbis opercularis Gould. 33. Tvpe Planorbis opercularis Gould.. Anatomy. p. pp. and A. 1929. Wis. C. As genus Menetus F. Menetus Walker. Pars 22. Conch. L. H. Cab. Univ. The vas deferens in typical Menetus enlarges as it enters the vergic sac to fprm an einphallus. and FW. As subgenus of Planorbis . pp. MENETUS H. 1650. Ann. and A. pp. Mus. Xo type cited but inckides both Planorbis opercularis and P. N. Type Planorbis opercularis Gould. Few Menetus. Pub. 1899) 1865. 1870. As genus Menetus J. N. 140-141. IX. 125. Menetus Germ. In Promenetus and Planorbula. Section of subgenus Hippeutis Agassiz 1923. Mich.. 1884. p. Sh. In other respects. No type cited but includes Planorbis opercularis and P. Type Planorbis opercularis Gould. Includes several unrelated species among which is Planorbis opercularis Gould. Con. C. I. Promenetus exacuous exacuous (Say) Promenetus rubeUus (Sterki) Promenetus exacuous Jtiegas (Dall) Promenetus hudsonicus (Pilsbry) Promenetus umbilicatellus (Cockerell) Promenetus imus (Vanatta) Geological Distribution. Menetus Wenz. They might belong in the genus 1926. Anatomy. G. 8. C. Subgenus of genus Hippeutis 1923. (Non Menetus of Chenu.^ll. N. are attributable to Promenetus. p. 1905. Genera of Rec. Tryon. Genus 1855. Baker. Mus. there is no such enlargement. 86. Alaska Moll. Xo. Studies. Baker. Pliocene to Recent fauna. XXII. first described as a group of the genus Menetus. 1928. Adams...species listed are true members of the genus found in Europe. 1883.. Fischer. II.\ll. XVH. Henderson. Moll. Y.. 2. Section of subgenus Hippeutis Agassiz 1918.. As subgenus Menetus Clessix. Syst. Promenetus. 262. Lye. in one of which two races are recognized. Acad. 1935.182 The Molluscan Family Planorhidae Species Considered as Valid. Cat. 46) was a surprise and shows that the group Promenetus includes species with both rounded and sharply carinated whorls. 94. Adams. the genus Promenetus conforms to the general characteristics of the subfamily. Planorbis opercularis Gould Menetus H. 6. 351.

1). or two-thirds of its length.Subfamilies. Type Planorbis opercularis Gould. 8). fig. 14). Acad. 8). ANATOMICAL CHARACTERISTICS PLATES 38 AND 39 Organs. The vas deferens is 3 mm. ultradextral. about 0. Menctus Pilsbry. Male Female Organs (fig. When expanded (fig. — Recent and Fossil As 86. p. The spermatheca (S) is ovate. There is a large . 1934. long and has a very narrow duct (SD) almost 1 mm. the prostate shows several small diverticula projecting from a single.aker. 2). 3). wide and 1 mm. left side with the spire whorls deeply immersed. B. long. long. C. and Subgenera 1931.5 mm. The vesicle proper has a few wide lobes. As genus Type Planorbis operopercularis 47. Genera. [Menetus opercularis. The oviduct (CD) is very short (about 0. or all may be single. Prostate (PRS) about 0. long. The cup cavity is provided with vertical folds.. Anatomy. 481. The foot is comparatively short. there is a short penial gland which is doubled over in the normal position (GL). Small. as in other Planorbidae. There is a slight constriction between the preputium and the vergic sac (VS). narrowed at the male genital opening. p. occupies 1 mm. about half the diameter of the sperm duct. the body whorl of large diameter compared wdth the inner whorls. Handbuch. this gland is trumpet-shaped. 1).3 mm. Sperm duct (SPD) about 1 mm. 22-24). Seminal vesicle (SV) about 0. XLIX. type of the genus. plate 38. 10) is elongated and sac-like. one or more of which may be divided two or three times at the end (as in figs. As genus Menetus F. sets of supporting GENITALIA. long. 2. and slightly enlarged as it enters the vergic sac. The verge (V) is elongated and tapering and normally as long as the vergic sac. Sci. vertical diverticulum. which enters the short and narrow vagina (VG). Animal. There are four muscles (SM) attached to the preputium. Type Planorbis Subgenus MENETUS SS.5 mm. there being usually five branches. of which the preputium.5 mm. T^])o Phowrbis opercularis Gould Shell (plate 79. 63. There is one narrow retractor muscle (RjNI) attached to the base of the vergic sac. cularis Gould. long. There is a dependent ring or diaphragm between the preputium and the vergic sac cavity (D in fig. . Gould. as in cooperi (fig. periphery or shoulder of body whorl more or less carinated aperture wide. Anatomy. The penial complex (fig. about 1. the preputium has two pilasters (PL). In section (fig. 6). tion under Hippeutis. more or less expanded. p. In general features the animal is like that of Planorhula. figs. long.4 nnn. of few rapidly enlarging whorls. The uterus (U) is about 0. Internally (fig. Color generally blackish. long with about a dozen main diverticula. Nat. outer lip usually thin. rather thick. the tentacles long and filiform. At the upper part of the preputium. the bell-like cup fiaring.2 mm. Proc. Phil.) and enters the ovisperm duct without notable decrease in size. Most of these are branched several times. subgenus of Anisus 1935. but the lower part (the ovisperm duct) is lined with small vesicles on the side. long and club-shaped. Nautilus. right side flat. 183 sec- Mcnclus Thiele. twice the diameter of the ovisperm duct and about the same length.

In section. 13 in plate 38. In section (fig. but the relative length of prcputium and vergic sac is the same. 7) is about 1 mm. sac-like penial gland (GL) with a short. fig. 14. the prostate is similar to that of opercularis (fig. Rounded swellings or vesicles extend down the ovisperm duct some distance. 11) but a lesser number in a specimen from British Columbia (plate 39. rounded vesicles (SO. long and about half as wide as long. shows some noteworthy differences (plate 39. There is a large. The albumen gland (fig. 5). plate 39. . the spermatheca (plate 39. callioglyptus. The ovisperm duct is not free at any portion of its length. The verge (plate 39. fig. is shown in fig. but distinct variety. The end of the verge is shown in fig. fig. In section (fig. The position of the albumen gland in relation to the other organs is shown in fig. 11. The seminal vesicle is twice the diameter of the ovisperm duct but the vesicles are longer and digitiform (SV). The other organs are similar to those of opercularis. 1. c. 7. 5. There are five sets of supporting muscles in a California specimen (plate 38. indicating the centrally located exit of the sperm canal. club-shaped diverticula. as in fig. 15. 7. fig. plate 38. plate 38) or it may be unbranched. fig. 1). plate 38 (British Columbia specimen). fig. there is a decided enlargement at the sunnnit of the vergic sac forming an epiphallus (EPI) as in some land snails. fig. 3. S) is much elongated and sac-like and the duct is narrow. widening somewhat as it enters the vagina. In specimens from British Columbia (plate 38. plate 39. The natural position of the penial complex beneath the female organs (vagina and uterus) is shown in fig. unbranched. This was only slightly developed in opercularis (plate 38. 10). the preputium has two large vertical pilasters (PL). callioglyptus and other members of the Planorbulinae. the gland shows a shallow cup (OC) at the outer end. Internally (plate 39. VD). fig. The prostate (PRS) has 15-17 diverticula (see also fig. 9). narrow. The ovotestis (OT) is composed of rather long. as is the case in cooperi and c. EPI. fig. In the female organs. 64. fig. the gland was bent upward and somewhat contracted. 12. fig. The penial gland of a Californian specimen of cooperi callioglyptus is shown in fig. This is better shown in a California specimen (fig. The penial complex (plate 38. 7. p. where the glands are in groups and branched. tube-like internal duct (DC) which enters a canal in the muscular ring or diaphragm (D). fig. 7a) figures the penial complex of Menetus cooperi callioglyptus (Vanatta) and this is similar to the figures in the . 9). There is but one retractor muscle (RAI). 9. In this specimen. plate 39. The vesicles are relatively large. V) of cooperi is wide with thick walls. 6). Hermaphrodite Organs. it shows a large cavity connecting with the enlarged vas deferens (epiphallus) which narrows notably at the entrance of the vas deferens (fig. ba(ilv infested with flukes. 1). California specimen) is rather broader than that of opercularis. Pilsbry (1934. A long duct runs through the fleshy part of the gland and connects with the short duct which emerges from the base of the gland. 10. A jicnial complex from a specimen collected in British Columbia. Menetus cooperi). lined with many vertical folds. they are seen to be in pairs. related. a British Columbia specimen) In section. 3.184 The MoUuscan Family Planorhidae carrefoiir (CF). but do not cover the entire length as in opcrcularis. The duct is about as long as the spermatheca. 8). but A is lined with small.

posterior part. 3). 4. This difference indicates that the two forms are distinct species. 5. The jaw is heavily. wide). 2 and 4 on plate 39 indicate the amount of variation that may take place in two individuals of the same species. in which the form of the penial gland and the size of the eiiiphallus differ. P. 2. and the pyloris rounded and diminishing to the intestine which forms a complete loop around the stomach. as figured on plate 38 (figs.Suhjamilies. The anal opening is placed at the left side of the pseudobranch. 1 in both plates 38 and 39) and the series appears simple. The verge is figured rather longer than was observed in specimens personally examined. A cross section of the kidney near the middle (fig. The lower part of the kidney is very close to the mantle margin. with a large. the pseudobranch is rounded. The side jaws are normal except . 2. but this organ is variable in form under different conditions. 7) is arched. makes another loop around the liver. In cooperi callioglyptus (plate 39. rounded extension of the radula sac at the lower. The pseudobranch of opercularis is wide. There is a large pneumostome. The salivary glands are about twice as long as the buccal sac and are much enlarged at the posterior end where they are attached. the anus opening at the lower edge (plate 38. A. and Subgenera — Recent and Fossil 185 present work. The diverticula in the natural position are hidden beneath the large diverticulum (as in fig. fig. The statement that the prostate alveoli (diverticula) are in a single series needs emendation. the gizzard rounded. The }x^culiar epiphallic enlargement of the vas deferens is shown in Pilsbry's figure. in a cross section they are seen to be composed of several brandies. the whole apparatus forming a loop. somewhat wider than high. coopcri. The buccal sac is pyriform. ending over the pseudobranch. The superior jaw of cooperi callioglyptus (plate 50. The stomach region is elongated. There is considerable difference between the genitalia of Menetus operculaj'is and M. This ridge is also found in opercularis. Genera. the ureter being in contact with it. extends backward in the body. The two forms of pseudobranch shown in figs. 7) of Menetus cooperi callioglyptus is short and very wide (about 2 mm. figs. narrow tube reflexed abruptly and directed upward into the mantle cavity. The ureter is a rather long. vertically striated. the vein leading to the pericardium much greater in diameter than the renal vein. There is a long and narrow blind sac or caecum. ovate. There is no superposed ridge. The oesojihagus is enlarged to twice its diameter posterior to the buccal sac. for while they ai^pear to be in a single series when viewed in position over the nidamental gland. but is not indicated in the figure. long and 1 mm. R). somewhat folded or lobed and the rectum extends through the middle. The pericardium is about 1 mm. figs. the oesophagus enlarged to form a crop. There is some slight variation between specimens from British Columbia and California. the ends attenuated. The stomach region is like that of Promenetus exacuous which is figured on plate 48. and with a high ridge extending o^'er the length of the rectum (R) and running down the midde of the pseudobranch as a low ridge. particularly in the penial complex. The kidney (plate 46. There is a central rounded bulge on the lower margin. long. 4). Digestive System. fig. 8) shows the lumen to be large. Respiratory and Renal Systems. 4. somewhat lobed. but in general the anatomical features are similar. and runs forward to the rectum. fig.

eleven specimens examined. Walton Clark. Puget Sound. Pacific coast region from to be confined to California. and over the stomach. California. The entocone. Only a few species are known. Vancouver Island. higher than wide. had parasites in liver. four specimens studied. as many as 200 larvae occurring in one specimen. but cooperi distributed over the nortliern California. Foster The material for the study of Menetus was received from the following sources: opercularis from Mountain Lake. The outer marginal teeth are very narrow and appear to be serrated with small cusps. Constitution. Hanna. albumen gland. the mesocone the longest. California. The parasites were most numerous over the stomach region. The lateral teeth (1-8) are sciuarish and tricuspid. British Columbia 144-145 Orcas Ishmd. Arthur Peake Dr. collected by Dr. D. Baker. 1859. Species Considered as Valid. Arthur Peake. the two wide. Quatsino Sound. The following species and races are at present known for the typical subgenus of Menetus. fig. D. Foster. a fossil . near San Francisco. The intermediate teeth (9-10) are narrower than the laterals and have one or two additional cusps above the ectocone. the cusps long and spade-shaped. Data for Radula Name cooperi callioglyptus cooperi Formula 20-1-20 20-1-20 Locality Rows Collector Quatsino. It is of the coast region. All organs were affected but the ovotestis and albumen gland were more often attacked. spade-shaped cusps not reaching the lower border of the base of attachment. from Orcas Island. In some specimens from both Washington and British Columbia all of the organs were obscured by a thick coating of mucus. The radula of Menetus cooperi callioglyptus (plate 67. cooperi callioglyptus. The marginal teeth (11-15) are very narrow. collected by Mr. Specimens of cooperi from Orcas Island. small mountain lake on Mt. collected by Mr. Washington. British Columbia. 1824. distinctively a genus Opercularis appears (=^ planulatus) is found from northern California northward to Vancouver Island. species. the entocone is split into from two to four small cusps and the ectocone splits into three to four small cusps. new name Menetus cooperi multilineatus (Vanatta) for planulatus Cooper. C. were largely infested with cercariae of trematode worms. G. mill pond at Crescent City. 3. H. Washington 155-161 Mr. fifteen specimens examined. Specimens of cooperi callioglyptus from Quatsino Sound. ovotestis. mesocone. and within this area is cjuite variable. The jaw of Menetus opercularis is similar. and ectocone become more nearly equal in length. Tyjiical Menetus is Vancouver Island south to The group does not extend far inland to the east. Geographical Distribution.186 The Molluscan Family Planorhidae that at the i^oint of attachment to the superior jaw they are bent backward more than usual. T. from British Columbia) has a large center tooth. The mesocone persists throughout the entire series of teeth and may always be identified. elongated. cooperi. Menetus opercularis (Gould) Menetus cooperi callioglyptus (Vanatta) Menetus cooperi F. Vancouver Island. received from Dr. not Menetus centervillensis (Tryon) planulatus Deshayes. T. Puget Sound. There are a number of local forms of limited distribution. two specimens examined. D.

Subfamilies. Animal. ANATOMICAL CHARACTERISTICS PLATE 40 GENITALIA. 390) includes in it species now referred to Tropicorbis. Male Organs tu. p. attaclied far back on the head. tentacles long and filiform. dilata- of a small number of relatively large vesicles placed on each side of the ovisperm duct. Thiele phices it under Anisus 1931. Foot short. The shell is carried at an angle of thirty to forty-five degrees or it may lie flat on the body of the animal. 12). is quite unlike either that of Planorbida or Promenetus. of few rapidly enlarging whorls right side flat or convex. Baker. however. C. enlarged somewhat at base. Seminal vesicle (SV) of M. Remarks. Typical Menetus is a very distinct genus characterized by noteworthy anatomical features. C. chief among which are the sac-like penial gland with its short. It is seldom carried erect as in Helisoma (plate 70. body whorl with a more or less well-developed carina. 13-15) Very small. Menetus has been subordinated to several groups as a section or subDall as a section under genus. aperture large. dilatatus New Subgenus Gould Shell ("plate 79. 9). wide. ultradextral. Pliocene to Recent fauna. Von Martens (1899. (1923. The penial gland. The combined free portion and that part beneath the prostate measure about 1 mm. in length. 360) and Pilsbry (1934. Anterior to the seminal vesicle. eyes on small swellings at inner base of tentacles. eight or nine.s (fig. two black lines extending down middle of head. bluntly rounded before and behind. bottom and sides of foot yellowish. narrow. Sperm duct (SPD) of small diameter. Prostate (PRS) with relatively few diverticula. In cross section. ( ) . and Subgenera — Recent and Fossil 187 Geological Distribution. the sperm duct being a trifle longer than the prostate portion. left side with inner whorls submerged by the body whorl. figs. Baker (1928. The anatomical figures herein published support this conclusion. F. 156). The ovotestis is also like that of Planorbula. the prostate shows the same multiplication of smaller diverticula on the under side as is present in Menetus opercidaris. Little is definitely the geological ancestry of this genus. The duct is also different The radula and jaw are like these organs in Promenetus and Planorbula. whole back dark. color mottled brown and cream. 351). though one was definitelv proposed bv Dall in 1870 (p. p. Genera. tentacles transparent. not much greater than that of the ovisperm duct. The prostate resembles that of Planorbula which has the same structural arrangement of diverticula. Subgenus MICROMENETUS Genotype Planorbis . The Adams brothers indicated no type. In an immature specimen of . fig. 64) have shown definitely that the group should rank as a genus. F. p. p. the ovisperm duct is bordered by a number of widely spaced vesicles for a distance three times made up the length of the seminal vesicle. lips thin. internal duct and the epiphallus-like enlargement of the vas deferens as it enters the vergic sac. p. usually placed just below the top of the right side. some. p. what dilated. 82). Germain as a subgenus of Planorbis known concerning . oval on base. 481 subgenus Hippentis (1905. Shape of foot varies during locomotion.

The uterus (U) is wide. 3). There is a carrefour which lies between the oviduct and the albumen gland (not shown in the figure). The vas deferens is a narrow tube ahiiost as long as the sperm duct. The cup containing the vertical folds is small and is placed at the end of the penial gland. which follows the wall of the preputium (DC) to the muscular ring or diaphragm separating the preputial sac from the vergic cavity. A single diverticulum filled with developing ova is shown in fig. 2) the gland was somewhat different. 10).5 mm. There is a fluted crest on the rectum (R) which. Internally duct.. The ovotestis (OT) is composed of relatively large. The oviduct (OD) is short and gradually narrows to meet the sperm duct. long. the gland being pushed upward. resembling a pipe or trumpet. There is one retractor muscle (RM) which is usually attached to the constriction between the preputium and the vergic sac (fig. Only a few rather large vesicles occur on the ovisperm duct. does not appear on the pseudobranch in the specimens examined. This is (fig. somewhat club-shaped diverticula arranged in pairs. There is a central duct running through the gland as in Menetus cooperi callioghjptus. In one (fig. 9) is about twice as long as of the foot in preserved specimens. shown to where the gland (GL) is a sac-like penial gland with a rather long better advantage in Menetus sampsoni (fig. V) is elongated. the seminal vesicle (fig. Respiratory and Renal Systems. the preputium (PR) is sac-like or elongate-pyriform and is about twice as long as the ovate vergic sac (VS). In sampsoni (fig. 12. The penial complex (fig. 1 differs from that organ in cooperi in being composed of four rounded swellings. 5. S) is short and sac-like and connected with the very wide vagina (VG) by a wide duct twice as long and half as wide as the spermatheca. The intestine makes a Hermaphrodite Organs. 11). The pseudobranch of dilatatus (plate few wide and extends below the margin somewhat folded on the side and in a Texas specimen {sampsoni. there is much elongated and gradually diminishes in diameter to the round duct. 12) is about 0. being more swollen when filled with ova ready for discharge. In another specimen (fig. In Menetus sampsoni. The diverticula vary in form. the vergic sac is longer and wider. The spermatheca (fig. The albumen gland (AL) is very large and wide (almost half is as wide as long) and is composed of large vesicles. 5). increasing to about twice the diameter of the vagina in the region of the prostate. 40. ) . 4) the preputium was much swollen and the retractor muscle was attached to the preputium some distance below the vergic sac. 6. as occurs so frequently in the genus Helisoma. 8) formed a hollow cylinder. There are several small supporting muscles on both sides of the preputium. This might have been abnormal. loop beneath the albumen gland (IN). 10. It is slightly enlarged near the vergic sac. The anal opening (A) is jilaced at the upper end of the pseudobranch. however. Female Organs. fig. The penial complex was observed in several different forms in the material examined. The verge (fig. The sperm canal has a central outlet. See also fig. the vergic sac was placed on the side of the preputium. In a Texas specimen (fig. figs. 7. It is . 188 The Molluscan Family Planorhidae cooperi. the prostate had but five rows of diverticula. where the still wider nidamental gland (NG) appears. ID. narrowing to a point at the lower end.

the entocone with three to four small subequal cusps. The pericardium is very wide. Intermediate teeth (8-10) becoming narrower and developing a split entocone and one or two small cusps above the ectocone.Subfamilies. about 1 mm. 5). the ectocone with several small cusps on the outer edge of the teeth. figs. not as wide as in typical Menetus. against the lower part of the kidney. wide. The radula (plate 67. and Subgenera — Recent and Fossil 189 long and 0. Lateral teeth 1-7) squarish. the mesocone longest. The kidney is wider than that of exacuous but narrower than that of cooperi or c. There is a slight bulging in the center of the lower cutting edge of the superior jaw in both dilatatus and sampsoni. The kidney (fig. The radula of sampsoni is practically of the same type as that of dilatatus (fig. all cusps sharp and spade-shaped. fig. 5). 5) is rather short. callioglyptus. Marginal teeth (11-14) long and narrow. (see plate 48. 6) are similar. ( . 4. but larger. 6) shows an oblong or ovate lumen with a small vein at each end. The side jaws are as long as the width of the superior jaw. The superior jaw (plate 50. dilatatus). fig. The digestive system is similar to that of exacuous The radula sac is essentially the same as in Menetus. The ureter is very long and is folded back tightly Digestive System. 9) of dilatatus is wide and low with vertically striated face. The marginal teeth become much smaller toward the edge of the membrane.3 mm. tricuspid. The jaws of sampsoni (fig. Center tooth squarish. A cross section near the middle of the kidney (fig. 4. There is no superposed ridge. Genera.

Discoidal. perhaps. Planorbijex Pilsbry. fig. A. 190 The MoUuscan Family Planorbidae races of the subgenus as follows: dilatatus (Gould) Species Considered as Valid. Paraplanorbis Hanna resembles Planorbijex in being small and keel. for the narrow umbilicus. 14. Genus Type by PLANORBIFEX Pilsbry. left side deeply umbilicate. The form of the shell is lenticular and there is usually a peripheral carina more or less well develoiied. with plate 40. vanvlecki Arnold. the last whorl more or less flattened. Species Considered as Valid. fig. 568.. its outline excised bv the preceding whorl. Sculpture of fine. Nat. with the last whorl carinate. this duct is short and enters the diaphragm directly without being attached to the wall of the preputium (plate 39. that of Menetus being 20-1-20 while in Micro- menetus it is 15-1-15. 1934 original designation Planorbis vanvlecki Arnold 1934. 7). Kettleman Hills. The fossil species Planorbijex vanvlecki Arnold. Basal part of the Tulare formaNorth dome. As subgenus of Menetus Pilsbry. the right side moderately convex. Genotype Planorbis Fossil 1935. The group here separated as Micromenetus differs from Menetus in the size of the shell which is always much smaller. 'Except . As far as examined the radulae of the two groups differ in formulae.) Menetus (Micromenetus) alabamerisis aims Pilsbry Menetus (Micromenetus) sylvanicus Pilsbry Menetus (Micromenetus) brogniartianus (Lea) Menetus (Micromenetus) buchanensis (Lea) Menetus (Micromenetus) sampsoni (Ancey) Menetus (Micromenetus) alabamensis alabamensis Pilsbry Menetus (Micromenetus) Vanatta uliginosus Geological Distribution. J. p. the periphery rounded. 86.. Remarks. a much larger planorb from Lake Titicaca. dilatatus dilatatus penndilatatus Menetus (Micromenetus. fig. Pliocene. Phil. The penial gland has a duct which is almost three times as long as the gland and is attached to the inner wall of the preputium for the greater part of its length (plate 40. 9).. hair-like striae. the receding left margin thickened within (Pilsbry) tion. Menetus (Planorbijex) vanvlecki Moll. Acad. typical Not exactly known but certainly from the Remarks. Non-Marine Shell (plate 81. but they appear constant. These are small differences. close. which has similar sculpture. California. Tlie pseudobranch in Micromenetus is also very long and narrow while in typical Menetus it is short and wide (compare jilate 38. fig. There are several distinct species and Micromenetus recognizable within the genus Menetus. Geological Horizon and Distribution. as genotype. Sci. p. 252. In typical Menetus. is the only species known for this genus. dextral in appearance. N. Henderson. Kings County. the absence of a peripheral and the thickened peristome. 6). Aperture strongly oblique. Pliocene to Recent fauna. Micromenetus differs from both Promenetus and Planorbula in the shape of the penial gland. Proc. 10). in diameter. none exceeding 4 mm. fig. this form resembles Platytaphius Pilsbry.

The species was referred by Dr. 1934. a very different shell. The small which. and is different from any known species of Menetus or Promenetus. p. but in carinate species of Menetus the keel is peripheral' (Pilsbry. 1934. As remarked under the genus Paraplanorbis. but the excised outline of the strongly oblique aperture. Genera. p. where the carina is peripheral or just below the margin of the left side. it appears better to regard Planorbifex as a separate genus related to Menetus rather than as a subgenus of Menetus. Cooper to Valvata virens. Planorbifex vanvlecki. sometimes it is scarcely noticeable. but Drepanotrema. it — Recent and Fossil 191 has a spire of closely coiled whorls like and umbilicate left side are like Menetus. The carina in the middle of the whorl of the right side is distinctive. as Arnold thought. varies in the amount of flattening of the base. and Subgenera narrowly iinibilicatc. and the thickened peristome. 568). is probably its nearest relative. . exclude a reference to the Valvatidae. 'The shell of the type species.Subfamilies. some of which have a size similar sculpture' (Pilsbry. It has certain resemblances to such forms as Valvata bicarinata Lea. 569).

ji. 1883. As genus Shell (plate 81. figs.3 mm. Type Choanomphalus maacki Gerst. half-moon-shaped superior jaw and two small lateral jaws. 1931. Type Choanomphalus maacki Gerst. Conch. pp. 1884. 232. Choanomphalus cryptomphalus has the formula 5-6-20-1-20-6-5 (63 teeth in row). Mollusken.. p. As genus Choanomphalus Lindholm. 190. St. and radula. the spire depressed but not flat. The shell resembles some species of Valvata in form.6 mm. N. 9) small. nine marginal teeth. p. Choanomphalus Germain. turbinate. IX. Sci. As genus Choanomphalus Westerlund. 63. p. The eggs 192 . L. Man. side (see Dybowski and Grachmalichi. Type Choanomphahis maacki Gerst.(4) (69 teeth in row) or sixteen lateral teeth. 508. 1875. Jour. Region Binnenconchylien. 527. The formula of Choanomphalus maacki is (4)9-5-16-1-16-5-9. Type Choanomphalus maacki Gerst. XXI. and four outer marginal teeth without distinct form. Conch. Dybowski. 160. V. 1909.. 1886.. five intermediate teeth. base widely umbilicated or with a small chink. Exped. p. The radula plate measures 1.. As genus Choanomphalus Clessin. Ann. In Choanomphalus valvatoides the formula is (3) -10-5-10-110-5-10. 1923 Characters those of the genus Choanomphalus Gerstfeldt Genus Type by 1859. Ser 7. Kosmos. 8. Petersburg. As genus 1923.(3) (57 teeth in row).. Choanomphalus Gerstfeldt. As genus Choanomphalus Dybowski and Grachmalichi. Mus. of few whorls rather rapidly increasing in diameter. p. aperture rounded. Die Gasteroi)oden Fauna Baikal-See. 52. Choanomphalus maacki Gerstfeldt 1870. As genus Choanomphalus Thiele. Type Choanomphalus maacki Gerstfeldt. Type Choanomphalus maacki Gerstfeldt. p. Rec. Zool. As genus Choanomphalus W. 93. Animal. Type Choanomphalus maacki Gerst. 7. pseudodextral. Similar to Plonorbis and genital openings on the left 870) . and.. Type Choanomphalus maacki Gerst. in length and 0. Type Choanomphalus maacki Gerst. Respiratory p. Wissens. Ind. 877. XXVII. 8. in width. p. in Korotneff. As genus Choanomphalus Dall. 1885. 1879.VIII. 358. jaw. Lye. p. N. S. The jaws are three in number with a large. Type Choanomphalus maacki Gerst. in tentacles. As genus Choanomphalus Tryon. Baikal-See. de Conch. As genus Choanomphalus Crosse and Fischer. p. Ergebn. 105. The smallest formula is borne by Choanomphalus schrencki which is 4-7-1-7-4 (23 teeth in row). 1925. Ueber Land imd Siisswasser-mollusken Siberiens.S. Handbuch. Fauna der palaarct. XX. and carries 175 rows of teeth. de Conch. Type Choanomphalus maacki Gerst. Syst. 353. XVII. IH. IV. Acad. 1859 original designation. CHOANOMPHALUS Gerstfeldt. pp. Cab. Y. In Choanomphalus bicarinatus the formula is 4-6-20-1-20-6-4 (61 teeth in row). GROUPS OF UNCERTAIN AFFINITIES Subfamily CHOANOMPHALINAE Germain. As genus Choanomphalus Fischer. Mem.. H. 482. are laid in capsules with four to five eggs in each capsule.

some twenty-five in number. Six genera and some twenty-three lesser groups are designated. Another species is found in Japan ( japonicus) Westerlund includes Europe. 835). of which fifteen are considered valid. The published information concerning the anatomical details is wholly insufficient for purposes of classification. It has been thought to be related to Choanomphalus by Sturany and other European conchologists. The Japanese species is placed in a separate section by Lindholm. The genus Choanoinphalus is a group of mollusks probably related to the Planorbidae but of uncertain status as to its proper position in the classification of the fresh-water pulmonates. on the other hand. Choanomphalodes. . Dall (1870) and Tryon (18841 placed the genus. 1. Crosse and Fischer in 1879 compared the genus with the American group Carinifex and suggested its group relationship with the fossil Valvata multiformis { = trochiiormis) The resemblance to Carinifex in both Choanomphalus and the Steinheim fossil itrochiformis) is superficial. Valvatoides. No mention is made of the cusps of the teeth. A. Geographical Distribution. pp. In 1909. It probably represents another peculiar form of Planorbidae and its anatomy should be examined. There seems to be great uniformity in the total number of teeth but some variation in the number of lateral teeth. principally between 150 and 350 meters. Type Choanomphalus amauronius Bourguignat 3. Siberia. from Lake Ochrida near the city of Ochrida. One species is found in the Angara River. between Greece an^l Turkey. in 1902 (p. Lindholm made a study of the Lake Baikal fauna and divided Choanomphalus into three subgenera. On pages 877 to 880. Choanomphalus not having the same characteristics of anatomy found in Carinifex and Paraphohjx. in the distribution of the genus. This range needs confirmation. Lake Baikal. that Segmentina may be related to Choanomphalus is obviously erroneous because that genus has been shown to be the type of a distinct subfamily not known to have any affinities with the Baikal Lake group. 871). 1927. Thessalia. Wladislaviidae. review all of the species of Choano))iphalus. Dybowski and Grachmalicki 1925. 120). The suggestion in the foregoing reference (p. 819-881 in an exhaustive monograph. is discussed.. Dj-bowski . ( ) . Westerlund. Germain (1923) places the group in a subfamily Choanomphalinae. type Choanomphalus japonicus Preston. Achoaiiomphalus Lindholm. Type Choanomphalus schrencki W. as noted below. in the subfamily Pompholiginae with Pompholyx {Paraphohjx) and Carrriifex. from two to ten meters. in European Turkey. W. thus maacki occurs at great depths. A form of Segmentina nitida is found in Lake Baikal and is given the name of angarensis by these two authors. placed Choanomphalus in the subfamily Planorbinae. Sulcifer Lindholm. but its relationship to Choanomphalus is questioned. p. occurs in comparatively shallow water. near Lake Baikal iamauronicus). Choanomphahis Sensu Stricto. The peculiar Planorbis paradoxus Sturany. .Groups of Uncertain Affinities 193 the outer marginals indistinct and uncountable (op. Dybowski and Grachmalicki present a new arrangement of Choanomphalus which is made a separate family. The different species occur at varying dei)ths. Type Choanomphalus maacki Gerstfeldt 2. cit. Remarks.

Valvatomphalus D. Genus Tj'pe POECILOSPIRA :\Iorch. p. Conch. Yoldi. and G. the carina placed at the upper side of the whorl in planorboid forms and forming a peripheral ridge in the forms with elevated spire. but in that case the name of Germain must be used because it was given at an earlier date than Wladislaviidae of Dybowski and Grachmalicki. 1929.. the last whorl rounded and in the same plane as the body whorl or greatly deflected. Type Valvata muUijormis Zeiten. {= Achomwmphalus Lindholm. These by Thiele (1931. Tvpe C. judging from the figures in Fischer's paper (1879. Type Valvota multiformis Zeiten Crosse and Fischer. As subgenus of Choanomphahis Shell (plate 81. ISQ^)^ Anomphalodes Tomlin. p. and group names is debatable. Type Choanomphahis bicarinatus Dybowski Anomphalus D. entire and . On this page Germain also questions the value of recognizing subgenera for the different species. particularly the form of the ovotestis. S. Until the details of the genitalia arc known. the group designation of Germain as a subfamily of Planorbidae should be retained. Type Choanomphahis maacki Gerst. are tabulated below. plate 4. 2. For the present. varying from fiat with depressed spire. As genus Poecilospira Tryon. it will be impossible to determine the true position of this group in the classification of the fresh water pulmonates. 1). 482) with the earlier names of Lindholm and the new names proposed by Tomlin for the preoccupied names. The plate of Dybowski and Grachmalicki monograph docs not show variation enough to warrant any such minute division. Cat. Type C. Dybowski Biangulatiis D. Umbilicus large and conspicuous. fig. Type Planorhis muUijormis (Zeiten). Choa7iomphaIus (SS). 1909). Aperture round or lunate. and the internal conditions of the penial complex. 4. The name Wladislaviidae can not be used because Germain in 1923 (p. to trochiform with greatly elevated spire. Type C. 6. 160. Cryptomphalus D. and S. p. discoidal.194 The Molluscan Family Planorbidae in 1909 are ignored. amauronicus Bourg. prostate. that when the anatomy is known the group might be found to be of family rank. 105. Whorls carinated. such as is common among many of our The value in of these generic species the North American species of Helisoma. as listed 1. {= Helicites trochiformis Stahl. and G. and G. 8-10) which are reproduced on my plate 81. 1909). and many of the figures suggest individual variation. The subgeneric names proposed by Lindholm six genera. and G. 3. (non Charpentier. 190) proposed the subfamily name Choanomphalinae two years earlier. Possibly the few groups and species enumerated by Lindholm can be recognized. and G. It is possible. XXVII. Dvbowski Platybasafis D. {= Sulci) er Lindholm. Conch. schrc7icki W. de Conch. 5. Jour. cryptoinphahis W. Just why all of the Lake Baikal species of Choanomphahis should not be included in the one genus is not apparent from a study of the shells and what little is known about the animals. 1853 by original designation Valimta muUijormis Zeiten 1853. Poecilospira 1884.. 1837)= Omphalocrypta Tomlin. of course. (non Meek and Worthen. Tvpe C anomphalus W. 1929. Dybowski . Dextral. figs. Ill. Poecilospira Morch. 1824) 1879.

Remarks. The planorbid fauna at Steinheim has become classical for studies of variation. a thick callus on the parietal wall and lip thickened in many specimens. for between 1824 and 1920 no less than forty-five names have been given to variations of this group. Perhaps Hyatt's studies in 1880 (pp. It is certainly not near Gyraidus and apparently does not belong in the subfamily Planorbinae. Morch. Germany. approaching Valvata in » 1 A form. 1-114. the forms grouped around multiformis (trochiformis) appear different from the Gyraulus-like shells. . pp. Hyatt refers all species to the genus Planorbis. Taf. p. 10-12). In a later paper (1920. and others shows that. 1601 includes all of the Steinheim fossils in the genus Gyrauhis. Steinheim near Heidenheim. recognized this difference and gave the assemblage the name of Poecilospira. There are some features in common between Poecilospira and Choanomphalus and its true position may be with the latter group. as suggested by Tryon in 1884. Gottschick. Horizon and Distribution. 155-216. He postulated a coldAvater fauna and a warm-water fauna. The work of Wenz also brings out clearly the fact that the fossils of this and nearby regions have been very much overnamed. This paper is well illustrated and shows the great variation in the group of shells known as Planorbis ))iultiformis. as we sometimes find in the Pleistocene faunas of the middle west. Gottschick reviews what is known concerning the fossils of this locality. It is not mentioned by Wenz (1923) in the list of fossils related to trochiformis. Also. careful study of the works of Hyatt.Groups of Uucertain Affinities 195 the outer separated from the body whorl. Wiirttenberg. it is not related to Carinifex or to any of the fossil relatives of this group. and apparently should be segregated in another group. plates 1-9 are the best known and show in large measure the great amount of variation which has taken place during the life of these mollusks while the Tertiary strata w^ere being formed in the old lake bed and on its shores. Wenz (1923. Later authors appear to have overlooked this name which does not appear in recent monographs or check lists. ^Miocene period. not only as individuals in a fauna. To what group Poecilospira is related is not definitely clear. but also in time between the earlier and later strata. while many of the so-called species appear to be referable to the genus Gyrauhis. in 1863.

leaf-shaped. XXII. phite 1-2.\ker. XXIV. The family is mostly exotic. 1931.\le Med. with few rapidly increasing whorls. 1934. Of medium size. but as one species previously referred to the Planorbidae has recently been placed in Bulinidae.. p. Proc. Rec. spire whorls flat. figs. No. Larambergue. 1933. Anatomy and family position. 1-12. p. Description of anatomj'. 33-35). Ind. pp. the inner whorls concealed by the last two whorls. sinistral. Type Planorbis exustus Desh. especially genitalia and radula. 1927. Oceania. No. p. Morph.and. More recently several authors have considered the group as of family rank. 86. p. 1921.\d. 199-219. 286-295. 1922. Jour. p. p.IX. space is given to a discussion of this species and the data for its inclusion in a family to which it was not previously believed related. As genus Indoplanorbis Axn. Iitdoplanorbis Ann. X'at.\sh.. 132). As genus Shell (plate 79. p.shad. 1. 'The animal is sinistral. Handbuch. 132). 1921. Planorbis exustus Pr. pp. Mus. 471. Japan. As genus Planorbis (PUmorbis) exustus Germ. the Ethiopian and Malagasy regions. 479. As genus 1921.\d.synonymy Indoplanorbis Anx. Zool. the body whorl large and greatly increasing in diameter toward the aperture. Ind. 1920. Acad. Rec.\. Ind. 112. As section of Planorbis Indoplanorbis F. the fluted or lobular })seudobranch and the shape of the ]ienial complex in the genitalia. XXV. Mus. Tyi)e Planorbis exustus Desh.. 578. p. the ^Mediterranean subregion as far east as Mesopotamia.. Account of anatomy. Celebes. Tvi)p Planorbis cxuslus 1923. Soc. LXIV. Rec. These are so different that they would appear to be characteristics of family rank. Animal (plate 21.. XXI. B. VIII. Type Planorbis exustus Deshayes. lips simple. and the Antilles (vide Pilsbry. 196 . pp. New Guinea. Phil. Its foot is relatively broad and short. As genus Indoplanorbis Rao. FAMILY BULINIDAE This group has ahnost universally been considered a subfamily of Planoris so treated by Pilsbry (1927. It Genus INDOPLAXORBIS and Aiinandale and Prashad. base with small umbilical opening. Jour. India. Remarks on taxonomic position Indoplanorbis exustus Larambergue. 1939). 2). Rec. aperture usually oblique.. Type Planorbis exustus Desh. Only one species occurs in the Antillcan region.. Indoplanorbis Pr.\ni). T^-pe Planorbis exustus Desh. Res. 5. XXII. Roc. 1939.\le and Pr. Bull. Mus. Mus. Type Platwrbis exustus Desh. distributed as far as known in Australia. 360. de France. The present work was prepared to include only the groups i)roperly belonging to the family Planorbidae. 514. Ind. Desh. 54. Two characteristic features of the group separate it from any known division of the Planorbidae.\nd. distinct from the Planorbidae (Germain.\sh. fig. depressed below the level of the body whorl. sharp. Ind. LV. C. pp. 1920 Deshayes Type by original designation. p. As genus Indoplanorbis Pilsbry.\le and Pk. broadly rounded in front and pointed behind. The shell is of Physoid shape but the radula is like that of Planorbis.ash. 1926. As genus Indoplanorbis Thible. 26-41.. Exhaustive account of variation and . Sci.

This condition is indicated in fig. 3) is made up of a short. ( . but it was of the shape figured in all but one of the specimens examined and the exception was regularly elliptical. long and nearly 1 nun. 1921. penial complex (fig. vertical. 1923. which position was not observed in any specimen examined. 9. the prostate lies against the uterus below the nidamental gland. which appears to be its natural position. narrow tube-like vergic sac (VS) which is five times the length of the preputium. 200. fig. long). The eyes lie at the inner base of the tentacles and are completely sessile. the portion near the ovisperm duct folded or coiled. p. In the figure of Larambergue (1939. the preputium contains two long. muscular pads or pilasters (]\IP) which extend the whole length of the preputial cavity. Internally (plate 21. be some variation in the shape of this organ. 215). 3). 8). p. See Rao.5 mm. the vergic sac is shown four times the length of the preputium. cylindrical preputium long) and a very long (17 mm. Both Rao and Larambergue describe the prostate as ellipitical in form. In Rao's figure (1923. 1). p. 10. In its natural position. 3. The mouth opens on the lower p. its low^r end broadly rounded. obviously. (3. Male Organs. wide and is composed of small lobes or tubercles surrounding the ovisperm duct. Bands of supporting muscles (SM) are attached to the preputium on both sides. narrow tube (about 12 mm. 578). 1) shows the vergic sac too short and the preputium too long as compared with the specimens personally examined. long.). fig. wide.5 mm. The sperm duct and vas deferens meet on the lower side of the prostate. fig. surface of the head in front of the foot' (Annandale and Prashad. The prostate is composed of many long. The verge (V) is a very long. ANATOMICAL CHARACTERISTICS (plate 21. In Rao's figure (1923. The tentacles are elongate and filiform. nar- The . the prostate diverticula radiating outward from this junction. its upper end narrowed to half the diameter of the lower is GENITALIA lobe. 288) shows the muscle attached to the vergic sac just above the preputium. The figure by Larambergue (fig. There is a single retractor muscle (R^I) attached to the vergic sac near the summit of the prejmtium. There is a small muscular ring or diaphragm between the vergic cavity and the preputial cavity. The ])rostate diA'erticula enter the sperm duct from which the vas deferens proceeds as a small tube less than half the diameter of the sperm duct. The seminal vesicle (SV) about 4 nun. Larambcrgue's figure 1939. The vas deferens is a very long. There may. The prostate (PRS) is 3. the vergic sac and the vas deferens are confused and the enlarged vergic sac is entirely too short. the retractor muscle is shown as attached to the vergic sac far above the preputium. A heavy nerve (N) innervates this muscle. causing the outer surface of the prostate to show twenty-four or more rounded projections. there are shown eight primary diverticula each of which is branched three or four times toward the outer end. The si)erm duct (SPD) is a small tube 7 mm.Family Bulinidae 197 The head is very broad and has its lower margin exi)anded and flattened. for a more extended account of external characteristics. long and nearly 2 mm. Each diverticulum is branched as shown in fig. In a cross section near the middle (fig. all of wdiich radiate fan-wise from the common meeting place of the sperm duct and vas deferens. branched diverticula. 215). p.

the differences may be attributable to methods of preservation. p. 5. 215) figures the verge (penis) as protruding from the vergic sac into the preputial cavity. N) It is clearly evident that this muscle pulls back the male intromittent organ after copulation. while in all specimens personally examined . The retractor muscle is attached to the vergic sac near its distal end and the large nerve lies beside it (RM. the sac lying on the body of the animal. A cross section of the preputium near its widest part is shown in fig. One of these specimens is shown in optical section in fig. In several specimens. The verge (V) and vergic sac (VS) extend through the preputium nearly to the opening of the preputium (PR). this bulbous portion of the verge was present in all specimens examined and a swelling of this region is shown in the figure of Larambergue where the long line for the symbol pe is directed. The writer figured and described a bulbous swelling within the vergic sac (plate 21. preputium and body during eversion. 5. 1933). A trifle more than half of the length of the vergic sac is everted. fig. 2 of plate 21. 6. Larambcrgue's figure 2 (reproduced as fig. the verge extended from the vergic sac and protruded into the perputium for some distance (plate 21. Rao (1923. plate 21). fig.198 The Molluscan Family Planorhidae freely movable within the vergic sac (VS). 1). the shorter portion remaining within the . The so-called supporting muscles of the preputium apparently become retractor muscles when the verge and the preputium are everted (fig. 293) describes and figures the penial complex in the everted position (see plate 74. p. In this specimen. He also figures the retractor muscle as attached to the vergic sac and not to the preputium. 7 of plate 22. greatly enlarged. 1. In other words. The branching of the retractor muscle and the connections of the nerve are shown in fig. It has been the writer's opinion that only the verge was protruded from the end of the preputium when the latter was everted from the male opening. The retractor muscle is shown by Larambergue as attached to the upper end of the preputium. B) which was thought to be the termination of the verge. This organ was enmisunderstood by Rao and was not fully comprehended by the writer row tube tirely (Baker. 2 on plate 73 of this work) shows that this is not the case and that the whole tube must be considered the verge. The bulbous portion of the verge is conspicuous and has moved downward. Larambergue (1939. the tube behind this swelling being considered as a part of the vas deferens. However. p. 288) correctly figures the relationship between the vergic sac (poche de penis) and the verge (tube penial). Two specimens examined by Baker had the preputium completely everted from the male opening. filling the greater part of the preputial cavity. Observations on the specimens dissected by the author do not entirely agree with those of Larambergue. These drawings of Larambergue were made from freshly anesthetized specimens taken in coitus. being almost twice the diameter of the proximal end near the preputium. as shown in fig. the latter being much longer than figured by Baker and extending the whole length of the vergic sac. The everted copulatory organ is enlarged at the distal end. 6). It seems probable that this bulbous portion (use unknown) is found in all examples of Indoplanorhis but in the specimens examined by Larambergue there was less contraction than in the specimens examined by Baker. Larambergue (1939. the preputium was greatly flattened so that the pilasters (AIP) became two flattened muscular pads. The vergic sac is shown extended from the everted end of the preputium. fig.

They radiate fan-wise from the ovisperm duct. The spermatheca is. as in all of the Planorbidae examined. narrow vagina (2 mm. The writer believes that the bulbous swelling (shown at B in fig. ) ( . 6. 4). fig. Rao 1923. distally branched. The ovotestis (OT) appears on the surface as a multilobed organ. 199 as shown in plate 22. 3). It might be that it marks the limit of the copulatory portion of the verge. long) by a very short duct. Female Organs (plate 21. but Larambergue figures it as having lateral swellings. 208) describes the lobes as occurring in groups of three or four. 9) suggest such a condition. evidently undeveloped ova. There are muscles on the preputiiun which probably aid in its retraction (plate 21. The uterus (U) is about 5 mm. as shown in fig. The oviduct (OD) is about as long as the uterus and vagina combined about 9 mm. attached to the long. although in a more lengthened condition. 4) groujied together in series of three to six folds. 5). fig. The albumen gland (AD. plaits. long. The pseudobranch (figs. p. The rectum (R) is on the side above the pseudobranch and the anal opening is between the pseudobranch and the pneumostome. these differences may be largely due to the preservation of the material examined. The schematic figures 8 and 9 in Larambergue's paper (cojiied on plate 74. long) It is 1 mm. 6). Several branched diverticula contained ripe ova at their distal ends. long joins the seminal vesicle to the ovotestis. in fact. 8. p. In the living animal. long and nearly four times as wide as the vagina. The ova were dark gray in color and with these there were several small. almost sessile. wide at the uterus end but decreases in diameter toward the albumen gland where it joins the sperm duct to form the ovisperm duct.Family Bulimdae this nmsclc. Respiratory and Renal Systems. smooth duct about 1 mm. as was observed in some species of Planorbidae. In the specimens examined it was entirely smooth. 7). but on the upper and lower surfaces of a leaf-like base are developed a number of projections with broad lobes. for it is also shown. long) pyriform. the folded portion of the pseudobranch is turned back against the mantle edge. Spermatheca (S) small (about 1 mm. figs. 7. As observed by Larambergue. was attached to the end of the vergic sac. The female organs are as figured in Larambergue's paper and also as described by Rao. 6. A small. a flattened. however. is convex above and concave below. in Larambergue's figure on page 288. plate 22) has some significance. more or less heart-shaped organ. fig. plate 21 is large and conspicuous and is not simply folded or leaf-like. each group of folds being separated from the other folds by a distinct depression or gap. 4. fig. The base of attachment of the folds is not modified by the folded series (see fig. or folds (fig. rounded bodies which were red in color. It has a narrow duct which enters the carrefour. but in the specimens examined there were two groups of six each on the under side of the base and two groups of three lobes and two groups of two lobes on the upper side. is about as wide as the uterus and is curved about the uterus and oviduct. long between the seminal vesicle and the oviduct. 7. The large nidamental gland (NG) is 6 mm. as suggested by the writer in a previous paj^er (1933. 1. plate 21. There is a small carrefour. The ovisperm duct (SO) is a narrow tube about 3 mm. and at B in fig. In cross section (near the anterior end) it is seen to be made up of seven or more main diverticula which branch twice at the outer end (plate 21. The pneumostome (PS) ( .

which resembles some of the large Hclisotna species (as Helisoma pilsbryi. The buccal sac is short. Intermediate teeth (6-11) narrower. highly vascular. The cross section somewhat resembles that of the kidney of Planorbarius corneus (plate 45. PS). The outer marginal teeth (27. The radula sac is represented by a wide. long and is sharply reflexed against the lower part of the kidney. thin lobe and fig. The kidney of Indoplanorbis is noteworthy for the size and width of the superposed ridge. Digestive System. fig. (fig.5 to 3 mm. wide ridge. fig. tricuspid. which is wider with larger glandular lobules. then makes another loop and passes forward to the enlarged rectum. The ureter is 1. The jaws (plate 50. p. and muscles. the posterior ]iortion of tiie salivary glands (somewhat less than half the length) is floubled backward beneath the anterior portion. The stomach resembles that organ in Australorbis glabratiis (i)late 48. 10) consist of a wide and low superior jaw. blind sac near the pyloric portion of the stomach. differing from the typical laterals in having a second small cus]) above the ectocone. The two salivary glands are rather long. The entocone is split into three to five small. fig. 10). the kidney resembles that of Helisoma trivolvis figured on plate 44 (fig. about small cusps above the ectocone. The reflexed ureter in the s})ecimens examined is not so long as figured by Rao 1923. In the natural position. Marginal teeth three times as long as broad. wide. squarish. 2). figs. The kidney is long and narrow (12 mm. pointing backward and upward into the mantle cavity. Lateral teeth (1-5). which has its exit in the anus above the pseudobranch. A large vein lies on each side of the lumen. The salivary glands are joined behind as in Helisoma and other planorbids. slightly bent downward at the ends. striated on its outer face. wide). subequal cusps. 200 is The Molluscan Family Plauorbidae is a large. 2. long. and high. The ducts are short and narrow. 1. In form. 18). 32) are still narrower and the lower edge of the reflection is minutely denticulated. flatly rounded bulge at the lower posterior extremity of the buccal sac. The lateral jaws are in the form of a question mark. fig. and the ectocone is smaller with two to three small cusps above it on the outer margin of the tooth. it is roughly heart-shaped.. The jaws of Indoplanorbis do not differ notably from the jaws of the subfamily Helisomatinae. The extreme outer marginal teeth are very small (33) ( . 12). Viewed from above. The center tooth of the radula (plate 66. There is a large caecum or ( . well-supplied with blood vessels. thick. all cusps wide and spade-shaped. 1) is higher than wide. the ectocone the shortest and i)laced high up on the reflection. The lumen is rounded and shows internal folds or septa. There is a narrow. 19) shows a high. rather narrow. plate 44. it is capable of considerable extension sjiread open. The tenth intermediate tooth has two 12-26) narrow. the reflection broadly bicuspid and reaching nearly to the lower margin of the base of attachment. passes backward to the posterior part of the liver. are very narrow and about as long as the width of the superior jaw. 206) A cross section near the middle (plate 45. It more nearly resembles the molluscan branchiuni than the pseudobranch of any of the other fresh-water pulmonates. In shown turned back and is The pseudobranch veins. 4. the mesocone the longest. about one and a half times as long as the buccal sac when fully extended. fig. The intestine first makes a loop around the stomach. the mesocone is larger and distinct.5 mm. 9. low ridge on the rectum which ends near the pseudobranch.

Rao. The genus as represented by the type PUniorbis cxustus Desh. It is the largest planorbid in India and the shell closely resembles certain species of the American genus Helisoma. French Indo-China. Remarks. is in doubt. S. 514). N. particularly in the form of the penial complex. There is some variation among the different rows of teeth. The most convincing characteristic is the pseudobranch. p. which is lobed in Bulinidae (Germain. The systematic position of Deshayes' Planorbis exustus has been a matter of interest for a number of years. and-Sumatra (vide Annandale). collected by Dr. p. The number Germain (1921.' Pilsbry (1934. 54) suggests its resemblance to Tropicorbis. its male complex exhil^iting characters not shared by any other group at present known. 27) places almost everything under exustus as syno- nyms of that species. 291. B. particularly the penial complex. 28. In both of these species. The specimens were received from Dr. Choi)ra and Dr. 5. The shell of Indoplanorbis exustus is distinctly sinistral. H. Bulinus contortus of Europe by Larambergue 1939) and Isidora globosa from Portuguese East Africa by Connolly. Baker (1933) stated that it was 'one of the most distinct groups of the Planorbidae. A careful study of all names in connection with a large series of specimens from various localities might establish several recognizable forms. Helisoma duryi seminolina. the deciding features being the lobulated pseudobranch and characteristics of the male genitalia. The radula figured on plate 66 agrees with that described by Rao (1923. .Fcunilij Bulinidae 201 The nulula formula of Indoplanorbis is 33-1-33 with 140 to 150 rows of teetli. 294) argues that the genus Indoplanorbis is a group of the family Bulinidae. Geographical Distribution. occurs throughout the plains of the Indian Empire east of the Indus. principally in the nmiiber of accessory cusps in the marginal teeth. 29). Burma. Little is known concerning the anatomy of the majority of the species of this family. of species or races referable to Indoplanorbis p.. Planorbis coromandelicus Sowerby and Planorbis indicus Clessin have been thought to be distinct by some conchologists. p. ]\Iore recently Larambergue (1939. An examination of its anatomy led Annandale and Prashad (1920) to consider it a new genus and Rao's investigations (1923) strengthened the view of its distinction as a generic group. especially in the form of the penial complex and the prostate (see plate 71.) The anomaly of a distinctly i^lanorboid shell in a almost exclusively of physoid shells is no greater than group made up is shown in the American subgenus Seminolina where one species. 3 and plate 75 of this work). fig. North Shan State. the Malay Peninsula. B. in Siam. The radulae examined were remarkably uniform. In Isidora globosa the lobed condition is well shown in Con( nolly's figure (plate 71. pp. of this work. The writer agrees with Larambergue in the statement that the group Indoplanorbis is a distinct genus of the family Bulinidae. contains all shapes of shell from distinctly physoid to normally planorboid. Prashad of the Zoological Survey of India. the genitalia are similar to those of Indoplanorbis. species. fig. The material examined consisted of six specimens from Hsipaw. 204). par- ticularly so in the young and immature stages of growth. Two species have been rather carefully studied. 1931. For a more detailed account of the distribution of this group see Germain (1921. pp.

although considered by Thiele. Nautilinus 1921. fiir XXV. Only an examination of the anatomy of the animal. in diameter. 28-30. p. 385. The figures given by Mousson (his plate 1. The single cusp of the center tooth removes clymene from the Planorbidae in which the center tooth always has two cusps. Thiele extracted a dried radula from a shell of this species and briefly described the teeth. Conch. p. possibly Polychaetes. Belgique. The center tooth is small and has one indistinct cusp. 324) says 'I am extremely doubtful whether this curious little Planorbis-likc shell should be associated with Hyalina. Islands. ser. I. Tryon (in Man.' The name does not occur in any other edition of Zittel's work. 481. Ancylus. The form of the shell is distinctly unlike any form of the family Lymnaeidae. Schweiz. The side teeth (laterals and marginals) have four sharp ctisps. XXII. etc' Pilsbry (Man. IX.). The formula is 20-1-20. p. GENERA WRONGLY REFERRED TO PLANORBIDAE Genus Type by 1872. die zierliche Palaeorbis is The only in Zittel's . As subgenus of Anisus Type A. It does not appear to be referable to any mollusk. is scarcely a member of the family Planorbidae. ii. will definitely settle the question of the taxonomic position of Nautilinus. Radula and generic position Nautilinus Thiele. Beneden and Coemans. p. 1867 Bull. As subgenus of Hyalina Thiele. Genus 1867.. Wollaston (1878. 424. pp. original designation. Grundziige der Palaontologie (Palaozoologie) Abth. so far as known to the writer. 111. et Coemans em. The radula resembles some groups of Ancylidae and Nautilinus might prove to be related to this family. Ill). 1886) places it in Zonitidae and says 'Its habitat is different from the Hyalininae generally. living in wet moss. LIII. 1931. Type Mollusk. Neue Denksch. 19. Hyalina clymene Shuttl. This genus. Archiv.. only 2 nun.) clymene Canary so figs. Teil 2. p.. (N. in which the center tooth of the radula is imicuspid. Nautilinus Shuttl. 1872 Gesell. p. in which the shell is particularly Planorbislike. 24) lists it among the land shells. Allg. Type Hyalina clymene (Shuttl. 31-33 enlarged) do resemble some forms of Gyraidus. Invertebrata. 172. perhaps in a similar manner to the American genus Neoplanorbis Pilsbry. Conch. natural size.X. II. based on a supposed land snail from Garachico. NAUTILINUS Mousson. centainly not to the Pla- 202 . Handbuch. figs. Palacorbis PALAEORBIS Beneden and Coemans. 390 recent reference to this group. w^iere the following comment is made: 'Hierhcr diirfte wohl auch die Planorhis sehr ahnliche Gattung.^ In 1921 (p. The shell is very small. The group is believed to include fossil Vermes. Bened. Acad. Tencrife. p. especially the genitalia. and Hydrocena. Reis aus oberkarbonischen und permischen Ablagerungen von Europa und Nordamerika gehoren. Hyalina clymene Shuttleworth Mousson. associated with Physa.

.Genera Wrongly Referred norbidae. The generic name does not appear. It is j^robable that some Planorbidae will eventually be found in Palaeozoic strata. which are to Planorbidae 203 unknown in and Pennsylvanian) and Permian periods both the Carboniferous (Mississippian of America. as far as known to the writer. since the family is so well developed in Mesozoic time. in any American work on fossil puhnonates.

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in text.

Magyarorszagi Planorbis-Feiek warkesziilekenek Alak-es Szovet-



Kozlemem-ek, 32 Kotet, 1-2 Fuzet, pp. 21-42, 4 figs, in text. Morphologic und Histologic des Geschlechtsapparates der ungariAUat. Kozlem..

schen Planorbis-Arten.


pp. 21-46,' 2 Tafeln.

Soos, Lajos (1917). Zur systematischen Anatomic der Ungarischen Pulmonaten. Musei Xat. Hungarici, XV, pp. 165, 100 text figs.



J. P. E. F. (1850). Die lebenden Schnecken Berlins, etc. Berlin. Pp. viii-|-120, 3 plates.

und Muscheln der L'mgegend

Studer, S. (1820). Kurzes Verzeichniss der bis jetz in unserm Vaterlande entdeckten Conchvlien. Xaturw. Anzeiger Allg. Schweiz. Gesell. Gesammt. Naturwiss., Ill,

No. 12, pp. 83-90. SwAiNSON, W. (1840).



on Malacolory. Pp. viii+419, many text



(1894-1900). Monograph of the land and fresh-water Mollusca of the British Isles. Structural and general volume. Leeds, pp. 454, plate vi, 743 text figs.
(1928). Mutations of land and fresh-water Molluska. Jour. Conch., pp. 97-116.

John W.


Thiele, Johannes (1921). Ueber die Radula und svstematische Stellung von, LIII, pp. 111-112. s.ystematischen Weichtierkunde. Jena. Erster Band, Erster und Zweiter Teil, pp. vi+778, 782 figs, in text. Tryon, George W. (1870). A monograph of the fresh-water univalve Mollusca of the United States (in continuation of Prof. S. S. Haldeman's work published under the above title). Pp. 238, plates irregularly numbered.

clymene Shuttl. Archiv.

Handbuch der

(1871). Review of Meek's paper in Proc. Acad. Xat. Sci. Phil., 1870, p. 59. American Jour. Conch., VI, p. 285. (1884). Structural and systematic Conchology. An introduction to the study
of the Mollusca, III, 453 pp., plates 92-140.


and fresh-water shells of the British Islands. Pp. viii+150, 16. 10 colored plates. London. (1857). A manual of the land and fresh-water shells of the British Islands. New edition by J. E. Gray. Pp. xvi+335, 12 colored plates, many text figs. London. VanCleave, H. J. (1932). Statistical analysis of quantitative collections as a means of interpreting life histories. Trans. Illinois St. Acad. Sci., Vol. 24, No. 2, pp. 228-234. (1934). The seasonal life history of an amphibious snail, Fossaria modicella, living on sandstone cliffs. Ecology, Vol. 16, No. 1. pp. 101-108. VanCle.u'E, H. J., and Altringer, D. A. (1937). Studies on the life cycle of Campeloma rufum. a fresh-water snail. American Nat.. LXXI, pp. 167-184. VanCleave, H. J., and Lederer, L. G. (1932). Studies on the life cycle of the snail, Viviparus contectoides. Jour. Morph., Vol. 53, No. 3, pp. 499-522. W.-iLKER, Bryant (1918). A synopsis of the classification of the fresh-water Mollusca of North America, north of Mexico, and a catalogue of the more recently described species, with notes. Miscel. Pub., No. 6, Mus. Zool., L'niv. Michigan, pp.
of the land

TuRTON, W. (1831).

A manual

plate, 233 text figs.

Ward, Henry

The Molluscan Family Planorbidae

B., and Whipple, Geo. C. (1918). Fresh-water Biology. New York. (Chapter XIII, Parasitic flatworms, pp. 365-424, by Henry B. Ward.) Wenz, W. (1923). Gastropoda extramarina tetiaria. In Fossilium Catalogus I: Animalio, Pars 22, pp. 1421-1734. Edited by C. Diener. Berlin. White, C. A. (1883). The Molluscan fauna of the Truckee group, including a new form. Proc. United States Nat. Mus., V, pp. 99-102, plate 5. lebenden Westerlund, C. a. (1885). Fauna der in der Palaarctischen Region Binnenconchylien. Lund, Fascicle V, pp. 135-|-14. (1899). Planorbis libanicus n. sp. Nachrbl. Deutsch. Mai. Ges., pp. 170-171. (1902). Methodus dispositionis Conchyliorum extramarinum in regione Rad. palaearctica viventium, familias, genera, subgenera, et stirpes sistens. Jugoslav. Akad., CLI, pp. 82-139. (1-58). WoLLASTON, T. V. (1878). Testacea Atlantica, etc. Pp. xi+588. London. Woodward, S. P. (1851-1856). A manual of the Mollusca. Published as follows: Pt. 1,
. . .

pp. 1-158, 1851;

quent editions.

Pt. 2. pp. 159-330, 1854; Pt. 3, pp. 331-486, 1856. Several subse(Raljih Tate has an appendix in 1869 in which Pompholyx

Zittel, Ivarl a. von (1910). Grundziige der Paliiontologie (Palaozoologie). I. Abteilung Invertebrata. Neubearbeitet von Dr. Ferdinand Broili. Munchen und Berlin. x4-607 pp., 1414 text figs.



*It was Mr. Baker's plan that Part II should contain full morphological and taxonomic descriptions together with comprehensive illustrations of the shells of all species of the Planorbidae known for North and South America and the West Indies. His voluminous working notes were not near enough objective organization at the time of his death to permit their being edited for publication. Any attempt at preparation of these notes by another hand could not ensure the reflection of Mr. Baker's matured judgment and ultimate intent in the casting of specific descriptions, in the portra.yal of indi\idual variability, and in the discussion of geographical distribution and ecological relationships. However, at the time of his death, he had completed fifty-four of the plates intended for this section and had prepared typewritten copy for them. In his desk were found plainly-marked photographic prints of other species sufficient to prepare six additional plates and pencil manuscript for descriptions of the figures had been written out. The entire group of sixty plates (82-141) is included as a supplement to Part I, even though the six plates at the end of this series represent working copy in which he might have desired to make some final alterations. For convenience of arrangement and reference, the plates of Parts I and II are presented in a single undivided series at the end of this monograph. In addition to the plates for Part II, descriptions ha\'e been prepared for nine new species and seventeen new varieties. The foreword was written by Professor Harley Jones Van Cleave, who also assembled and edited the descriptions of the new forms and edited the explanations of the plates.— H.E.C.


preparation of detailed descriptions of the species and critical evaluation of the synonymy of the Planorbidae of the Western Hemisphere. This study was the natural outgrowth of the long years of detailed morphological studies which led him to the evaluation of anatomical detail as expressed in the broader phases of taxonomy in the main body of the present volume. The second volume in this series was to have treated the species of the Planorbidae in a manner similar to his treatise on another large and important family of fresh-water snails, the Lymnaeidae. Though published in 1911, his volume on the Lymnaeidae of North and Middle America after more than thirty years still stands as the leading taxonomic and distributional study of the lymnaeid snails of the
in the

FOR ALMOST A QUARTER OF A CENTURY, Frank had been engaged



Western Hemisphere. Though such a program of study involved intimate familiarity with the voluminous, scattered literature and wise interpretations of matters of disputed priority and synonymy, these labors paled into insignificance when compared with the self-imposed task of dissecting and microscopically
studying representatives of all the species available through the wellrecognized repositories of collections. To these customary sources were added large quantities of material from individuals throughout the world with whom kindly cooperation had established intimate personal and professional contacts. Large numbers of dissections and long series of shells gave opportunity for bridging the gap between the earlier field of Conchology and the newer science of Malacology to both of which Mr. Baker had made notable contributions. As he reached final decisions on the validity and synonymy of the species in the various genera, he prepared photographs of the shells or of original drawings of the type specimens wdien shells were not available. These photographs he arranged as plates and for each he prepared a detailed descriptive legend. He had completed plates illustrating the shells of the genera HeUsoma, Carinijex, Parapholyx, Planorhula, Menetus, Drepanotrema, Tropicorbis, and Taphius. In his office at the time of his death were hundreds of photographic prints of other genera and species intended for use in this monograph. Some of these he had so marked and keyed with numbers that it has been possible to assemble six additional plates which are included at the end of the series. With this inclusion, all of the genera which he apparently intended to figure are included though many of the species are not portrayed in the detail which he established in those earlier parts which he had completed. The explanations of plates 136 to 141 are compiled from rough pencil notes and must be accepted as the author's tentative arrangement of work in progress which might have been changed somewhat in

the final editing. Some of the species not included in the plates of the api:)endix have representative shells figured on plates 72 to 81 of the illustrations for the body of the monograph. However, these figures are usually of only the type species in each genus and do not carry out the plan of comprehensive comparisons set for the second section of the monograjih as originally projected by the author.



The Molluscan Family Plauorbidae

In the sixty plates included in this appendix (plates 82 to 141), many type specimens and other critically determined materials are figured, and information as to the present whereabouts of these critical materials is made available. In the members of the family Planorbidae, where synonymy has been so involved and w^iere intergradations in shell characters have been so confusing, the interpretation of range of individual variability by one who has devoted years to the study and had the singular advantage of access to all the important collections, will be appreciated by all naturalists who have the need for studying snails. For convenience of reference, the plates in the appendix are numbered in direct continuation with the series of plates in the body of the monograph. In the explanation of the plates, the repository of the figured specimens is usually indicated by the following abbreviations:
A.N.S. Academy of Natural Sciences of Philadelphia B. Frank Colhns Baker Collection C.A.S. California Academy of Sciences M.C.Z. Museum of Comparative Zoology, Harvard University U.C. University of Colorado Museum U.I. University of Illinois, Museum of Natural History U.M. University of Michigan Museum U.S.N.M. United States National Museum

— — — — — —

In connection with the foregoing list of repositories, it should be here recorded that the Frank Collins Baker collection of fresh-water snails and the specialized library accompanying it, after his death, was deposited in the United States National Museum. This was in keeping with his expressed desire. It should be further explained that the collections attributed to the University of Illinois are designated by two series of letters. Those 'U.I.' entries having the letter 'Z' preceding the accession numeral are in the zoological research collections of the Museum of Natural History in the University of Illinois. Paleontological specimens, indicated by insertion of the letter 'P' immediately before the accession number, are now deposited in the Paleontological Collections of the Illinois State Geological Survey, in Urbana, Illinois. Citations of magnification in the explanation of the plates are usually only approximate. In final preparation of the cuts, it was in many instances unavoidably necessary to make slight modification of the author's indicated amount of reduction from the original plates so as to conform to the available page size. In listing the names of the species in the explanation of plates in the appendix, Mr. Baker placed the name of the author of the species immediately after the specific name and frequently gave no indication by the use of parentheses in instances where the si~)ecific name was recombincd with a generic name other than that to which the species was originally ascribed. It should be recalled that the preliminary draft which he prepared was never finally edited by him. Through the very generous cooperation of a number of the most widely recognized American authorities in the study of the Mollusca, the use of parentheses for author names has been carefully checked. Some errors or inconsistencies may yet be found because the literature is extremely scattered and there are no comprehensive check lists of the species of Planorbidae. As previously explained, the manuscript for the section in which Mr. Baker had expected to provide full descriptions of all species and varieties

of the Plan()rl)i(l;R' of the


c()mi)leted. In the unfinished fonn, there were included the preliminary drafts of descriptions of many new species and new varieties intended for incorporation in Part II. After it had been decided that the plates for Part II should be included as an appendix to the morphological studies, it was discovered that the legends for these plates made frequent reference to previously unpublished specific and varietal names. His manuscript descriptions for many of the new species and new varieties were so well organized that there could be no doubt regarding his intention. Consequently, liis jireliminary descriptions have been edited and are included as a separate section of Part II. A few of the varieties and species which he tentatively considered as new could not be found in his files. For these it is assumed that he had never prepared definite descriptions. In order to avoid nomenclatorial difficulties, these names have been deleted to prevent their becoming nomina nuda. In each such instance the undescribed and unnamed form has been referred to under its proper genus as a questioned species or questioned variety.

New World

had never been

vohnninous long-hand notes which he

left in



and new varieties were edited from longhand notes -**- prepared by Mr. Baker. During the past quarter of a century he examined every important collection of Planorbidae housed in the museums of America and received extensive field collections from many private collectors as well. In the handling of this huge quantity of material he came to recognize fundamental likenesses and differences at vaiying levels.



in the foreword, the following technical descriptions

In addition to his study of the shells, his intensive work on the morj^hology of the snails gave him a broad background for the recognition of natural
the one side, he expressed his conclusions in the broad new Planorbidae as set forth in Part I of this volume. There he evaluated the characteristics on which families, subfamilies, genera, and subgenera may be recognized with safety. On the other side, his attention to the limits of individual variation in characteristics of the shell furnished the basis for the detailed recognition of specific and varietal boundaries. The descriptions of new forms and the redescription of all known species were to have been the final goal of Part II of this study. From his incomplete notes the following descriptions include only those for which he had prepared tentative definitions. On other species and groups.
classification of the


which he obviously thought of as new, he had not committed himself through the preparation of manuscript notes and in the assembly

of illustrations.

The descriptions of nine new species and seventeen new varieties are arranged in the same sequence of genera and subgenera as outlined in Part I (see pages xiv and xv of the Contents) All tabular arrangements of shell measurements are expressed in

Plate 134,


Tropicorhis shivtieki,



12-14, 28

Shell solid, small, of three and one-half whorls. Upper surface showing three and one-half whorls which are subangulate above, sutures very deep. Lower surface showing three and one-half whorls which are subangulate in the middle, sutures very deep. Whorls rounded on the i)eriphery. Apical whorls sunken on both surfaces. Whorl slightly deflected at aperture. Aperture Innately rounded, outer lip thick with callus. A callus on parietal wall connecting the extremities of the outer lip. Sculpture of coarse growth lines with fine spiral lines. Apertural lamellae six, those of sigmoid form extremely short and thick, only slightly more than half the length of those found in obstructus.


Descriptions of


Species and




Material. Collected by B. Shimek in 1893. Four specimens deposited in United States National ]\luseum, Accession no. 534290. The holotype is shown on plate 134 as fig. 12; figs. 13, 14 are paratypes. Tropicorbis shimeki resembles albicans but is smaller, has a greater axial height, and has subcarinate whorls. The parietal lamellae are distinctly shorter and thicker than in other forms of the genus. T. shimeki is smaller than declivis and its axial height is greater; it also has deeper
sutures than are found in declivis. Additional representatives of T. shimeki were collected by Orcutt in Coatzocoales, Mexico (United States National Museum, Accession no.

Type Type

Locality. Ometopc, Nicaragua.

219696). This species




honor of Professor B. Shimek.

Plate 96,


Helisoma anceps anticostianum,




Shell differs from typical anceps in the low axial height, the wide and shallow spire depression with bluntly angular carina, and the shallow and wide umbilical region of the new variety. The aperture is but slightly expanded. "Whorls about four, evenly and regularly coiled, with faint
spiral lines.


The Molluscan Family Planorbidae
thickened and expanded, white, bordered by reddish band inside.

Parietal wall with white callus.



Aperture Height

Aperture Diameter

6.5 6.1 6.6

11.7 12.0 11.6

5.5 5.1 5.5

3.3 3.0 3.3

Holotvpe Paratvpe Paratype

Material. Holotvpe and seven paratypes, deposited in the United States National iMuseum, Accession no. 227858. Holotvpe figured on plate 83, fig. 19; paratypes figs. 20-22. This variety may be distinguished by its small axial height, depressed shape, wide and shallow spire depression, flat basal whorls with carina near periphery. It resembles the variety poUtum from Canada, but that variety has rounded basal whorls without marked carina, its spire depression is wide with more flat-sided whorls, and dorsal carina is more acute. The basal part of the aperture is rounded or almost flat in politum, but has sharp, wide angle in bartschi. Bartschi differs from sayi in lower axial height, less sharply angled shallow and wide basal whorls (with carina not near edge as in sayi) spire depression. Bartschi differs from anceps in lower relative axial height (compared with diameter) wider spire depression; basal whorls flatter; carina nearer periphery (near center in anceps). Helisoma anceps bartschi is apparently a common form in the Potomac
; ;

Type Type

Locality. Brook at Great Falls, Virginia. Collected by Soelner.


Helisoma anceps idahoense.
Plate 84, Plate 101,




figs. 11,

Spire with carina in center. Base concave, basal whorls flat to sutures which are well marked. Whorls four. Spire depression deep and funnelshaped, the whorls flat-sided. Carina well marked. Whole shell is coneshaped, the whorls sloping upward to the spire carina. Color yellowish horn. Sculpture of regular growth ribs crossed by distinct spiral lines. Aperture auriform (reversed), sharply angled above, flat or flatly-rounded below. Lip sharp. Distinct columellar callus. Color of aperture whitish or slightly


callus often within the lip edge.
Shell Height

1863. In logs. the front half of the body whorl with a faint ridge about in the center of the whorl. Holotype (plate Baker Collection. fig. often raised into slight ridges. rounded carina. Idaho. Shell Height . ^vider than high. spire deeply sunken. Pend Oreille River. Sculpture of fine. Idaho and Alontana. straight or but slightly curved below. thread-like riblets. New Variety 16-18 showing three and one-half to four whorls. now and paratypes in 84. higher than wide. Material. Aperture ovate. Ridge disappears on the last third of the body whorl which is rounded. Color of shell pale yellowish to dark brown. showing three and one-half to four whorls. The lip edge is white on the outside. obtusely angled above. the depression widely cone-shaped rising to a vaguely defined. Locality. Plate 83. Umbilical region flattenecl. no. anceps idahoense is apparently common some places it occurs on muddy sand or on old HeXisoTTia anctps ^oliiutn. Outer lip thickened. Surface polished in living specimens. Sand Point. H. especially toward the aperture. Fine impressed spiral lines present in all specimens examined. C. bordered within by a reddish-brown band. Shell flattened.Descriptions of New Species and New Varieties 221 Type Type r. 26) deposited in United States National in Museum. figs.

figs. Shimek Collection. New 16-21 Species Plate 119. the holotype is figured on plate 119 as fig. The sides of the depression slope sharply downward. Sutures well marked. Bennett. especially the body whorl. Shell Greater Lesser Height Diameter 19. fig. 404719.0 6. Some living Cuban specimens have a somewhat narrower spire than the typical form here figured but these lack the greatly increased diameter of the body whorl characteristic of cubense and the aperture is not auriculate or thickened by a callus as in mature and nearly mature specimens of cubense. 1927.1 Diameter 16. Helisoma carihaeum cuhense.2 4. are deposited in the United States National Museum. Type Type Locality.3 Aperture Diameter 4.1 16. and are not like those of tenue which are flat and carinated at the edges. Seattle. Holotype figured on plate 94. 5. not rounded.0 3.n&e.222 The Molluscan Family Plauorbidae Locality. Material. B. 1. the lip sometimes (as in the holotype) thickened by a heavy callus. Sutures are not as deep as in caribaeum. 23-25.1 6. Accession no. Accession no. umbilical opening distinct. the spire whorls flattened. and three of the paratypes are shown as figs. immature.5 12..0 14. Sculpture as in caribaeum. Basal whorls rounded. Aperture more nearly auriculate than in caribaeum. the last whorl with a more or less sharp carina or ridge. especially like that of tenue sinuosum. widely spaced. in marl between layers of peat. increasing more rapidly in diameter.0 13. figs. collected April 19.0 14.0 3.1 Aperture Height 8. The basal whorls are more concave than in caribaeum. rapidlv enlarging. Sutures not impressed as in caribaeum.0 6. paratypes. broadly funnelshaped. Collected by P. HeXisoma colum\)ie. fig. deposited in United States National Museum. Base of shell concave.5 4. Cuba. the whorls. H. Axial height somewhat greater than in typical caribaeum.0* 4. Zapata Swamp. Of these. Type Type Lake Wasliington. Nuclear and first whorl rounded as in caribaeum and of the same size. by H. not evenly rounded as in typical caribaeum.0 11.A. Spire depression flattened. 2-4. the umbilical region concave. the inner whorls bordered by a sharp carina which becomes distinctly central in position on the last whorl. Material. Sculpture of coarse riblets.perture contracted in holotype liy thickened lip. Plate 119. figs. New Variety 22-25 Shell differing from that of typical caribaeum in having the spire depression much narrower.7 7. of the whorls increases like that of tenue. the body whorl flattened near the spire depression. Randolph. as in typical sub- . Evidently a Pleistocene fossil. 22. Shell with four to four and one-half whorls.0 6. Five full whorls. 504360. Eight specimens.1 Holotype Paratvpe Paratvpe Paratvpe *. and there is often a carina at the edge of the whorls. but inner whorls are like those of caribaeum in that The form they are rounded. Washington.

A callus on the parietal wall renders the aperture continuous. Aperture ovately rounded. The basal part of the aperture is flatly rounded. The aperture may be rounded above or there may be a sharp V-shaped angle. the li)) riiu thickened and broadly reflected. There is usually. There is normally an indication of angulation at this point. Several old lip expansions may show on the body whorl back of the aperture. Color whitish. a border of dark chestnut or reddish brown within the aperture. . tinged with chestnut in some specimens. depenfling upon whether the dorsal carina is sharp or rounded.Descriptions of Xew Species and New Varieties 223 crenatum.

It differs from the latter in its expanded lip and more tightly coiled whorls of the umbilical region. Figured on plate 91. Knee Lake. rounded above and below. Helisoma pilshryi prehlei. Sculpture as in pilsbryi. paratypes as figs. H.0 26. 17. New Variety 13-17 Shell small. horn colored. Expressed in percentages of height to diameter. Deposited in United States National Museum. bearing the label of the old Binney collection no. The aperture is widely expanded to form a reflected lip. 8. the one-half whorl broken away. 180279. causing the spire depression and umbilical region to appear smaller than in pilsbryi in which the whorls are rounded more regularly. Accession no. 13. . Canada. Height at Diameter at Shell Expanded Expanded Lip Lip 17. kennicotti is pression sixty-five per cent and infracarinatum. as compared with diameter. only one and one-half turns before entering the umbilicus. Differs from infracarinatum in the absence of a basal carina and in having a narrower and more sunken spire depression. Helisoma subcrenatum perdisjunctum^. Plato 91. 15. In contrast. It differs from injracarinatum in its smaller size. Whorls slowly increasing in diameter. Preble in the 'North West Territory. figs. Accession no. 16. holotype as fig. the body whorl increasing more rapidly in diameter. New Variety 6 to 8 Helisoma pilsbryi prehlei differs from typical pilsbryi in its greater relative height in comparison with diameter of the shell. 14 immature shells. fifty-six per cent. 29231. Plate 96. Figures on jilate 89. The spire deis much smaller. Three specimens. The whorls in prehlei are more tightly coiled. The whorls of the spire depression are subangulated. narrower. 3 whorls below. greater relative axial height. Type Material. Knee Lake. collected by E. preblei differs from subcrenatum in absence of all indication of a carina on the umbilical side. The base of the new species is likewise narrower. Type Locality. holotype as fig. figs. Keewatin. not carinated. Lake Isle la Crosse.224 The Molluscan Fainily Planorhidae Helisoma kennicotti is a very distinct species.0 27. Canada. whorls three and one-half to four.' deposited in United States National Museum. The type material was i:)reviously called infracarinatum but it is nearer to pilsbryi. the shell forming an inverted cone. in infracarinatum tliere are two full turns visible.0 last *Bioken specimen with four and one-fourth whorls. 6. Whorls. Manitoba. evenly rounded. 9272 and there recorded as Planorbis trivolvis. English River. p. Sculpture of distinct growth lines. Material. This is not true of pilsbryi. and deeper in kennicotti. discoidal. four and one-half. Type Type Locality. A. paratypes as figs.0 16. not sharply angled as in infracarinatum. 7. and the whorls dip into the umbilicus at a sharper angle.

Aperture rounded above and below. A distinct callus on the parietal wall. slightly expanded where it joins the body whorl. Shell . A slight thickening inside outer lip. bordered by brown band. often making the aperture and lip continuous.Descriptions of Xeiv Species and Xeiv Varieties 225 finer than in disjunctum.

figs. Accession no. New Jersey. Shell resembling michiganense in the low axial height.226 The Molluscan Family Planorbidae Locality. New 16 Variety 12. upper whorls irregular. Washington. Material. 13. 13. dalli.2 . base 'reamed out' similar to that of rudentis. D. t. Umbilicus much smaller than in rudentis and also smaller than in michiganense. but the body whorl is irregularly coiled and overlaps the penultimate whorl. 15. In addition to the type locality. C. Lower whorls rounded. deposited in United States National Museum. paratypes as figs. Shell Maximum Diameter Lesser Height Diameter Aperture Height Aperture Diameter 6. HelisoTTia campanulatuTn Plate 111. Collected by Foreman. 124989. 11 and 12. and whorls irregular. Four whorls showing on base. Type Type and Virginia. marshalli has been identified in collections from various localities in Maryland. 14 and 15 of immature shells. Holotype on plate 87 as fig. Five whorls. fifis. H. blunt carina on upper whorls. Sculpture and aperture as in michiganense. New York.

Sculpture of heavy. about as in some individual In the following table the measurements were taken just behind the aperture. Umbilicus small. which is constricted and then dilated.Descriptions of Neiv Species and N'eiv Varieties 227 Helisoma campanulatum rideauense. evenly spaced but not as heavy as in wisconsinense . the inner whorls raised slightly above the body whorl. shells of ca»ipanulatiuu. Plat (> New Variety- 109. sutures well marked and whorls rounded. Base with three to four whorls visible. in same plane.^. deep. Aperture as in campanulatum. riblets. not reamed out as in dalli and rudentis. of five whorls. 21-24 Shell very large. fig. Shell Height . rounded. sutures deeply marked. spire flattened.

Some individuals of costata may have slightly these are never like those of dalli. more tightly appressed to the columella.^. Klamath Falls. but not markedly angular in some specimens. sinitsini. Shell . Seventeen specimens. Whorls three. Parapholyx klamathensis Plate 123. Shell similar to that of klamathensis but smaller. not as effuse. Columellar callus white. Aperture angulate below. lig. New Variety 2-10 more globose. Sculpture heavier than in klamathensis. 27. Spire more elevated than in typical klamathensis. paratypes as figs. Lip bordered internally by a white margin which in old specimens forms a thickened callus. in a few specimens rib-like. klamathensis of typical variety has a thin lip without a callus. Oregon. growth lines more distinct. never greenish. The Molluscan Family Planorbidae membranous ribs. 219749. columella callus heavier. Accession no. Some specimens with diagonal markings like pores. Color brownish horn. The umbilicus usually closed. rarely having a small chink. 28-30. Type Type Locality. Interior of aperture reddish-brown.228 ture. The holotype is figured on plate 115 as fig. but Material. as in klamathensis. the shell thicker and more solid. collected by Henshaw. In contrast. are deposited in the United States National Museum. aperture rounder.

jiaratyjies on the same plate as figs. sutures distinct. 11 and 12. H. the spire whorls in the same plane and not so sunken toward the nucleus as in wheatleyi. Smith.0 2. turban-shaped.3 4.3 1. GENUS Plate 118. as fig. Fifteen specimens. umbilical opening wdth the whorls rounded. Parapholyx pusilla differs from Parapholyx rievadensis in that the new species has a more rounded shell. Aperture continuous by the presence of a parietal callus. The new species differs from P. Apertural lamellae as in armigcra not as in wheatleyi. One hundred twenty-five specimens taken from a Pleistocene deposit by Elbert Howard^ are deposited in the United States National Museum. 308925.7 1. This is apparently an extinct sj^ecies.0 1. Columella callus narrow. The individual from this lot chosen as holotype is shown on plate 117 as fig.9 Diameter Aperture Height 3. Type Type Locality.4 2.1 7. Sculpture coarse. Six miles west of Pyramid Lake. figs. lower lip rounded at base. lips simple. fijis.5 4.8 2. of five whorls.4 Holotvpe Paratvpe Paratvpe Type Nevada. 11-12. not expanded. 10.3 2. Washoe County. Externally Planorhula smithii resembles wheatleyi but the newly recognized species is readily distinguishable by the absence of the L-shaped . often costate.5 2. with three whorls. not flat sided and without the sharp carina distinctive of wheatleyi. 1. with elevated and turban-shaped second whorl and the lower portion of the aperture lacks the sharp angle caused by the columcllar callus in Jievadensis. Alabama. 10-12 Shell resembling that of wheatleyi but larger. Accession no.0 4. Type Material. New 1-6 Species Plate 117. Jackson County. the second whorl standing on the body. scalariform. separated from the columella region where it often projects leaving a distinct umbilical chink. The holotype is figured on plate 119. no dorsal ridge as is developed in wheatleyi. Shell Maximum Diameter Height Aperture Height Aperture Diameter 4.8 2. are deUnited States National Museum. Shell small. Accession no.0 5. collected by H. Sculpture of distinct spiral lines. Crest behind aperture not as heavy or as thickened as in ivheatleyi.3 2. solida in the form of the spire.5 2. Figures 2 to 5 on the same plate are of paratypes.0 6. 321196.9 5. New Species figs. plate 119.5 5.1 2.0 4. Planorhula Planorhula smithii.0 1. Locality.Descriptions of Xew Species and A^eiv Varieties 229 Parapholyx pusilla.0 Holotvpe Paratvpe Paratype positee! in the Material. Aperture roundly ovate.5 5.5 Aperture Diameter 1. aperture and base of lip. Shell Greater Lesser Height Diameter 7.

0 0.1 Diameter Diameter Aperture Height Aperture Diameter 1. 19-21. but in this variety the base and sculpture are as in typical exacuous. Menetus cooper i Plate 121. Some specimens begin with coarse growth lines and end with ribs. and the heavy (in some instances. Umbilicus with rounded edges. It somewhat resembles indianensis but lacks the axial height of that race and the latter also lacks the lip callus of smithii. Base convex near the suture but flattened toward the periphery. The carina is not acute (excepting in immature specimens) but is bluntly rounded.230 The Molluscan Fa7nily Planorhidae second labial lamella.8 1. Accession no.0 Aperture Height Aperture Diameter 1. flattened. 23 and 24 on same plate. Sculpture ranging from coarse growth lines to evenly spaced ribs.9 0. Smithii is larger than crassilabris and has a much heavier lip callus. Aperture triangular. Type Type Locality. Aperture much wider than high. and base often malleated as in cooperi. New 23-25 Species Shell acutely lenticular.0 Holotype . Menetus coloradoensis. Smith. Basal whorls separated by deep sutures. spiral lines very fine. jmrplish within and somewhat pearly.4 Diameter 4. crassilabris.1 5. two paratypes as figs. Shell of four whorls. plate 123. Planorhula smithii is named for Herbert H. The whorls tumid near the sutures. 10114. Whorls usually rounded but with slight indication of carina at shoulder as in cooperi. figs. with deeply inpressed sutures. with three whorls. Color blackish brown.6 1. Upper surface flattened. near Eldora. Swamp at head of Eldora Lake. Colorado. the swollen or tumid base near the suture. plate 122. Most shells are smaller than cooperi. not usually modified by carina. but lower lip has a long columellar region and is convex. upper and lower lips arching downward and upward to meet the periphery which is not sharp but is narrowly rounded. costate) sculpture. Aperture rounded.5 Holotype Paratype Paratype Material. 13. Umbilicus rather wide and deep. GENUS Menetus Plate 122. The new species is nearest to kansasensis but differs in having a more tumid base and a different type of rib-sculpture. Heavy spiral sculpture. Several spiral ridges below shoulder. 26 Specimens like cooperi but with heavy callus just within lip. Upper lip projects just above the periphery and is flat. Deposited in the University of Colorado ALiseum. Holotype shown as fig. rounded carina on the periphery which is impressed both above and below the periphery a short distance from the edge. New Variety fig.9 1. fig. Menetus exacuous megas has a pinched periphery. Shell Greater Lesser Height Diameter 2. Menetus coloradoensis differs from eracuous in the pinched nature of the periphery.6 2. Shell A Greater Lesser Height 1. figs. 25 on plate 122.

0 2. Museum Oregon. Lower surface flatly convex. It differs from Menetus cooperi multilineatus in the position of the carina. Menetus dalli. The lower surface is swollen over the umbili- . but the umbilicus is distinctly different in crassilabris. lenticular.0 3. One collection from Oakland was associated with typical cooperi.0 2.3 3. Type Type Locality. However. Oakland. Sculpture of rather coarse growth lines. Accession no. W^ashington. 5. of yellowish horn color. Washington. Peripheral ridge stands out and elevated above the shell as a cord. 19. One of these. and mountain swamps at Olga. H. Evidently this was based upon the presence of a lip callus. is deposited in the Museum of Comparative Zoology. and three paratypes are shown on the same plate as figs. Other collections have been determined as centervillensis. California.0 2. Shell Greater Lesser Height Diameter Diameter Aperture Height Aperture Diameter 3.0 2. the umbilical or spire region small. New figs. figs. A collection of thirty-six specimens. Lake Washington at Seattle. California. 20. crassilabris lacks a peripheral carina and cannot be allied with oregonensis.4 Holotype Paratype Paratype Paratype Material. is shown on plate 122 as fig. tional Type Type Locality. Menetus cooperi planospirus. Puget Sound. The original labels of some lots in the National Museum carried identification as oregonensis. Portland.2 3. 214590. Seattle. Washington. three whorls visible. the whorl bordered by a raised revolving ridge at the edge. Umbilicus deep. one of which is selected as holotype and the remaining specimens become paratypes. designated as holotype. sutures deeply incised.6 2. Species Plate 140. Base convex. Menetus cooperi crassilabris has been identified in collections from Colma. and 6. San Mateo County. Washington. this one being doubtfully referred to cooperi. Menetus cooperi planospirus is characterized by its very flat spire and the presence of a raised carina placed at the upper edge of the whorl. 3. Aperture modified by carina. and in having a flat rather than a convex spire. 31583. occupying one-sixth of the diameter of the shell. Holotype is shown on plate 121 as fig. Spire whorls sunken below general level. about one-fourth the diameter of the shell. and one paratype as fig. Souther. 22-24 Shell small. crossed by heavy spiral lines. Upper surface flat. Plate 122. taken by W. Of the twenty-four specimens in the type lot all but one had a callus. Orcas Island. 4. with a bluntly angular periphery. The last named were erroneously identified as Menetus cooperi calioglyptus. Harvard University. Upper surface flatly convex. Twenty-four specimens are in the United States Nabearing Accession no. New Variety 3-6 Shell with three whorls.Descriptions of New Species and New Varieties 231 Material.

.5 .

1.0 .




duct from albumen gland DC. PD. vergic sac VD. PL. body wall of gland pericardium duct of prostate prostate diverticula pedal ganglia pilasters papilla in C. tubular part of kidney U. blind sac or caeciun BY. carrefour CG. verge VC. aorta AP. duct of gland DG. vergic papilla VS. external opening of vas deferens F. cilia PRM. head heart IC. kidney L. VO. foot FL. uterus UO. vas deferens in vergic sac VF. seminal vesicle T. RD. ureter V. vergic appendage BR. growth lines GW. PG. VT. jiseudobranch PA. velum MC. new shell forming OC. FPR. rectum CR. mantle VG. RK. anus AG. opening between and preputium VP. accessory gland GA. opening of sperm canal SJ. stylet GF. saccular part of kidney SM. buccal retractor muscle BS. FO. eye EPI. VX. supporting muscle SO. liver M. canal ventricle in penial gland . PC. shell SK. jaw K. intestine or space for intestine on surface of albumen gland J. S. renal vein ridge ridge on kidney RL. superior jaw G. sperm duct ST. epiphallus-like swelling EUD. visceral ganglion VL. oesophagus ARM. PF. sperm canal SD. buccal pouch P. PR. ganglion GR. sarcobelum SC. cavity of gland connecting with gland duct IN. pneumostome PY. crop D. fiagellum RA. muscle to vergic sac vein vergic sac MX. ovisperm duct SP. nidamental gland NS. outer cup of gland OD. duct of preputium DV. gland appendage SH. wall of preputial sac STM. columella muscle CMC. albumen gland AO. salivary gland SCO. tentacle TK. lung LJ. radula RM. section through neck NG. auricle B. abdominal ganglion AL. PP. nerve X. folds in vergic sac GZ. ovotestis of liver duct into stomach BG. spermathecal duct SG. canal CF. spermatophore SPD. male opening muscle of neck MP. duct entering vergic sac E.ANATOMICAL SYMBOLS USED ON THE PLATES A. mantle collar MO. suspensarv muscles of verge VM. oviduct OE. pulmonary vein anterior retractor muscles AU. folds in wall GL. cerebral ganglion preputium preputium posterior retractor muscle CL. HT. diverticula of ovotestis diverticula of prostate retractor muscle spermatheca SB. prostate PS. mouth X^. gizzard H. muscular pads or pillars in preputium ( pilasters) MR. OLV. vergic sac MT. blood vessel BW. bulbous termination of verge NA. buccal ganglion BP. cut portion of mantle and columella muscle PRS. external opening of ureter UR. vas deferens VDV. diaphragm DA. lateral jaw LV. vagina VGL. muscular ridge or diaphragm MS. female opening FOV. collar CM. penial gland of gland cavity GX. pyloris R. opening OT. stomach SV.

Section through ovotestis. 8. Jankowski. Poland. Penial complex with muscles. 3. Section through verge and vas deferens. from A. a pond 25 miles southeast of Warsaw. 6. in length . Line near figure indicates mm. 5. The dark 2. Junction of oviduct and vas deferens to form ovisperm duct and connection with albumen gland and carrefour. and vas deferens showing a prostate canal separated from the vas deferens. oviduct. 4. spaces are folds in the anatomical Entire genital system spread out.) From 1. Head and foot showing form of pseudobranch. preparation. Vertical section through penial complex to show small size of verge.238 The Molluscan Family Plauorbidae PLATE 1 Planorbis planorbis (Linn. Section through prostate. 9. 7. Section through o\'otestis showing di\'erticula embedded 1 in the liver cells.

Illustrations of the Anatomy and of Shells 239 PLATE 1 .

of verge. 14. Vertical section of penial flagella. Vertical section of gland or appendage. Line near figure indicates cates . in length. Poland. 12. 3. 8.). a meadow attached. From pool in meadow in Jabtonna. 6. of 7. Hippeutis complanatus (Linn. 9.240 The Molluscan Family Planorbidae PLATE 2 Segmentina and Hippeutis 1-6. Gland or appendage 10. 5. complex showing relationship or appendage. Outline of albumen gland (from above). General dissection of genitalia omitting the penial complex. 4. 11. Figure 14 has same magnification as fig. 7-15. Jankowski. one for prostate (PD) and one sperm duct (SPD). Cross section of pilaster to which appendage is 2. 15. Upper part of vergic sac showing two flagella and the vas deferens. north Warsaw. 5. Two for diverticula of ovotestis. Segmentina nitida (Miiller). Upper view of cup of gland or appendage. Figures 6 and 11 are greatly enlarged. gland. Jankowski. 3. near Warsaw. 4. collected by A. 8. Vertical section of preputium showing position of gland-like appendage. in preputium. showing vergic appendage and outlet of sperm canal. of figs. 15. Cross section of prostate showing two canals. Head of animal showing form of pseudobranch and pneumostome. 1. Penial complex from the outside. From swamps on from A. Poland. and 13. 16 km. . 12 also indi- and 9. End End of verge of verge showing position of vergic appendage.size 1 mm. Line at left of fig.

Illustrations of the VD FL Anatomy and of Shells 241 PRS .

6. Head showing position and form of pseudobranch. Line near figure indicates sents size of figs. is Line at left of fig. 7. a turfy ground east of Warsaw. 7 also repre- 3 and Figure 9 same magnification as fig. From meadow near Warsaw. From Drewnica. Hermaphrodite complex. km. Cross section of prostate showing prostate duct sei)arate from sperm duct. in length. from A. mm. 8. of rectum. Segmentina nitida kowski. north- Head showing position and form of pseudobranch. Hermaphrodite complex dissected and spread 1 out. 1. Cross section of prostate and ducts of prostate and vas deferens. 4 5-10. 8. Poland.). Jan- External view of penial complex. Poland. collected by A. 9. Pseudobranch showing position Cross section of ovotestis. 3. (Miiller).242 The Molluscan Family Planorhidae PLATE 3 Segmentina and Bathyo))iphalus 1-4. Bathyomphalus contortus (Linn. 10. 4. Penial complex from the outside. . 10. 2. Jankowski. 5.

Illustrations of the Anatomy and of Shells 243 PLATE 3 .

4 and Yen). view from side showing internal canal. 3. General dissection of genitalia. Preputium cut open to show penial gland. New genus Fan Memorial Institute. Penial complex showing long duct of penial gland and long and peculiarly formed 5. Peiping. the organs separated and extended. China. Terminal end of penial gland. Albumen gland showing position of stomach. 6. Portion of penial gland showing position of ducts. Preputium cut open Verge in vergic sac. 8. from 1. 9. Figures 3 and 4 are greatly enlarged. . Line near figure indicates 1 mm.244 The Molluscan Family Plauorbidae PLATE Pingiella peipinensis (Ping Peiping. appendages. 7. imder side. 4. to show penial gland and relationship of long duct. 2. Terminal end of penial gland. China. in length.

lUustratioiis of the Anatomy and of Shells 245 .

Entire genitalia dissected and organs separated. Peiping.6. Line near figure indicates 1 mm. Spermatheca of different shape. from Fan Memorial Outline of albumen gland. China. 2.246 The Molluscan Family Planorbidae PLATE Pingiella 1. hemisphaerula (Benson).5. 4.7. 5 and Polypylis Region of the (Ping and Yen). 6. Pingiella peipinensis 1. China. 7. Figure 6 is greatly enlarged. 3. in length. 5. 2-4. pseudobranch. Section through prostate and ducts. Region of the pseudobranch. Peiping. . new genus. Polypylis Institute. Distal end of verge showing outlet of canal at the side.

Illustrations of the Anatomy and of Shells 247 PLATE 5 .

6. 12 km. Aiiisus spirorbis (Linn. 4. 5. northeast of War- saw. Line near figure indicates 1 mm. 18 km. Penial complex showing long and narrow vergic sac. and somewhat crushed. muscular and jiilasters. upper part. sperm canal at side of verge. Entire genitalia dissected and separated. Figures 2 and 5 are greatly enlarged. Jankowski. Penial complex of specimen with vergic sac shortened and club-shaped. Penial complex in vertical section showing stylet. Foot. 9. 7. 10. From stream in meadow in Jabtonna. Poland. 11. From a trench in Struga. Albumen gland. Poland. . 8. Warsaw. collected by A.). Cross section through prostate and o\ iduct.7-13. 6. 12.). Stylet. and jmeumostome. showing papiUa in preputium.248 The Molluscan Family Planorbidae PLATE 6 Anisus vortex and Anisus spirorbis 1-4. St5det at end of verge with outlet of Two diverticula of ovotestis. Section through prostate and sperm canal showing two ducts. Section through upper portion or preputium and vergic sac. stylet and outlet of sperm canal at the side of the verge. i)apilla. Jankowski. 13. by A. Anisufi vortex (Linn. 3. Pseudobranch showing relationship to rectum. north- east of 1. collected 5. pseudobranch. in length. drj' 2.

Illustrations of the Anatomy aiid of Shells 249 PLATE 6 .

5 is only for mm. collected by A. collected by A. in Muscular ring or diaphragm stylet protruding. Poland. Stylet. Poland. upper part of preputiimi showing papilla with 4. Figure under seminal vesicle shows 3. 6. 6. Anisus septemgyratus (Ziegler). Foot.size in length. 3 and . this organ stretched out. and pneumostome. Genitalia dissected and the organs separated. From small pond in Bielany Park. . pseudobranch. 4 km.250 The Molluscan Family Planorbidae PLATE 7 Anisus leucostomus and Ainsus septemgyratus 1-5. Line near figure indicates 1 fig. 1. Anisus leucostomus (Millet). left of of figs. Jankowski. Jankowski. 5. northwest of Warsaw. End of oviduct and vas deferens showing where ovisperm duct joins the ovi- duct and vas deferens. Warsaw. From turfy meadow in Drewnica. 2. Line at 4. Outline of genitalia exclusive of the penial complex.

Illustrations of the Anatomy and of Shells 251 PLATE 7 .

11. from Fan Memorial In- Section of penial complex cut open to show verge. Small figure at right shows insertion of vas deferens at junction of vergic sac and flagellum. by Dr. South Shan State. 3. 5. 5. the fiagella. China. 9. Pseudobranch and rectum. Cross section of penial gland. 6. Vertical section of penial complex showing verge. . N.11. A spermatophore. 7. Annandale. Figures 1. Burma. Penial complex. and duct. India. 8. collected 1. showing the large Peiping. Penial complex. 10. Intha capitis Annandale.10. 6. size of 11. Distal end of verge. From Inle Lake. gland. 9. Outline of albumen gland. hemisphacnda (Benson). 2. ventral view. Figure 4 is greatly enlarged. Line near figure indicates 1 mm. Polypylis stitute. dorsal view. 4. and 7 have same magnification as fig. Head of animal showing tentacles and eyes.252 The Molluscan Family Planorbidae PLATE 8 Intha and Polypylis 1-8. Dorsal view. diaphragm and pilasters. ^'entral view of entire genitalia. in length.

Illustrations of the Anatomy and of Shells 253 .

Genitalia dissected and organs separated. A. 3. from Dr. From Lares. Hoff- 4. 7. Note the small number of ijrostate 3. Puerto Rico. Cross section of portion of ovotestis to show multiple branched diverticula. 2.254 The Molluscan Family Planorbidae PLATE 9 Drepanotrema and Australorbis 1-3. Head and man. diverticula. 1 is also for . in length. W. Drepanotrema anatinitm (Orbigny). fig. Penial complex showing form of flagellum. 1. 5. 9. A. Australorbis globratus (Say). 4-10. Note branched form of prostate diverticula and multiple nature of ovotestis diverticula. Line near of fig. foot showing form of pseudobranch. 6. Puerto Rico. Hoffman. from Dr. Outline of albumen gland showing depression occupied by intestine. W. Distal end of verge showing opening at center. Pseudobranch and pneumostome. Entire genitalia dissected and organs separated. 8. Line near figure indicates left 1 mm. From Viejo. 10. One diverticulum of prostate showing multiple branching at end. Section of penial complex showing long and narrow verge and vergic sac.

Illustrations of the Atiatomy cuid of Shells 255 PLATE 9 .

Upper part of penial complex showing relationship of flagellum. pilasters. vergic sac. 4.256 The Molluscan Family Planorbidae PLATE 10 Drepanotrema hoffmani From 1. vas deferens. F. collected by Dr. vas deferens. Penial complex showing attachment of short flagellum. Puerto Rico. Hoffman. Section of penial complex showing verge. 3. Penial complex of another specimen. 5. Distal end of verge showing central outlet of sperm 1 canal. etc. Genitalia dissected and organs separated. muscle. 6. Baker Isabela. Line near figure indicates mm. pseudobranch. of Animal removed from shell showing pigmentation of mantle. Fore part of body showing head. flagellum. retractor 8. etc. in length . A. Albumen gland showing attachment of ducts. C. vas deferens. and position of stomach (STM) in body. 7. shape tentacles. etc. etc. 9. head and 2. W.

Illustrations of the Anatomy and of Shells 257 PLATE 10 .

The pseudobranch and rectum are also shown. (Pfciffer) collected by Dr. 2. 1. C. Head and Head and foot with prepiitium and verge extended from male genital opening. Line near figure indicates 1 mm. in length . Aguayo.258 The Molluscan Family Planorbidae PLATE 11 Drepanotrema lucidum From Havana. retractor muscle in body of animal in specimen with preputium and verge extended from male opening. Cuba. Penial complex with very long flagellum. Relationship of vas deferens. 7. etc. Diagram of penial complex showing preputium and \crge extended from male genital opening. 3. Genital system dissected and organs separated. foot with only preputinm extended from male genital opening. flagellum. G. 6. 4. 5. Penial complex showing flagellum. vas deferens.

Illustrations of the Anatomy and of Shells 259 PLATE 11 .

7. pneumostome. from above. Distal end of verge showing central opening of sperm canal. Orleans. and 9. Genitalia dissected and the organs separated. 5. frilled ridge 4. Line at left of fig. Body of snail showing pigmentation Albumen gland from above. 6. E. rectum. or crest over rectum. Muscular ring or ridge between vergic sac and preputium (diaphragm). Portion of genitalia at junction of oviduct and sperm duct with ovisperm duct. in length. Also entrance of albumen duct into carrefour and thence to oviduct. . 11. 10. 1 is also for figs. and of mantle. 11 is also size for fig. Same as fig. 3. Penial complex in section to show verge and pilasters in vergic sac. 2. Lme at left of fig. collected by Dr. From below. Line near figure indicates 1 mm. Penial complex showing enlargement of preputial sac. 6.. La. 8.260 The Molhiscan Family Plauorhidae PLATE 12 Tropicorbis havanensis (Pfeiffer) From Xew 1. 7. 8. showing relationship of all organs. 7. 9. Carroll Faust. Body of snail showing pseudobranch. Single diverticulum of prostate showing canal distinct from that of sperm duct.

Illustrations of the Anatomy and of Shells 261 PRS PLATE 12 .

Portion of genitalia dissected and organs separated. 1.5. W. 1. A. Line at right indicates 0. 7. 262 The Molluscan Family Planorhidae PLATE 13 Tropicorbis 1-3. Organs as in fig.. for in length. Head animal showing pseudobranch and imeumostome as well as ridge over rectum. Tropicorbis obstructus (Morelet). collected by Dr. 1 mm. 3. Tropicorbis riisci (Dunker). Albumen gland from below showing duct (DA). Line at left of 4 and 5 is both figures. Puerto Rico. Also presence from sperm duct. Genitalia dissected and organs separated. external view\ 8. from Dr.5 mm. Faust. From Lares.. Penial complex. Vas deferens showing branching to form sperm duct and prostate duct (greatly enlarged) 9. La. 2. in length. of Line near figure indicates figs. Cross section of ovotestis in gravid stage with ova in diverticula. . 4. Cross sections of prostate diverticula of prostate duct distinct 6. 4-10. 10. Hoffman. E. Cross section of ovotestis in non-gravid stage. showing bending at end. C. From New Orleans.

Illustrations of the Anatomy and of Shells 263 PRS PLATE 13 .

264 The Molluscan Family Plauorbidae PLATE 14 Gyraulus albus (Miiller) From pond 1. 11. Albumen gland from below. 3. Poland. Fore part of animal showing pseudobranch and pneumostome. a suburb of Warsaw. 6. sperm duct. Distal end of ovotestis showing grouping of diverticula. 4. sperm canal outlet at side of verge. and venation of some of these organs. external view. Penial complex. 6. for figs. Line at lower right corner 11. and general relationship of these and other organs. 5. Stylet at end of verge with 7. Genitalia dissected and organs separated. Section through ovotestis near front end. in length. etc. Jankowski. 7. . 10. oviduct. and Figure 8 line is one-sixth of 1 mm. pneumostome. Fore part of animal showing pseudobranch. 10. 2. stylet. Penial complex sectioned to show verge. in length. Portion of genitalia showing attachments of albumen gland duct. in Krolikania. Three diverticula are shown. 9. 5. from A. Section of prostate showing distinct canals for prostate and sperm duct. 9. 8. 1 Line near figure indicates of plate is mm.

Illustrations of the Anatomy and of Shells 265 PLATE 14 .

California. 8-10. from Dr. Distal end of genitalia showing relationship of ducts. Same as fig. 4. 4. Genitalia dissected and organs separated. Fore end of body showing pseudobranch. From right side. Lake Nipissing. Golden Gate Park. in length. and 7. from left side. Genitalia dissected and organs separated. From Bass Creek. Line near figure indicates corner of plate 1 mm. 9. 3. rectum. external view. Ontario. A single diverticuhun of the prostate Two diverticula of the ovotestis.266 The Molluscan Family Planorbidae PLATE 15 Gyraulus 1-7. etc. Line in upper left is for figs. 6. From Stone Lake. San Francisco. . 5. Fore end of body showing pseudobranch. 10. 7. Penial complex. 8. Hanna. 4. D. Gyraulus hirsutus (Gould). Gyraulus (Gould). A portion of the intestine is shown beneath the albumen gland. LaRocqiie. Distal end of genitalia showing connection of various ducts. received from A. 2. Canada. pneumostome. 5. G. 6. 1. vcrmicularis with duct separate from the sperm duct.

Illustrations of the Anatomy and of Shells 267 PLATE 15 .

L 2. 11.268 The Molluscan Family Planorhidae PLATE 16 Gyraulus deflectus obliquus (DeKay) From Taylor Lake. in length. collected by A. Line at indicates size of figs. Stylet. left 8. Also male opening behind tentacle. 7. External view of penial complex showing relationship of parts. Masham. Terminal diverticula of ovotestis. Distal end of verge shows sperm canal outlet at side. side. 4. Terminal diverticula of ovotestis from Section through diverticula of ovotestis. Section of prostate six diverticula from front end. 13. 10. Penial complex of specimen with bulbous termination of vergic sac. LaRocque. Canada. 1 Line near figure indicates mm. Pseudobranch showing network of blood Another form of pseudobranch. 12. showing de\-eloping ova. 15. Quebec. etc. middle and 12. vessels. 7. 9. 3. 8. 11. Section of prostate near middle. 5. Penial complex in vertical section. 6. 5. left of i)late . albumen duct. Section through portion of penial complex at junction of vergic sac and preputial sac to show thickened portion with papilla. Body and foot of animal showing pseudobranch. Distal end of genitalia showing position of sperm duct. 6. oviduct. etc. 14. Genitalia dissected and organs separated. 9.

IllustrdtioDS of the Anatomy and of Shells 269 11 12 PLATE 16 .

Figures 12. Two diverticula of ovotestis. From Wainwright by Dr. 4. From left side. Genitalia dissected and organs separated. position. 5. Two diverticula of ovotestis in gravid condition. Gyraulus parvus (Say).270 The MoUuscan Family Planorbidae PLATE 17 Gyraulus 1-5. and have same magnification as fig. Canada. Baker. from left side. near Oshkosh. Outline. From Winnebago Lake. in length. Genitalia dissected and organs separated. 1. C. Albumen gland showing attachment Three side. 10. Line near figure indicates 11 1 mm. stylet. Wisconsin. Distal end of genitalia showing duct connections. etc. 9. etc. 8. Gyraulus circumstriatus Alberta. near Mott Lake. of oviduct. Park. 5. col- lected 6. Penial complex in natural 2. Penial complex opened to show verge. . by F. papilla. 3. Portion of fore end of animal showing pseudobranch. di^'erticula of prostate with prostate duct below and sperm duct at the 11. Animal removed from shell showing position of several organs. 6-12. collected (Tryon). 10. Swales. Distal end of genitalia. Pseudobranch and pneiunostome. 7. 12.

Illustratious of the Anatomy and of Shells 271 .

Poland. side Sperm canal with 11. collected by Dr. 6-11. 1 mm. a suburb of Warsaw. Line in center of 7 is also for 8. left of fig. outlet in center of verge. End of verge showing absence of stylet and jjresence of a fleshy papilla. Armiger crista (Linn. by A. Gyrauhis similaris (F. Head showing form Albumen gland from Genitalia dissected and organs separated.). From Smartweed Lake. 4 measures one-fifth of fig. 5. From pond in Krolikarnia. Genitalia dissected and organs separated. 7. 3. 1. Distal end of genitalia showing arrangement of durt. Stjdet. Line near figure usually indicates mm. End of verge from showing relationship of 1 jiapilla and canal. near Tolland. side. Penial complex in section showing verge. collected 6. Frank Smith. Head and foot of animal showing pseiidobranch. 8. Jankowski. Line at fig. 10.5 mm. 9. Figures 10 and 11 are much enlarged. Outlet of sperm canal is 4. C. Colorado. shown at side of verge. Same line equals 0. in length. fig. .272 The Molluscan Family Planorbidae PLATE 18 Gyraulus and Arniiger 1-5. 2. and preputium. Baker). for 9. Penial complex in .'<. stylet.section showing verge without stylet. of pseudobranch.

Illustrations of the Anatomy and of Shells 273 .

6. Gyraulus latestomus F. Cross section of one diverticulum of prostate. Albumen gland from below. showing loop Penial complex. 7. in length. Anterior end of body showing pseudobranch and ])neumostome. Genitalia dissected and the organs separated. . LaRocque.274 The Molluscan Family Planorbidae PLATE 19 Gyi'aulus 1-3. Figures 1 and 7 are greatly enlarged. Distal end of genitalia from left side. 8. 4. B. Canada. Ontario. of intestine. 5. 1. Cross section of one diverticulum of prostate showing separate canals for prostate duct and sperm duct. From Calcutta. C. 2. from Dr. India. 9. Line near figure indicates 1 mm. Gyraidus co7ivexiusculus (Hutton). 4-9. 3. Baker. Prashad. Algonquin Park. From Cedar Lake. Genitalia dissected and organs separated. Pseudobranch and pneumostome. collected by A.

Illustratioihs of the Anatomij and of Shells 275 PLATE 19 .

6 represents 0. . Vergic sac opened to show verge and canal in 5.south of Warsaw. 3. 10. 11. in length. verge. etc. pilasters. . 5 km. 8. and nidamental gland. 1. Retractor muscles in tyjjical i)enial specimen. and rectum with ridge. pneumcstome. complex opened is to show position of jienial gland. Cro. retracted. 6. Line near figure indicates 1 nmi. Line at right of fig. Penial appendage. in length. 4.5 mm. One diverticulum of ovotestis with ripe ovum (greatly enlarged). Portion of head . Genitalia dissected and organs separated. Jankovvski. 9. End of verge showing Albumen gland. 7. by A. Penial complex from outside showing muscles. 1 is also for fig. 13. section through prostate.276 The MoUuscan Family Planorhidae PLATE 20 Planorbarius corneus (Linn. 2. Line at left of fig. Morysinek. 2.)- From branch of Wista River at collected 1.showing jiseudobranch. uterus. Cross section of ovotestis in gravid specimen. showing particularly the form of jirostate and its junction with the vas deferens and sperm duct. Section through appendage at C in fig. Penial appendage 12. Penial gland and appendages in extended position. Pohuid.

lUustratioihs of the Auatouiy and of Shells NG 277 .

9. State. B. and diverticula of gland. Figures 9 and 10 are greatly magnified. 3. contributed by Dr. 5.. sperm duct. 2. Under side of preputium everted through male opening in the neck of the animal. 6. 3. Pcnial complex in natural position. Rao. and 10 were published in Jour. Genitalia dissected and organs separated. Cross section of ovotestis showing Cross section of prostate. LV. Two diverticula of prostate. Portion of prostate showing relationship of vas deferens. . 1. N. S. Line near figure indicates 1 mm. North Shan Chopra and Dr. Figures 1.278 The Molluscan Family Planorbidae PLATE 21 Indoplanorbis exustus (Deshayes) From Hsipaw. Mori)h. Burma. 10. India. se\'eral gra\id diverticula. Fore part of animal showing pseudobranch and pneumostome. plates 1 and 2. collected by Dr. with preputium protruded from male opening and lying on the neck of the animal. Pseudobranch from side showing branched or pleated condition. in length. 4. H. 8. B. Pleated pseudobranch and pneumostome. 2. Prashad. 7.

Illustrations of the Anatomy and of Shells 279 PLATE 21 .

Line near figure indicates 1 mm. 9. Penial gland with cup turned back to show folds in inner cup. 6. Helisoma anccps pcrcarinatiDii (Walker). Section of vergic sac through verge. in length . 12. 10. sac. Retractor muscle of verge showing branching to vergic extended condition as . Compare with 8. 11.11. fig. Morph. Penial complex of specimen in breeding condition with preputium extended from male opening in neck. Greatly enlarged. Verge. Line near figure indicates 1 mm. 7. 1. A'ariation in form of muscles and in position of verge in preputium. 5. 2-4. Also enlarged portion of vas deferens (epiphallus). End of verge . Cross section of preputium in fig. Prostate showing attachment of prostate duct to sperm duct. Helisoma anccps latchfordi (Pilsbry). Helisoma anccps (Menke).showing papilla and sperm canal outlet. Section of vergic sac through bulbous termination of verge. 7. These figures were published in Jour. 2-4. in length. Indoplanorbis exustus (Deshayes). Figure 5 is enlarged. 12. 1.280 The Molluscan Family Plauorbidae PLATE 22 Indoplanorbis and Helisoma 5-9. Continued from plate 21. 1 on plate 21. Figure 6 has same magnification as fig. 7. 10.-^hown in Also nerve connection.

Illustrations of the Anatomy and of Shells 281 .

Diaphragm in preputial sac view from below. Canada. and rectum. 5. Baker and Leslie Brewer. in length . Helisoma anceps latchfordi (Pilsbry). LaRocque. 6. Penial complex of specimen in which gland in preputium has been pushed 8. Collinsville. lected by A. 11. pneumostome. 1. Genitalia dissected and organs separated. 9. Cross section of ovotestis. 3. upward.6. folds. From Meach Lake. Cross section of prostate showing multiple form of diverticula. Cross section of penial gland showing cup. 2. Section through diaphragm showing entrance of gland duct into vergic sac cavity just above diaphragm. Albumen gland from below. Connecticut. and duct. Line near figure indicates 1 mm. 4. Helisoma anceps (Menke). From Gillespies Pond. Exterior view of penial complex showing short duct on outside of preputium. 5. 7. col- 10. Region of the neck showing pseudobranch.282 The Molluscan Faintly Planorbidae PLATE 23 Helisoma anceps (Menke) 1-4. Section of penial complex to show position of gland in preputium.7-11. collected bj^ F. Quebec. C.

Illustrations of the Anatomy and of Shells 283 ~PRS PLATE 23 .

Young specimen with whorls. Illinois. Wisconsin. Section through penial gland in specimen in normal. Wisconsin. Penial complex in immature specimen. 15. except where otherwise stated. in length . C. non-breeding condition. Baker. upward. New York. rectum. 10. 5. Penial complex in vertical section. pncumostome. of penial IL External view 12. 8. Specimen with preputium pushed Illinois River. Penial gland somewhat contracted at cup end. near Peoria. Wisconsin. Section through penial complex in region of diaphragm. 4. Penial complex. Albumen gland from below. Penial complex. Genitalia dissected and organs separated. Region of neck showing pseudobranch. indi- cating great length. 7. Pond near Sturgeon Bay. Section through prostate and uterus near middle of gland. Pool near Green Lake. by F. Side view of albumen gland. Devils Lake. Section through ovotestis near anterior end.284 The Molluscan Family Planorbidae PLATE Helisoma 24 trivolvis (Say) Collected from Oneida Lake. 13. Specimen with portion of preputium containing gland pushed upward. Line near figure indicates 1 mm. L 2. 3Vi> 3. 6. Section through penial gland showing folds in cup. complex showing duct uncoiled and stretched etc. 14. out. 9.

Illustrations of the Anatomy and of Shells 285 PLATE 24 .

penial Penial complex with preputium pushed upward. Penial complex with vergic sac drawn to side of preputium. Canada. Baker).286 The Molluscan Family Planorbidae PLATE Helisoma pilsbryi 1-7. Section through prostate and uterus. Ontario. in length . collected by A. 2. 8-13. 10. Line near figure indicates 1 mm. collected by A. LaRocque. 25 (F. Penial complex of immature specimen. Penial complex of specimen with large preputium. Outline of genitalia with organs separated. Baker) Lake. Outline of genitalia dissected and organs separated. Cahn. Canada. Ontario. R. C. R. 12. 3. 13. Unless otherwise stated specimens are from Basswood River Rapids. 8. A. C. 11. C. Canada. C. showing length gland duct. 4. Penial complex showing great length of penial gland duct. 5. From Rideau River of Rapids. Cahn. Baker. Wisconsin. Ottawa. 7. Baker. Penial complex from outside. by Dr. Pseudobranch and pneumostome. 6. by F. Penial comjilex of specimen with preimtium pushed upward. collected From Bamiji Lake. Cross section of penial gland. Helisoma pilsbryi injracarinatum F. in normal position. Barron County. 9. Pseudobranch and adjacent organs. Helisoma pilsbryi (F. From Chetek collected 1.

Illustrations oj the Anatomy and of Shells 287 PLATE 25 .

Outline of genitalia dissected and organs separated. 2.288 The Molluscan Family Planorhidae PLATE 26 Helisoma 1-5. Penial complex with penial gland duct pulled out to show great length. collected by F. Line near figure indicates 1 mm. Helisoma iruncatum (Miles). the preputium pushed upward. is 12. Baker. position of pseudobranch. near middle of gland. 8. From Paul Lake. Genitalia dissected and organs separated. 3. Penial complex with penial gland duct in normal position coiled on surface of preputium. Cross section through prostate and o\iduct. From Winnebago Lake. Cross section through oviduct. and rectum. Helisoma horni (Tiyon). pneumostome. Cross section of penial gland. 1. 4. 5. C. Cross section of ovotestis near anterior end. 6. 9. Penial complex with penial gland duct spread out. collected by Prof. D. 11. Preputium partly pushed 13. Outline of penial gland. 10. and nidamental gland. Wisconsin. British Columbia. 7. 6-13. Cross section of prostate and oviduct near anterior end. in length . near Oshkosh. sperm duct. Rawson. Penial complex in breeding condition. S. Head showing upward.

Illustrations of the Anatomy and of Shells 289 PLATE 26 .

collected by Dr. 14. Section through ovotestis near anterior end. From Sturgeon Lake. Section through diaphragm showing relationship of penial gland duct to vergic sac. C. Penial complex of Helisoma campayiulatum sniithi (F. showing variation in form. Penial complex. Ontario. Also variation in muscles. Baker. Cahn. 13. and rectum. Canada. 11. Michigan. Helisoma campaiudatinn canadense F. Lake. especially the preputium.290 The Molluscan Family Planorbidae PLATE 27 Helisoma campamdatum (Say) From Oneida 1. Penial complex with penial gland duct uncoiled to show length. Pseudobranch. 2. C. 3. A. 4. R. near Brewerton. Section through prostate near middle of gland. Albumen gland from below. Genitalia dissected and organs separated. 5-7. C. Penial complex showing penial gland duct coiled between preputium and vergic sac. Baker). Section through penial gland. collected by F. 8. New York. Section of penial complex showing relationship of penial gland to verge. from Douglas Lake. 12. 10. -14. Baker. Line near figure indicates 1 mm. 9. in length . pneumostome.

Illustrations oj the Anatomy and oj Shells 291 PLATE 27 .

9. 3. Section through penial gland. Helisoma scalare Andrews. From Lake Butler. From Tooele Co. Genitalia dissected and organs separated. received from Dr. Cross section of prostate near middle of gland. in length . 6. Penial complex with preputium pushed upward. A. sac. (Jaj'). 4. collected by J. Pseudobranch and pneumostome. E.292 The MoUuscan Family Plauorhidae PLATE 28 Helisoma 1-5. 6-14. Helisoma oregoncnse (Tryon). Line near figure indicates 1 mm. Florida. 11. 7. Hen- Penial complex showing long penial gland duct. the penial gland duct extended. Penial complex in normal position.. 10. 8. Section through ovotestis near anterior end. 13. Penial complex in section to show relationship of organs. Section through prostate near middle. Genitalia dissected and organs separated. 5. 12. Utah. 2. Penial gland viewed from above. 14. derson. Entrance of penial gland duct into vergic Section through penial gland. 1.

Illustratio))s of the Anatomy and of Shells 293 PLATE 28 .

LaRocque. showing penial gland and verge everted. by E. Tennessee. complex with penial gland duct uncoiled. 2. 6. . In 10 line equals 0. Faust. macrostomum (Whiteaves). 9.specimen in normal condition. 3. 13. Helisoma Penial complex. Same locality as fig. 4. Baker. 8. Helisoma chautauquense F. collected by A. From Lake Gauvreau. Baker) ^ lentum m Say. From Xew Orleans. collected by F. C. Penial complex in section showing preputium pushed upward and penial gland duct vmcoiled. Peoria. 11. Verge much retracted. 14. Head and neck Verge in fig. Helisoma lentum (Say). From Cambridge. Penial complex with penial gland duct coiled on preputium. Penial complex of immature specimen. New York. lip-like processes sur- 10. 14.. 7. C. collected by F. Penial complex showing large size of preputium. Penial Penial 15. 14-16. Co.294 The MoUuscan Family Planorbidae PLATE 29 Helisoma 1-6. Wisconsin. 20. 22. C. Helisoma trivolvis pseudotrivoivis (F. of animal rounding the opening. Illinois. Penial complex with penial gland and verge everted.5 mm. C. Penial complex of adult . From St. Joseph. C. Penial complex with penial gland duct uncoiled. fig. Illinois. 12 removed from penial complex. 17. Exterior 21. Penial complex in section showing penial gland putium. Champaign 1. 5. 19. Penial complex with penial gland everted. 16. 18. complex with preputium pushed upward. Helisoma lected trivolvis fallax J. End of verge showing exit of sperm canal with small. trivolvis Canada. Baker. From Bayfield. Verge fully extended with sections in two places as indicated. in length. Baker. Massachusetts. view of penial gland. Baker. Louisiana. Exterior view of penial gland. collected by F. 19-22. 17. Line near figure indicates 1 mm. collected 7-13. trivolvis From Reelfoot Lake. Neck of animal showing penial gland everted. (Haldeman). col- by W. Clench. natural position in pre- Penial complex with penial gland duct spread out to show length. 12. From Illinois River. C. Penial gland in section.18. From Chautauqua Lake.

Illustrations of the Anatomy and of Shells 295 .

of Form of verge in a specimen from Smartweed Lake. Head showing pseudobranch. 12. north 9. Section of vergic sac From Smartweed Lake. Penial complex with vergic sac cut open to show form of verge. Carpenter). Penial complex with penial gland duct uncoiled. near Gypsum. Penial complex with vergic sac cut open to 17. Devon. 15. 11. Cross section of prostate about midway of its length. Penial complex showing short penial gland duct in immature specimen. 14. Henderson. Colorado. collected by J. (Ingersoll). Helisoma subcrcnalum (P. Penial complex with penial gland duct uncoiled. Colorado. 5. 13. Collected by J. 6-8. From one mile west Montana. of 10-19. in length . pneumostome. Colorado. near Smartweed Lake. Section through penial complex showing verge and penial gland. Albumen gland from below. Figures 17-19 are from specimens collected at Cottonwood Pass. Genitalia dissected and organs separated. Helisoma plcxatum 1. Penial complex in section showing form of penial gland and verge. Cross section of penial gland. From Columbine Lake. Line near figure indicates 1 mm. Colorado. 18. Teton River. Colorado. Genitalia dissected and organs separated. Idaho. Penial comiilex in section to show form of gland. 19.296 The Molluscan Family Planorbidae PLATE 30 Helisonia 1-9. 2. From Rexburg. showing form of \erge in different individuals. Henderson. 16. 4. and rectum. From Columbine Lake. P. 3. show verge. From Lower Los Lagos. 10. Section of penial gland.

Illustrations of the Anatomy and of Shells 297 .

shell 12 nun. specimen about one-fourth grown. Baker. shell 23 mm. Cross section of prostate and uterus at X Genitalia dissected and organs separated. Baker. 11. Baker. Penial complex in specimen somewhat older than fig. in length . Helisoma corpulent lun (Say). C. 6. C. in diameter. A. 17. Penial complex and penial gland duct in natural position. shell 16 Penial complex of adult animal. From Lake Vermilion. From Kahnipiminanicock Lake. collected 19. 3. Penial complex of specimen with shell 14 Croix. 15. Cahn. Minne. Penial 22. Section of penial gland showing cup and penial gland duct. Helisoma corpulentinn multicostotum F. From Knife Lake. mm. Penial complex of immature specimen. 14-18. From Lake La Croix. Penial complex of animal about half-grown. Penial complex of immature specimen. Penial complex. in diameter. 20. in Duct short. collected by Dr. diameter. Canada. mm. 3. Cahn. 2. Minnesota. 18. Canada. 16. Line near figure indicates 1 mm. R. Helisoma whiteavesi F.5 Penial complex of specimen almost adult. mm. R.5 Penial comjilex of half-grown animal. mm. collected by Dr. in diameter. shell 5. in diameter. 14. C. by A. From Greenfield Pond. uncoiled. Paul Bartsch. 5. collected by F.298 The Molluscan Family Planorbidae PLATE 31 Helisoyna 1-7.?ota and Lake La Croix. R. shell 14. Penial 21. Penial complex with preputium pushed upward and penial gland duct imcoiled.5 10. From Lake La 8-13. mm. 12. 19. 1 mile south of Wil- mington. 7. Ontario. Penial complex showing preputium pushed upward. Ontario. Baker. Cahn. Helisoma magnificiun (Pilsbry). Penial complex of another half-grown specimen. Penial collected by Dr. C. 4. Cross section of ovotestis. in diameter. Ontario. 1. shell 14. Cross section of penial gland. Lake. complex with penial gland duct uncoiled. gland in section. in diameter. 13. 9. the penial gland duct uncoiled. Helisoma corpulentum vermilionense F. From Knife Lake. From Knife Lake. complex in section. Duct not shown. Duct imcoiled. North Carolina. Duct long. From Mille Lacs. in fi^. Penial complex with small vergic sac. A. Verge of different shape. 20-22. 16. 8. From Knife From Knife Lake.

Illustrations of the Anatomy and of Shells 299 PLATE 31 .

pneumostome. 17. 14. Cross section through prostate. Klamath Lake. 4. Pseudobranch. and nidamental gland. 3. Puget Sound. Genitalia dissected and organs separated. 5. 8. P. in length . three miles northeast of Bakersfield.10. Helisoma traskii (Lea). C. Red- by Dr. 2. Baker. uterus. Position of verge in vergic sac. wells. Section through penial gland. lands. Helisoma subcrenatum (P. From Wacos Bay. California. 6. Helisoma tenue californiense F. From pools from artesian Shandon. Section of prostate and uterus. Hanna. From Prospect Park. Penial complex with preiiutium pushed upward. Helisoma collected by J. 13. Penial complex cut open to show verge. 16. collected 11. (Cooper). Variety. collected by Dr. Pseudobranch. collected 7. Henderson. From slough near Kern River. Line near figure indicates 1 mm. Hendenson. Oregon. 15. 12. near Genitalia dissected and organs separated. Penial complex in normal position with penial sland duct uncoiled. California. Penial complex with penial gland duct uncoiled.300 The Molluscan Family Planorbidae PLATE 32 Helisoma 1-7. occiclentale 9. Ingles. G. G. Berry. and rectum. Carpenter). From Pass Lake. 11-17. pneumostome. 9. California. 1. S. 8. Penial complex cut open to show form of penial gland. Washington. by L. Penial complex with preputium pushed ujjward. Penial gland viewed from the outside. D. Penial complex with vergic sac at summit of preputium. collected by J. Fidalgo Island. 10. Bifid tentacle. and rectum. S.

Illustrations of the Anatomy and of Shells 301 PLATE 32 .

collected by T. penial gland protruding from male opening. Florida. Gainesville. High Springs. in length . fig. shell 6. 10. From Santa Fe fig. Penial gland. 19. Exterior view of penial gland. Head and neck with as fig. 9.302 The Molluscan Family Planorhidae PLATE 33 Helisoma 1-7. collected by T. 16-19. Penial complex of normal specimen. Florida. A very wide example of penial complex. Penial complex showing swollen duct of penial gland and position of verge in vergic sac.5 mm. Penial complex. Alachua Co.14. half-grown specimen. 5. very wide example. 12. Penial complex of immature specimen. Penial gland in section showing the long. 12. Same locality as fig. Penial complex with swollen preputium. A normal form showing short penial gland duct. Penial gland in section. Helisoma duryi (Wetherby). 8-10. Shay. F. Van Hyning. From Van Hyning. 12. River. From Silver Springs. F. Same locality as 12. 4. 16. Florida. 15. collected by W. Penial complex. near Paines Prairie. 2. Helisoma duryi Van Hyning.. 3. From Tamiami Trail. Line near figure indicates 1 mm. Florida. 13. Florida. Genitalia dissected and organs separated. 6. 15. 1. 7. collected by Penial complex. 8. Variation in form. A 11. by W. near typical form. 9. Penial gland of Same locality as fig. Penial complex from exterior with penial gland duct uncoiled. Same locality as 18. 14. Penial complex with preputium pushed upward. shallow cup.13. collected From canal at Boynton. Penial complex with penial gland protruding from male opening. Helisoma duryi normale Pilsbry. partly contracted. Penial complex. Helisoma duryi normale Pilsbry. Shay. From near Gainesville. varying toward intercalare (Pilsbry). T. 40 miles west of Miami. 12. eiidiscus Pilsbiy. Florida. in diameter. Same locality 17. 12. 11.

Illustrations of the Anatomy and of Shells 303 .

Albumen gland from below. Section through penial gland to show duct and folds. IL Vagina and spermatheca 12. of different form. Genitalia dissected and organs separated. 2. and duct to albumen gland. 2. Line near figure indicates 1 mm. Smith From Upper Klamath 1. A. sperm duct.304 The Molluscan Family Planorbidae PLATE 34 Carinifex ponsonbyi E. ^'ertical ridges 8. 4. Oregon. 6. Cross section of ovotestis near proximal end. Portion of genitalia showing relationship of oviduct. 7. Cross section of prostate and uterus. Lake. Figure 12 has same magnification as fig. Cross section of prostate near distal end. 9. Penial complex cut open to show verge and penial gland. in length. ovisperm duct. 3. Henderson. . 10. Shows unusual position of spermatheca. Head and foot showing pseudobranch and pneumostome. Cross section of preputial sac to show or pilasters. 5. Penial complex showing muscle system and short penial gland duct. collected by J.

Illustrations of the Anatomy and of Shells 305 PLATE 34 .

306 The Molluscan Family Planorbidae PLATE 35 Carinifex 1. 6. From Jackson Lake. Simimit of verge showing attachment of retractor muscles. 7. From Upper Klamath Lake. 1. 4. 1 Line near figure indicates mm. collected by 2. sperm duct. Genitalia dissected and organs separated. Wyoming. gland duct. 6. Portion of genitalia showing relationship of oviduct. Henderson. Smith. Carinifex ponsonbyi E. etc. Another form of in fig. A. Carinifex jackso7iensis J. in length . Penial complex showing variation in muscle system. ovisperm duct. Penial complex showing multiplication of retractor muscles and short penial 3. Henderson. Same organs as Albumen gland. 5. J. spermatheca. Pseudobranch and pneumostome. 9. 8. 2-9. Oregon. from above.

Illustrations of the Anatomy and of Shells 307 PLATE 35 .

Cross section of ovotestis about 5. At 11. 3. From Crater Lake. Cross section of ovotestis toward posterior end. collected by 1. Baker. Line near figure indicates 1 mm. and nidamental gland. From Upper Klamath Lake. Oregon. Pseudobranch and related organs. in length .308 The Molluscan Faiyiily Planorbidae PLATE 36 Parapholyx 1-9. J. Penial complex cut open to show verge and penial gland with its short duct.12. 4.11. 10. 9. Henderson. 7. 6. collected Vergic sac and upper part of preputium showing \-erge. Henderson. Parapholyx cffusa diagonalis by J. muscular ring or diaphragm. Oregon. midway of its length. shell. C. 10. Penial complex showing nerve and blood vessels. Verge showing central position of J. uterus. Genitalia dissected and organs separated. Section of penial gland showing pipe-like cup and canal. Whole animal removed from 2. Henderson. and entrance of penial gland duct into diaphragm. 8. Penial complex showing retractor muscles and short penial gland duct. Cross section of prostate. Parapholyx ej]usa klamathcnsis F. X gland is cut open to show canal. exit for the sperm canal. 12.

Illustrations of the Anatomy and of Shells 309 PLATE 36 .

Genitalia dissected and organs separated. 11. 10. Wisconsin. sperm duct. in length. internal duct. Section of penial complex showing penial gland spread out and opened. 6. Section of penial gland. near Madison. 8.xtended laterally. Section of ovotestis to show paired condition of diverticula. Penial complex showing muscles and general form of vergic sac and preputium. 5. 2. 1 and 2. End of verge to show outlet of sperm canal. appearing as though digitate. 12. with a short. collected 1. Head and foot showing pseudobranch and pneumostome. 7. by J. 1 mm. 4. Long line at right of plate for figs. . 6.310 The Molluscan Family Planorbidae PLATE 37 Planorhula armigera (Say) From Murphy Creek swamp. immature specimen showing wide form of preputium. Long line at left of plate is for is and 8. 3. 7. 9. Verge in vergic sac. Portion of genitalia showing relationship of o\iduct. and carrefour. Morrison. with albumen gland in natural position. which are e. Penial complex of small. E. Section of prostate near middle of gland. Section of prostate near middle. ovisperm duct. Line near figure indicates figs.

Illustrations of the Anatomy and of Shells 311 PLATE 37 .

Section of prostate near distal end. Also from Quatsino Sound.2. Section of ovotestis.4-10. collected 1.312 The MoUuscan FainUy Planorbidae PLATE 38 Menetus 1. 4. Section through penial complex showing natural position of penial gland. 5. 15. Penial gland with cup expanded. British Columbia. Section of prostate about midway of its length. 2. 7. Penial complex. Walton Clark. Exterior view of penial complex. Hanna. 14. received from Dr. 14. 6. Line near figure indicates 1 mm. California sjiecimen. 3. and 15. Entire prostate. British Columbia specimen. 3. End of verge showing central outlet of sperm canal. D. British Columbia specimen. 6. 2. 13. G. Peake. 14 is also for figs. 8. collected 3. Menetus opercularis (Gould). City. Cali- fornia. by H. Head showing position of pseudobranch. British Columbia specimen. From Mountain Lake. Line at left of fig. . 11. 9. 10 is also for figs. From Crescent by A. Menetus cooperi calUughjptus (Vanatta). San Francisco. British Columbia specimen. Genitalia dissected and organs separated. Line at left of fig. 5. Albumen gland in natural position overlying stomach. Cross section of prostate and uterus near front end. California. 7 and 8.11-15. Penial complex from the outside. Albumen gland. 12. California specimen. 10. in length.

Illustrations of the Anatomy and of Shells 313 .

Puget Sound. collected by Dr. Penial complex cut open to show verge. Penial complex of specimen badly infested with Trematode worms (flukes).8. British Menetus cooperi callioglyptus (Vanatta). Line near 9 also for fig. of jiseudobranch. Head showing form specimen. 1. 7. 2. Also form of penial gland in preputium.314 The MoUuscan Family Planorhidae PLATE 39 Menetus 1-6. 9. penial gland with duct. C. 6 is also 5 and 8. Washington. Line near figure indicates for figs. Head showing pseudobranch. Dale Foster. 5. California specimen. See plate 38. T. 6. Vancouver Columbia. Seminal vesicle and ovisperm duct. fig. California. and position of retractor muscle. California specimen. collected by Arthur Peake. 7. preoccupied). California specimen. enlargement of its short. Note peculiar form. Island. British Columbia specimen.9. 10. 8. Baker {= planulat us Cooper. From Quatsino Sound. California specimen. Penial gland in section. Genitalia dissected and organs separated. in length. Penial gland attached to wall of preputium.10. Penial complex showing enlargement of vas deferens as it enters the vergic sac to form an epiphallus. Columbia 4. Menetus cooperi F. 7. 3. British Penial complex in natural position under female complex. California specimen. 1 mm. . Also from Crescent City. internal \as deferens (epiphallus). From Orcas Island. Line above is fig.

Illustrations of the Anatomy and of Shells 315 PLATE 39 .

Menetus dilatatus (Gould). 6. Line at right of fig. 1. 2. in cylindrical form. fig. Texas specimen. Penial complex in natural position.5 mm. Genitalia dissected and organs separated. Line at left of fig. External view of penial complex. 9. vesicle. Texas form. 12. Ovisperm duct and seminal Missouri specimen. Missouri. 4. 9. Baker. cut. 10 equals 0.8. 11. Small specimen shows re\erse side 12. 11. Texas specimen. C. near Stanton. From six miles northeast of Dallas. collected by Leslie Hubricht. Diverticulum of ovotestis with ova in upper part.6. 10. by F. Penial complex shown in gland. Also from Meremac River. collected 3. Massachusetts. by Dr. Texas 10. P. in length. 8. 5. Texas. All figures are greatly enlarged. ConnectiBaker and Leslie Brewer. Penial complex cut open to show verge and form of i)enial gland. Also from Hyannis. A broad form. 12 equals 1 mm. Massa- chusetts. 1. .y F. C. specimen from Union\iIle. Pseudobranch Pseudobranch of prostate of of specimen from Hyannis. Penial complex with preputium pushed u]>waid. 4 cut open to show position of verge and i)enial 7. the preputium inflated.2. in length. Texas specimen. Penial gland. 7. and vas deferens.316 The Molluscan Family Plauorhidae PLATE 40 Menetus 3-5. E. collected b. Pseudobranch specimen. collected Menetus sampsoni (Ancey). Cheatum. From pond in field near Unionville.

Illustrations of the Anatomy and of Shells 317 PLATE 40 .

4. Line near figure indicates 1 mm. Line at left of figs. Baker. Penial complex. Albumen gland from the side. 10. Swollen gland causing bulge in preputium.318 The Molluscan Family Planorbidae PLATE 41 Promenetus exacuous (Say) From Winnebago Lake. Genitalia dissected and organs separated. 5. . 3. 9. C. Ovotestis viewed from left side of animal. Cross section of ovotestis about midway of its length. showing diverticula. Wisconsin. 6. 8. 4 and 5 is for both of these figures. jienial animal showing verge and gland ]>rojecting from the male genital opening. Head and neck Head of of animal showing pseudobranch. showing a single retractor muscle. in length. collected by F. Penial complex. 7. 10. L 2. Cross section of prostate near anterior end. Figure 7 has the same magnification as fig. external view. Penial complex opened to show verge and penial gland. near Oshkosh.

Illustrations of the Anatomy and of Shells 319 PLATE 41 .

Penial complex in natural position. Penial gland in preputium which has been cut open and the pilasters separated. collected by Dr.3. by Dr. 5. 2. and peculiar ring near end. Rawson. 7.5. 3. bia. and 13 also for and 11. 9. Penial gland from above showing trough-like duct leading to muscular ring diaphragm at upper part of preputium. collected 1. 8.ship of ducts. sperm canal. Promenetus exacuous (Say). An unusual form with exit for split diverticula. 9. 6. Received from Canadian National Museum. Cross section of ovotestis near posterior end. Stiles.13. Terminal end of verge showing central jiendage.8-11. 10.7. 10. 12. Cross section of ovotestis near middle. Alberta. S.320 The Molluscan Family Planorbidae PLATE 42 Promenetus 1. Alberta. fully Pseudobranch expanded. Side view of preputium showing penial gland and end of verge passing through the diaphragm. From Paul Lake. small fleshy ap- 6. 8. 13. Promenetus umbilicatcUus (Cockerell). . Cross section of prostate. Pseudobranch and jineumostome. 1 mm. Head or of animal showing pseudobranch. From Wainwright Park. Received from Canadian National Museum. 11.4. Promenetus exacuous megas (Dall). 7. Line near figure indicates figs. 4. D. Portion of genitalia showing relation. collected 2. Wainwright Park. is in length. Stiles. Section of penial gland. British Colum- by Dr.12. Vertical line bordering figs. From Mott Lake.

Illustrations of the Anatomy and of Shells 321 .

in length. . Penial gland showing channel-like duct. Section of penial gland. 7. also carrefour. viewed from above. Wainwright Park. 8. Penial gland viewed from the side showing to 10. 6. Cross section of ovotestis. 6. 11. Cross section of prostate and uterus about midway of length. 4. Albumen gland from below. Figures 5. 1. Alberta. etc. diaphragm.322 The Molluscan Family Planorbidae PLATE 43 Pronienetus umbilicatellus (Cockerell) From lake southwest of Dr. 10. Portion of genitalia showing union of oviduct and sperm duct. collected by Received from the Canadian National Museum. mm. Figures 9. Genitalia dissected and organs separated. and 8 have 12 same measurements as fig. Penial complex opened to show penial gland. and have same measurements as fig. 7. verge. Head showing pseudobranch and pneumostome. 3. 9. 3. Mott Lake. Note developing ova. 5. how gland is folded and base attached preputium at lower end. Penial complex in natural position. Termination of \erge showing small fleshy appendage Line near figure indicates 1 at end. 11. Stiles. 12. L 2.

Illustrations of the Anatomy and of Shells 323 .

C. Louisiana. About middle. Near middle. Canada. Near middle. New York. H. H. //. Ontario. Lake H. tenue H. Pilsbry. (Inger. truncatum (Miles). H. Shandon. Oneida Lake. Baker. scalare (Jay). anceps (Menke). H. occidentale (Cooper). t. Lower end. C. Lower pilsbryi (F. Baker. Connecticut. Co. //. (raskii (Lea). Baker. Near lower end of kidney. Butler. duryi normale Pilsbry. a. Meach Lake. corpulcntum (Say). Arrow indicates point at which section was made. 2. Wisconsin. Near middle. subcrenatnm middle. Winnebago Lake. of the 14. (P. Near middle. H. 18. Woods. 26.soll). Collinsville. Carpenter). H. Ontario. North Carolina. 9. Ontario. Bayfield.. latchfordi (Pilsbry). lentum (Say). Lake Butler. Florida. H. Klamath Lake. California. H. Gillespies 8. 15. H. infracarinatum F. Lake scalare (Jay). H. Klamath Lake. C.324 The Molluscan Family Planorbidae PLATE 44 Kidney of Helisoma 1. H. multicostntum F. Line near figure indicates 1 mm. calij orniense F. magnificum (Pilsbry). Pond. duryi normale Canal at Boj-nton. Baker). 24. Oregon. Buena Vista Canal. Oregon. C. . Florida. Oregon. Near middle. Sections of kidneys 6. Lake Near middle.. Near middle. end. Canal at Boynton. Ontario. trivolvis (Say). Florida. P. About middle. Lower end. Paul Lake. anceps (Menke). Near middle. Above middle. oregonense (Tryon). British Columbia. Columbine Lake. 17. depressum F. trivolvis (Say). Wisconsin. H. New Orleans. Collinsville. macrostomum (Whiteaves). Near Middle. of the Woods. Oneida Lake. 5. C. 7. 4. p. 3. California. Connecticut. Basswood River Rapids. corpulentuyn (Say). Near Wilmington. 12. H. IL H. horni (Tryon).. H. H . Florida. Near middle. H. Baker. Washington. Colorado. Quebec. in length. 19. Skagit Near 22. Near middle. Tooele H. 10. Crooked Pine Lake. Lake Chetek. McMurray. t. At middle. o. Near middle. New York. 20. c. 21. H. H. H. Gillespies Pond. Wisconsin. plexatum H. Canada. 25. 16. 13. 23. Co.

lUustratioiis of the Anatomy and of Shells 325 PLATE 44 .

8. Australorbis glabratus (Say). Near Oshkosh. Section at E in in fig. H. Planorbarius corneus (Linn. 9. IL Australorbis glabratus (Say). 4. Baker. C. etc. 16.. Section at Australorbis glabratus (Say). H. E. California. wisconsinense (Winslow). campanulatum wisconsinense (Winslow). fig. Indoplanorbis exustus (Deshayes). . Carinifex ponsonbyi E. Section 12. in in I. 3. F. Australorbis glabratus (Say). B. Four spherical bodies from ridge of kidney. 14. Section H. Line near figure indicates 1 mm. Smith. tenue californie7i. Greatly enlarged. part. fig.). Section at at D C B I. C. Oregon. fig. D. Baker. Section through lower end. 5.326 The Molluscan Family Planorbidae PLATE 45 Kidneys of Planorbidae 1. campanulatum. A in L L fig. Planorbula armigera (Say). Puerto Rico. tenue californiense F. H. Section through upper H. Planorbula armigera (Say). Carinifex pojisonbyi E. Helisoma campanulatum wisconsinense (Winslow). C. in length. indicate points of kidney through which sections have been made and are figured. From Poland. A. 13. Section through middle. 19. 4. L fig. Letters A. 6. Baker. Klamath Lake. Wisconsin. A. Section near middle. tenue californiense F. 15. Section at B A fig. Section through lower end. 2. 17. Smith. From India. Australorbis glabratus (Say). in in 10. at C. Section near middle. 7. Section near middle. Section at Australorbis glabratus (Say). 4. Near Redlands.

Illustrations of the Anatomy and IN of Shells 327 PLATE 45 .

). C. 6. Klamath Lake. Tropicorbis 12. Menetus Menetus dilatatus (Gould). 5. Section near middle. Near Warsaw. in length.). Promenetus exaciious (Say). Baker. Hippeutis complanatus (Linn. Oregon. long.).3 mm. Tropicorbis havanensis (Pfeiffer). 2.). indicated by arrow.).). Baker. Line is 0. dilatatus (Gould). by arrow. 0. C. Planorbis planorbis (Linn. Connecticut. Puerto Rico. Line equals 18. Drepanotrema hoffmani F. Baker. Hippeutis complanatus (Linn. Poland. (Dunker). M. near Oshkosh. P. British Columbia. Section at jioint 20. C Baker. Drepanotrema lucidum Lsabela. 3. . 13. cooperi callioglyptus (Vanatta). Planorbis planorbis (Linn. 7.5 19. n. mm. Near Warsaw. Section near middle.3 long. Section near middle. Poland. Vancouver Island. 9.5 (Linn. 16. 4. Section near middle. in length. Wisconsin. 8. 15. riisei riisei 10. 14. Poland. Winnebago Lake. Bathyomphalus contortus Bathy<>mi>halus contortus equals 0. Tropicorbis (Dunker). mm. Puerto Rico. Near Warsaw. Section near middle. effusa klamathensis F. Except where otherwise mentioned the indicates 1 line near the figure mm. at ])oint indicated Drepanotrema hoffmani F. (Linn. Line mm. C. Promenetus exacuous (Say). in length. Menetus cooperi callioglyptus (Vanatta). Unionville. Section (Pfeiffer). Line is 0. Section near middle. Section near middle. Section at point indicated by arrow. Section near middle. 17.328 The Molluscan Family Planorbidae PLATE 46 Kidneys of Planorbidae 1. Parapholyx effusa klamathensis F.

Illustrations of the Anatomy and of Shells 329 .

5. Line equals 0. C. 14. hemisphaerula (Benson).5 mm.). India. Line equals 0. Section near middle. 16. 19. vortex (Linn. Section near middle.). 10. Intha capitis Annandale. Section below middle. Canada.5 mm. 26. Section near middle. South Shan State. Line equals 0. 47 of Planorbidae Gyraulus parvus (Say). Gyraulus siniilarls (F. Line equals Gyraulus vermicularis (Gould). Gyraulus deflcctus obliquus (DeKay). 8. Wisconsin. Near San Francisco. Section near middle. ni length. .sconsin. Armiger Anniger crista (Linn. 6. Section below middle. Section near middle. 5. IL Gyraulus Section below middle. Line equals mm. Section near middle. Gyraulus circumstriatus (Tryon). 22. 9. Near Quebec. Line equals Gyraulus convexiusculus (Hutton). 7. 20. 21.). 0.5 23. Poland. Wi. 24. Gyraulus hirsutus (Gould). Line equals 0.5 Anisus spirorbis (Linn. Section near middle. Line equals 0.5 nnu. Gyraulus vermicularis (Gould). crista (Linn. Section near middle. Gyraulus convexiusculus (Hutton). Polypylis hemisphaerula (Benson). Ayiisus spirorbis (Linn. Section near middle. Baker). Gyraulus deflcctus ohliquus (DeKay). Section nun. 13. California.5 stated the line near the figure indicates 1 mm. Poland. 2. below middle. Near Wansaw. 0. 18. Line equals 0. Polypylis 25. China. India. 27. 15. nun. latestornus F. Pohmd. fig. (Miiller). mm. Section near middle.). Wainwright Park. Near Warsaw. Winnebago Lake. 3.). Ayiisus Near Warsaw.5 mm.330 The Molluscan Family Planorbidae PLATE Kidneys 1. Anisus leucoslomus (Millet). Gyraulus hirsutus (Gould). Section near middle. Canada. nmi.5 Gyraulus circumstriatus (Tryon). Line equals 0. Peiping. Baker.5. Alhoifa. Segmenlina nilida Segmentina nitida (Miiller). 0. Intha capitis Annandale. Figures 6 to 11 have same magnification as 12. 17. Anisus vortex (Linn. C. Burma.). 4.25 Linless otherwi. Gyraulus parvus (Say).

Illustrations of the Anatomy and of Shells 331 PLATE 47 .

(Pfeiffer).). Region of stomach from above. Entire digestive trivolvis (Say). 15. Nerve ring of Helisoma trivolvis (Say). Bathyomphalus contortus (Linn. 7. Region of stomach from above. AuMralorbis glabratus (Say). 4. Planorbis planorbis (Linn. of 6. From above. system except liver. IL Drepanotrema hicidum 12. Nerve 18. Region of stomach from above. Region of stomach from above. Region of stomach from above. 3. Region stomach from above. 2. 8. Nerves and blood from left cerebral ganglion. Gyraulus albus (Miiller). Region of stomach from below. Line near figure indicates 1 mm. of Segmentina nitida (Miiller). Region Promenetus exacuous (Say).). Promenetus exacuous (Say). Planorbis planorbis (Linn. Carinifex ponsonbyi E. vessels to penial complex. stomach from below. Helisoma Region of the stomach from below. from above. Armiger crista (Linn.).). Region of stomach from below.332 The Molluscan Family Planorhidae PLATE 48 Digestive System of Planorbidae 1. 10. Hippeutis complanatus (Linn. 13. Gyraulus parvus (Say). A. 14.). Region of stomach from below. Helisoma anceps (Menke). 9. in length . 16. 17. Smith. Portion of digestive system from above. 5. Helisoma trivolvis (Say). Atistrnlorbis glabratns (Say). Region of stomach from above. Region of stomach from above. Region of stomach from above.

Illustrations of the Anatomy and of Shells 333 PLATE 48 .

campanulatum smithi 12. Klamath Lake. pilsbryi injracarinatum F. Line at lower extreme left 0. in length. 19. C. Helisoma scalare (Jay). Superior jaw. Oregon. 5. Michigan. Utah. Helisoma subcrenatum (P. in length. 10. IL H. Colorado. North Carolina. 13. 8. Superior jaw. 2. H. indicates striation. Helisoma pilsbryi (F. Paul Lake. 49 of Planorbidae Helisoma anceps (Menke). of Helisoma occidentale (Cooper). Helisoma campatudatum smithi (F. 14. Helisoma magnificum (Pilsbry). Wisconsin. for 7 to 16. Helisoma traskii (Lea). Superior jaw. Helisoma oregojiense (Tryon). Superior jaw. Wisconsin. A. Gypsum.. 4. C. Ontario. J. Carinifex jacksonejisis J. Small fig. Wyoming. Wisconsin. 4. 15. 16. C.5 is 18 and 19 and represents fig. Grand Mesa. Vertical striation on jaw. Line at right of for figs. H. Lake Winnebago. Rideau River. Oregon. 1 indicates 1 mm. Michigan. Douglas Lake. Highly magnified. Colorado. Helisoma plexatum (Ingersoll). Superior jaw. Bamiji Lake. Oshkosh. Line at right is for figs. Forty miles west Miami. side. 20. Baker). Devils Lake. California. Helisoma horni (Tryon). 7. P. 2. Tooele Co. 3. Helisoma trivolvis (Say). Middle 5. fig. 17. (F. Baker. Florida. Jaw and cartilage from below. C. duryi normale Pilsbry. 3. Chetek Lake. 6. Baker). Baker). Chetek Lake. 6. 9. Henderson. Helisoma truncatum (Miles). Carinifex ponsonbyi E. Near Wilmington. Lake Butler. Carinifex jacksonensis Henderson. British Columbia. Outlet of Kern Lake. Florida. mm. Superior jaw. and line is . Jackson Lake.334 The Molluscan Family Planorbidae PLATE Jaws 1. Jaw and cartilage from the 18. Carpenter). Douglas Lake. Figure 17 is diagrammatic. Klamath Lake. Canada. Ontario. Helisoma corpulentum (Say). Smith. Wisconsin. Jaw from above.

Illustrations of the Anatomy and of Shells 335 .

Menetus cooperi caUioglyptus (Vanatta). Taylor Lake. Canada. Tropicorbis obstructus (Morelet). North Shan State. 10. Vancouver Promenetus exacuous (Say). Wisconsin. . Burma. C. C. Pkmorbula jenksii (H. Unionville. British Columbia. 13. Wisconsin. 16. 3. Intha capitis Annandale. 2. Lake Nipissing. 25. Louisiana. Indoplanorhis exustus (Deshayes). Puerto Rico. Ayiisus spirorbis (Linn. Missouri. 27. Baker. Armiger crista (Linn. Henderson. Near Warsaw. n. Near San Francisco. Poland. Poland F. 19. Gyraulus albus (Muller). Planorbis planorbis (Linn. Porapholyx effusa klamathensis F. Porapholyx effusa diagonalis J. Alberta. Baker. 6. 21. Cuba. 26. California. Gyrauhis vermicrdaris (Gould). Winnebago Lake. Gyraulus deflectus obliquus (DeKay). Wainwright Park. Connecticut.336 The Molluscan Family Planorbidae PLATE Jaws 1.). Segmentina nitida (Muller). Quebec.). Sizes are disproportionate. California. Island. Wainwright Park. (Pfeiffer). 5. Poland. 24. Unionville. Australorbis glabratus 14. Baker. Oregon. 23. Alberta. Wisconsin. 9. Carpenter). Canada. Figures are greatly enlarged. Tropicorbis riisei (Dunker). (Say). Superior jaw. Muri)hy Creek. Puerto Rico. Burma. 17. Planorbarius corneus (Linn. Gyraulus hirsutus (Gould). near Madison. Winnebago Lake. Shandon. Menetus dilatatus (Gould). 50 of Planorbidae Helisoma tenae calijornicnse F. Oregon. Poland. Ontario. Gyraulus parvus (Say). Near Warsaw. Near Warsaw. 28. Near Warsaw. Poland. Connecticut. 12. Drepanotrema hoffmani Drepanotrema lucidum Puerto Rico. 15. 18. Gyraidus circumstriatus (Tryon). Menetus sampsoni (Ancey). Klamath Lake. Wisconsin.). Near Stanton. Promenetus umbilicatellus (Cockerell). 4. F. Planorhiila armigera (Say). 7. New Orleans. 8. 20.). C. Near Warsaw. 22. Crater Lake.

Illustrations of the Anatomy and of Shells 337 PLATE 50 .

51 of Helisoma scalare (Jay) Egg capsule containing 28 eggs. embryo embryo embryo in egg No. Young snail two days out of egg No. 3. 7. 14. 8. 13. March 14. 7. 12. No. Young snail just out of egg No. Line at lower left indicates 1 mm. March 11. 11. 11. March 16. 20 on March 14. March 15. Embryo in egg No. 1. March March Post-trochophore stage in egg No. March 10. 9. Veliger stage of Veliger stage of Veliger stage of 6. in length . 7. 4. 1. in egg in egg No. Each egg is marked to correspond with the numbered figures below. 7. Young snail in egg No. March 12. The 24-cell stage of segmentation in egg No. 9. 1 showing heart and stomach. 11. March 10. 9. 3. 8. 8. 8. Line near figure indicates 1 mm. A similar young snail was observed in egg No. Shell growth distinct. 11. Shell now shows both growth Veliger stage of lines and spiral striae. March 4. Embryo in egg No. observed on March 1. Post-trochophore stage in egg No. in length. Top view. 3. Gastrula stage of segmentation in egg No. March Side view. embryo in egg No. March 9. March 8. 5. Front view. 1932. 5.338 The Molluscan Family Planorbidae PLATE Development 1. 2.

Illustrations of the Anatomy and of Shells 339 .

5. normale Pilsbry. Nine eggs laid Feb. Florida. eggs laid eluryi May 27. Spermatozoa from active group. 14-16. Four eggs laid Jan. Buena Vista Canal. 1932. Helisoma 6. Ten eggs laid April trivolvis 1931. 1931. Sixteen eggs laid Jan. Florida. 1. Helisoma duryi normale Pilsbry. 6. 1931. Lake Butler.340 The Molluscan Family Planorbidae PLATE 52 Nidifieation in Planorbidae 1-3. Gainesville. Louisiana. 25. 10. 14. 6-8. o\otestis. 1931. 11. Eupyrene spermatozoa from the Line near figs. Nucleated ovum from ovotestis. Wainwright Park. Two eggs laid Three eggs laid 8. lentum (Say). 1931. 14-16 indicates 25 microns in length. California. 17. 40 miles west of Twelve eggs 12. New Orleans. Five eggs laid Dec. 17. Florida. laid Feb. 4-5. 13. Miami. Fourteen eggs laid April 2. eggs laid March 13. 26. traskii (Lea). 9. Helisoma 13. Helisoma scalare (Jay). . Helisoma duryi cudiscus Pilsbry. 3. 5. 1931. Florida. 1-13 indicates 1 mm. Twenty Ten eggs laid June 1933. 9-11. Typical (eupyrene) spermatozoa as seen in 16. Tamiami 12. 10. 7. Gyraulus circumstriaius (Tryon). 15. L Five 2. 1932. Alberta. Silver Springs. 13. 1932. Trail. Helisoma 4. Dec. Alcoholic specimen wath 50 eggs laid on adult shell. 1932. outlet of Kern Lake. 1933. Dec. Line near figs. side and from in length. abo\'e.

Illustrations of the Anatomy and of Shells 341 PLATE 52 .

Helisoma anceps (Menke). marginal teeth. marginal teeth. center 9. 80. 6. counting from the front end of the radula. C. the first lateral tooth being nvuiiber 1. Bayfield. 9-13. pathologic center tooth. 6. The numbers following indicate the number of the tooth in this row. marginal teeth lateral on left side of 8. center tooth. 11-16. 1. 4. 8. Chautauqua Lake. Helisoma anceps (Menke). lateral teeth. side. center tooth. teeth. lateral teeth. HcILsoma anceps sayi F. 2. intermediate tooth. and 75 on this plate) indicates the number of the transverse row. marginal beside intermediate teeth. Helisoma anceps (Menke). center tooth in 50th row. 9. marginal teeth. The line at the lower part of the plate indicates 25 microns. intermediate teeth. 98. Wisconsin. 90. 8-14. first left it. 6-8. 2. Unionville. 7. 15. 90th row. intermediate teeth. 5. on right tooth. 7. 1-5. the number under the central tooth (as 15. tooth with two ectocones. 118. 50. center tooth (of 118th row) with 3. Baker. 8-14. counting from the central tooth. row. 7. 75. pathologic lateral . lateral teeth. marginal teeth. 6.342 The Molluscan Family Planorbidae PLATE 53 Radulae of Helisotna In the plates of radulae. center tooth. 1-5. 1-5. 1-5. marginal teeth 10. 80. New York. 1. Connecticut. intermediate teeth. 50. 98. lateral teeth. Maple River. Michigan. 75th row.

Illustrations oj the Anatomy and of Shells 343 PLATE 53 .

2. 60. 1-6. 1-3. center tooth. marginal tooth.. Quebec. nnceps percarinatum (Walker). 60. Canada. 10-23. 100th row. center tooth. intermediate teeth. 9. marginal teeth. pathologic (2 entocones) intermediate tooth. teeth. Line at bottom of plate indicates 25 microns in length. 344 The Molluscan Family Planorbidae PLATE Radulae of 54 Helisoma See remarks on plate 53. L Helisoma 100. 7-9. lateral teeth. marginal teeth. marginal 3. 1-6. Big Muskallonge Lake. intermediate teeth. 1-6. 51. 1-6. Ontario. Douglas Lake. 10-19. Bamiji Lake. lateral teeth. C. intermediate teeth. . 4. with accessory cusps above the normal center cusps. Meach Lake. intermediate teeth. 10-18. center tooth. 85. Helisoma anceps latchjordi (Pilsbry). marginal teeth. 10. lateral teeth. Helisoma anceps royalense (Walker). Wisconsin. center tooth. Helisoma anceps cahni F. Michigan. center tooth. 10-26. lateral teeth. Baker. lateral an abnormal center tooth from another membrane. 7-9. 7-9. 7-9. 60. lateral teeth. Canada.

Illustrations of the Anatomy and of Shells 345 .

102. lateral 8-10. From 11-13. 7-11. Tennessee. teeth. Chautauqua Lake. intermediate teeth. 70. C.y) ^ pseudotrivolvis teeth. lateral teeth. near lentum Say. Helisoma trivolvis {Sa. center tooth. Reelfoot Lake. intermediate teeth. lateral teeth. center tooth. 8-11. Louisiana.346 The Molluscan Family Planorbidae PLATE Radulae See remarks on plate 1. intermediate marginal 2. 8-10. F. marginal teeth. teeth. Line at bottom of plate indicates 25 microns in length. Helisoma trivolvis (Say). intermediate teeth. Wisconsin. Sturgeon Bay. 1-6. teeth. 1-7. 1-7. marginal teeth. New York. center tooth. marginal 7. Baker. lateral teeth. Helisoma 16-25. 3. trivolvis (Say). New York. 6. 107. trivolvis lentum (Say). Helisoma trivolvis lejitum (S&y) ^ pseudotrivolvis teeth. 1-7. C. 55 of Helisoma 53. New Orleans. center tooth. 13. 4. teeth. marginal teeth. Tennessee. 1-8. lateral 8-10. intermediate teeth. 9-11. with two entoconic cusps. Baker. . Oneida Lake. lateral marginal teeth. 11-18. center tooth. Reelfoot Lake. 14. marginal teeth. intermediate teeth. 1-7. Helisoma chautauquense F. Helisoma 96. 150. 12-15. New Orleans. 50. 12-15. 4. 11-14. Baker. trivolvis lentum (Say). F. 4th lateral tooth from another row. C. Helisoma 60. center tooth. 5. 50th center tooth of another row. Louisiana.

^ Illustrations of the Anatomy and of Shells 347 11 12 1 ^^150 I 2 3 4^-~~— 1234 PLATE 55 .

5. marginal teeth. 4. Little Helisoma winsloivi 70. 1-13. 1-8. C. 115. intermediate teeth. Baker. Helisoma pilsbryi injracarinatnm F. 2. Bayfield. 14. center tooth. Baker). 15. . 1-8. Varying toward diate teeth. 132. Helisoma pilsbryi infracarinatiim F. C. marginal teeth. 14.348 The Molluscan Family Plauorbidae PLATE 56 Radiilae of Helisoma See remarks on plate 1. 9-13. 10-14. 15-21. center tooth. 10-12. 53. intermediate teeth. Baker). lateral teeth. marginal teeth. C Baker. Wisconsin. 1-9. intermediate teeth. Rideau River. W'isconsin. Helisoma pilsbryi (F. Basswood River Rapids. Ontario. pilsbryi. lateral teeth. Ontario. (F. C. marginal teeth. lateral Chetek Lake. 100. intermediate teeth. Wisconsin. lateral teeth. 17-19. 15. center tooth. Line at bottom of plate indicates 25 microns in length. 1-13. marginal teeth. Helisoma trivolvis macrostovium (Whiteaves). 14-23. 3. 53. teeth. Arbor Vitae Lake. lateral teeth. interme- 16-29. center tooth. 13-23. center tooth.

Illustrations of the Anatomy and of Shells 349 PLATE 56 .

85. with many split cusps on teeth. teeth. 6 and 7 with bifid entoconic cusps and 5. In 16 of fig. marginal teeth. intermediate teeth. 90. the small numeral 2 indicates the position of the mesocone in a of the tooth: marginal tooth. 13-25.350 The Molluscan Family Planorbidae PLATE 57 Radulae of Helisoma See remarks on plate 53. ectocone or outer cusp. split intermediate teeth with modified ectocones. lateral teeth. Helisoma corpulent um (Say). Wisconsin. teeth. center lateral teeth. marginal teeth. Near Wilmington. 1-7. Winnebago Lake. 1 indicate the following parts entocone or inner cusp. center tooth. 2. 90. near Oshkosh. 2. 1-14. L Helisoma truncatum teeth. 24-41. Line at bottom of plate indicates 25 microns in length. Abnormal membrane. 1-8. intermediate teeth. marginal The small numerals on the 1. mesocone or middle cusp. center tooth. 1-8. tooth. 6. North Carolina. lateral teeth. 3. Wisconsin. Lac 48. 9-11. (Miles). 9-11. intermediate 13-20. 16-23. la Croix. 4. and 7 with ectoconic cusps. first lateral of fig. 1. 12-21. lateral teeth. base of attachment to radula membrane. Canada. center tooth. Winnebago Lake. Normal membrane. Ontario. Helisoma truncatum (Miles). . 9-12. Helisoma magnijicum (Pilsbry). marginal 4. 3.

Illustrations of the Anatomy and of Shells 351 16 17 1 22 25 25 26 27 28 32 41 PLATE 57 .

Lake. 17-19. 23. 6.352 The Molluscan Family Plauorbidae PLATE Radulae See remarks on plate 53. C. Baker. intermediate teeth. (Ingersoll). lateral teeth. 1-16. and 4 with 10-13. how and 5. Minnesota. 101. Baker. intermediate teeth. intermediate teeth. mesocones worn or pathologic. 20-35. (Ingersoll). Idaho. . a pathologic tooth in another row. Helisoma plexatum 44. a lateral tooth on the side with split cusps. lateral teeth. Line at 58 of Helisoma of plate indicates 25 microns in length. Canada. Helisoma plexatum 13. center tooth. center tooth. The two 13 figures show two rows. Helisoma plexatum The 120th row in another membrane from a Teton River specimen. marginal teeth. near Rexburg. modified marginal teeth. Colorado. C. Teton River. lateral teeth. a lateral tooth.soll). Ontario. C. 3. 9-12. intermediate teeth. 12. 14-15. center tooth. 13-21. 4. 13-16. being an intermediate tooth in one row 1 1-8. 2. 60. Ontario. marginal teeth. Kahniinminanikok 15-41. bottom L Helisoma whiteavesi F. (Inger. 11-14. marginal teeth. Helisoma corpulent um inidticostatum F. this tooth differed in a marginal tooth in another row. marginal left teeth. Helisoma corpuhntum vermilionense F. 1-7. marginal teeth. Canada. Smartweed Lake. center tooth. Baker. 105. 16-18. lateral teeth. Vermilion Lake. 1-7. Mille Lacs.

Illustrations of the Anatoiny and of Shells 353 PLATE 58 .

19-28. San Bernardino Mts. 13. P. marginal teeth. Grand Mesa. Colorado. 11-12. intermediate teeth. 5. 1-15. C. the lateral teeth with extra cusps on both entocones and ectocones and with mesocone much modified. lateral teeth. 14-21. Columbine Lake. tooth. San Bernardino Mts. Abnormal teeth. marginal teeth. center (Ingersoll).. 13-15. Carpenter). Washington. 60. lateral teeth.. Helisoma tenue calijorniense F. 18-22. Variety. C. Normal teeth. 60. lateral teeth. Helisoma plexatum 95. 1-10. 6. 2. Puget Sound. tooth. Baker. 12. 1. California. intermediate teeth. Line at bottom of plate indicates 25 microns in length. intermediate teeth. center tooth. 11. San Bernardino Mts. 3. 96. Helisoma tenue calijorniense F.. Colorado. C. 13-17. marginal teeth. Helisoma tenue calijorniense F. center (Ingersoll). center tooth. 12. intermediate teeth. 1-11. lateral teeth. 4. Helisoma plexatum 60. 1-10. Baker. California. intermediate teeth. Baker. center tooth. marginal teeth. 1-12. lateral teeth. An abnormal 11th lateral tooth. . marginal teeth.354 The Molluscan Family Planorbidae PLATE 59 Radulae of Helisoma See remarks on plate 53. Helisoma subcrenatum (P. California. Pass Lake. 16-18. 12-17.

Illustrations of the Anatomy and of Shells 355 PLATE 59 .

1. center tooth. Paget Sound. Paul Lake. 4. P. Helisoma subcrenatum (P. 1-9. Lost Lake. Helisoma traskii (Lea). lateral tooth with bicuspid entocone. Outlet of Kern Lake. 1-7. 80th row. 88. abnormal center tooth.. marginal fourth tooth in another row. center tooth. center tooth. intermediate teeth. Outlet of Kern Lake. 5. teeth. teeth. Skagit Co. 15-20. Carpenter). Carpenter). 10. 9. Washington. Variety. lateral teeth. Puget Sound. Washington. lateral teeth. intermediate tooth. 82nd row. marginal tooth. 1-9. center tooth. P. 90. Helisoma horni (Tryon). Carpenter). Washington. 17. 12-22. 13.356 The Molluscan Family Planorhidae PLATE 60 Radulae of Helisoma See remarks on plate 53. Helisoma traskii (Lea). marginal Helisoma subcrenatum (P. 20. marginal teeth. marginal teeth. Line at lower rig. subcrciialum (P. 12. 11. intermediate tooth. first California. British Columbia. 90. intermediate tooth. 7. 10. lateral teeth. 7. 15-32. lateral teeth. 4. Lost Lake. 11. 10-14. P. Variety. . Outlet of Kern of plate indicates 25 microns in length. 9. California. intermediate teeth. California. 5. 12. 3. 10. 2. marginal teeth. 60. 6. Helisoma traskii (Lea). intermediate teeth. L Helisoma 73.

Illustrations of the Anatomy and of Shells 357 PLATE 60 .

intermediate teeth. 13-20. center tooth. P. intermediate teeth. 45. 9. lateral teeth. 9-12.. intermediate teeth. 1-8. Helisoma subcrenatum (P. Klamath Lake. 1. center tooth. 9-12. lateral teeth. Line at lower right of plate indicates 25 microns in length. Cottonwood Pass. Helisoma oregonense (Tryon). Carpenter). Gypsum. Utah. 2. 58. 10. Colorado. 13-23. Helisoma occidentale (Cooper). Tooele Co. 11-21. lateral teeth. 1-8. .358 The Molluscan Family Planorbidae PLATE 61 Radulae of Helisoma See remarks on plate 53. 1-8. 3. marginal teeth. Oregon. marginal teeth. 95. marginal teeth. center tooth.

^ ^^ 18 61 .Illustrations of the Anatomy and of Shells 359 C^X 3 PLATE .

9. 9. intermediate teeth. 10. 50. 11-20. 1-8. 11-19. Arbor Vitae Lake. 11-15. 60. Helisoma campanulatum smithi (F. 60. New York.360 The Molluscan Family Planorbidae PLATE 62 Radiilae of Helisoma See remarks on plate 53. center tooth. 1-8. Wis- intermediate teeth. 8. Line at lower right of plate indicates 25 microns in length. center tooth. 10. Helisoma campanulatum canadcnse F. Baker). 4. Oneida Lake. . C. marginal teeth. 9. 52. lateral teeth. consin. Bamiji Lake. marginal teeth. 10. wisconsincnse (Winslow). 1-7. 3. Baker. intermediate teeth. lateral teeth. 9. Ontario. marginal teeth. 1-8. Helisoma campanulatum. intermediate teeth. lateral teeth. center tooth. C. 10-20. Little 2. L Helisoma campanulatum (Say). marginal teeth. Michigan. center tooth. Douglas Lake. Canada. lateral teeth.

Illustrations of the Anatomy and of Shells 361 .

Numerals 1-11. intermediate tooth. Cambridge. 77. center tooth. lateral teeth.362 The Molluscan Family Planorbidae PLATE 63 Radulae of Helisoma See remarks on plate 53. center tooth. Massachusetts. 13-19. 12. C. 4. 1-11. Florida. 1-10. 60. center tooth. 11-23. marginal teeth. Helisoma tenue californiense F. . 60. California. 11. intermediate teeth. marginal teeth. lateral 2. 12-22. 84. Helisoma trivolvis lentum (Say). intermediate teeth. 12-20. 10. marginal teeth. lateral teeth. marginal teeth. 12. Shandon. lateral teeth with multiple cusps on entocone and ectocone. Helisoma trivolvis fallax (Haldeman). Baker. 11. Line at lower right of plate indicates 25 microns in length. 10. 1. center tooth. Helisoma duryi eudiscus Pilsbry. 3. teeth. Louisiana. lateral New Orleans. 1-7. teeth. intermediate tooth. 1-9. Silver Springs. 100th row.

Illustrations of the Anatomy and of Shells 363 r"^ 14 15 17 4 63 PLATE .

Lake Butler. center tooth. teeth. 10. Florida. marginal teeth. 12. Trail. Line at lower right of plate indicates 25 microns in length. marginal teeth. lateral teeth. 12-20. intermediate tooth. center tooth. Florida. 11. 1-6. 9-11. 1-10. Paines 55. 12-21. intermediate tooth. 1-11. 89. center tooth. 12th tooth (marginal) in another row of teeth. 13-29. Helisoma duryi normale Pilsbry. Florida.364 The MoUuscan Family Planorbidae PLATE 64 Radulae of Helisoma See remarks on plate 53. 2. Helisoma duryi normale Pilsbry. lateral teeth. marginal lower right. intermediate tooth. center tooth of another specimen. marginal teeth. center tooth . Lake Butler. intermediate teeth. . 4. Florida. 1-9. L Helisoma scalare (Jay). lateral teeth. 1-8. 140. 12. Prairie. Tamiami 110. 86. 11-26. 40 miles west of Miami. lateral teeth. lateral teeth. Helisoma scalare (Jay). 3.

Illustrations of the Anatomy and of Shells 365 PLATE 64 .

105. intermediate teeth. 9. 1-6. 11. marginal teeth. teeth. intermediate teeth. Parapholyx. Planorbula jenksii (H. center tooth. 4. intermediate teeth. marginal . 1. 9-11. center tooth. Wisconsin. 11. Wyoming. lateral 120. lateral teeth. near Madison. C. center tooth. 128. 12-19. 1-5. 1-10. Unionville. Planorbula armigera (Say). Baker. Connecticut. 13-24. and Carinifex See remarks on plate 53. 9. Line at lower left of plate indicates 25 microns in length. 12. Klamath Lake. marginal teeth. 13-19. intermediate tooth. Jackson Lake. Carpenter). Parapholyx effusa klamathensis F. Cariyiifex jackso7ic7isi. Murphy Creek. teeth. F. 3. marginal teeth. Oregon. lateral teeth. Henderson. 2. 12-19. center tooth. 1-8 lateral teeth. 60.366 The Molluscan Family Planorbidae PLATE Radulae 65 of Planorbula.<i J.

Illustrations oj the Anatomy and of Shells 367 128 1 2 3 4 <» 9 /'^:=:) .

intermediate teeth. 23-26. 27. 32. in same row. outer marginal teeth Planorbarius corneus (Linn. 11. marginal teeth. marginal teeth in same row. 15-38. Line at lower right of plate indicates 25 microns in length. 39-42. 1-5. North Shan State. 1.). 12-15. 2. Burma. 16-21. last intermediate tooth in another row. 33. 1-9. Near Warsaw. teeth. 10-14. 60. center tooth. marginal teeth. lateral intermediate teeth. Indoplanorbis exustus (Deshayes). Poland.368 The Molluscan Family Planorbidae PLATE 66 Radiilae of Indoplanorbis and Planorbarius See remarks on plate 53. . outer marginal teeth. marginal teeth in same row. lateral teeth. 60. center tooth. 6-10.

Illustrations of the Anatomy and of Shells 369 »* 17 18 23 25 26 27 32 33 PLATE 66 .

1.370 The MoUuscan Family Planorbidae PLATE 67 Radiilae of Planorbidae See remarks on plate 53. 10. 1-6. 50. 1-7. 11-15. marginal teeth. 11. British 3. marginal teeth. teeth. center tooth. Stanton. 50. Vancouver 55. marginal 6. intermediate tooth. 1 to 5. 11. Columbia. 8. 115. intermediate teeth. 14-20. 4. Anisus spirorbis (Linn. teeth. A. 1-7. 60. 1-8. Smith.). marginal teeth. center tooth. lateral teeth. intermediate tooth. Wainwright Park. 1-9. intermediate teeth. 11-14. 12-15. 8-10. 60. 45. intermediate teeth. lateral teeth. marginal teeth. 11-27. 60. Connecticut. 5. 12-15. 10. center tooth. Wainvvright Park. Menetus dilatatus (Gould). center tooth. 9. teeth. Near Warsaw. intermediate teeth. lateral teeth. 8. 1-6. 11-14. 10.). intermediate teeth. marginal center tooth. Oregon. Unionville. Menetus cooperi caUioglyptus (Vanatta). marginal . Poland. center tooth. Missouri. Line under fig. 7-10. lateral teeth. Alberta. Alberta. Island. center tooth. 9-11. lateral teeth. lateral teeth. Poland. Promenetus umbilicatcUus (Cockerell). 11-15. 1-6. 9. Klamath Lake. Menetus sampsoni (Ancey). Planorbis planorbis (Linn. 6 is also for figs. 1-10. lateral teeth. Camiifex ponsonbyi E. marginal 2. intermediate teeth. 7. teeth. lateral teeth. Line at lower right of plate indicates 25 microns in length. Near Warsaw. Promcnetus exacuous (Say). center tooth.

Illustrations of the Anatomy and of Shells 371 10 11 12 '< ^ 10 11 11 12 ii " 11 12 13 14 ^^ts^Ts 12 li 14 45 1 PLATE 67 .

60. intermediate teeth. 5. Drepanotrctiia hoffmani F. lateral teeth. Australorbis glabratus (Say). 9. Louisiana. 13. 5-9. center tooth. 6. Puerto Rico. marginal teeth. Puerto Rico. marginal teeth. 1-7. 15. marginal teeth. Baker. 7-12. lateral teeth. lateral New New Orleans. 7. 60. intermediate teeth. 8. 60. 1-13. 16-26. intermediate teeth. center tooth. center tooth. lateral teeth. 60. Near Warsaw. 5. C. Gyraulus albus (Midler). Tropicorbis obstructus (Morelet). 1. Poland. 52. marginal teeth. marginal tooth. 6. 11-14. 1. 50. 8. lines for figs. Drepanotrema anatinum (Orbigny). intermediate teeth. Cuba. 7. intermediate tooth. 15. 1-6. intermediate tooth. center tooth. 13. Lares. 1-4 are 10 microns in length. 13. 12. intermediate teeth. Poland. Line at lower right of plate indicates 25 microns in length for figs. intermediate teeth. marginal tooth. 8. 2. Louisiana. lateral tooth. 14. 1-5. center Orleans. 9. lateral tooth. Tropicorbis havane^isis (Pfeiffer). 1. 6-8. Bctthyomphalus contortus (Linn. 14. lateral teeth. . 5. Havana. 3. tooth. Puerto Rico. 10. marginal teeth.). marginal teeth. lateral teeth. center tooth. center tooth. Puerto Rico. 55. center tooth. Near Warsaw. 1-8. 1. teeth. 50. center tooth. Drepcmotrema lucidum 50. Tropicorbis riisci (Dunker). 10.372 The Molluscan Family Planorbidae PLATE 68 Radiilae of Planorbidae See remarks on plate 53. 8. (Pfeiffer). 9. 4. 11-21. 12..

lUustratiojhs of the Anatomy and of Shells 373 6 8 II 12 13 '* gA<^^ 52 1 60 1 ^^K ^ < V M 14 ^ 15 20 22 PLATE 68 .

5. Near San Francisco. Tolland. lateral teeth. 50. 1-8. Canada. Segmentina nitida 60. Taylor Lake. 1. lateral tooth. Canada. Gyraulus hirsutus (Govild). 8. (Miiller). 6 to 9 indicate 10 microns in length. Lines near figs. marginal tooth. center tooth. center tooth. 1-8. 4. 2. 55. near Oshkosh. marginal teeth. 13. 50. 13-16. 10. lateral 10. 6. Burma. 9. 9. center tooth. marginal teeth. 7-10. 1-8. center tooth.saw. intermediate teeth. Gyraulus parvus (Say). intermediate teeth. 1-6. marginal teeth. center tooth. Ontario. marginal teeth. North Shan State. 60. center tooth. 7-10. Winnebago Lake. intermediate teeth. intermediate tooth. Quebec. 13-18. Wisconsin. Michigan. center tooth. Intha capitis Annandale. 1. lateral teeth. Near War. 1. 1. lateral teeth. 1 to 5 indicate 25 microns in length. 7. 13. 12-16. marginal teeth. lateral tooth. center tooth.374 The Molluscan Family Plariorbidae PLATE 69 Radulae of Planorbidae See remarks on plate 53. marginal teeth. Gyraulus vermicularis (Gould). (Miiller). marginal teeth. Gyraulus deflectus obliquus (DeKay). 1-6. 5. 60. C. North Shan State. 6. 11. 12. California. teeth. lateral teeth. 3. 12. 60. marginal teeth. Burma. 1. intermediate teeth. Armiger imbricatus 55. lines near figs. 13-18. Lake Nipissing. 6. Polypylis calathus (Benson). . 8. Poland. Colorado. intermediate teeth. 15. 11. 8. center tooth. 50. lateral teeth. 9-11. Gyraulus similaris (F. Baker). 11. lateral teeth.

Ulustrations of the Anatomy and of Shells 375 12 15 50 n *- 2 60 1 n 13 55 1 60 1 10 11 5 50 1 Ts 10 12 6 55 1 PLATE 69 .

hirsutiis New (Gould). Animal crawling up side of aquarium jar. Gyraulus kee. Wisconsin. in length . Planorbula (H. jeriksii Top view jar. view of crawling animal with parts indicated. of animal. 7.sac. Adult with mm. New York. 3. Canadian specimen. near Alton Bay. 13.y). F. Louisiana. Head and tentacles of crawling animal. Connecticut. Florida. Lake WinnipesauHampshire. . Menetus Menetus dilatatus (Gould). 13. aquarium Helisoma subcrcuatum (P. 9. Young Animal. scalare (Jay). shell. Drawn from aquarium specimen. Carpenter). Hclisoma campanidatum (Say). side 2. Near Bayfield. Side view. Halfmoon Pond. P. Buccal radular sac and the buccal sac. Animal crawling on stick in aquarium 10. Carpenter). Helisoma trivolvis (Sa. Massachusetts. New Orleans. View as seen crawling up of aquarium jar. dilatatus (Gould). New Hampshire. Helisoma scalare (Jay). Connecticut. Hclisoma trivolvis jallax (Haldeman).376 The Molluscan Family Planorbidae PLATE 70 Animals of Planorbidae 1. Cambridge. shell 10 Lake Butler. Shows pigmentation of mantle through 5. Hclisoma trivolvis macrostomum (Whiteaves). Unionville. The outline of the radula is shown in the Lme near figure indicates 1 mm. 11. Buccal sac with parts indicated. long. March Helisoma from aquarium specimen. From of pool near Lake Wentworth. Gyraulus albus (Miiller). Oneida Lake. Unionville. Tropicorbis havanensis (Pfeiffer). Twelve hours out of egg. Side 8. 6. Animal crawling on bottom 12. jar. Drawn 4.

Illustrations of the Anatomy and of Shells 377 PLATE 70 .

Fig. Fig. Reproductive organs (genitalia). upper sac (vergic . seen from above. X etc. Isidora globosa (Morelet). Genitalia showing spermatheca. 11. adowensis Bourg. stomach. plate 7. Copied from Pilsbry. Planorbis (== Afroplanorbis Sci. South its Africa. adoweyisis p. as noted above. plate 8. X 8. 5. oesophagus. Alimentary canal and salivary glands seen from above. G. the verge indicated by dotted lines. lQ-\2. 10. Copied from Connolly. and first loop of X 6. prostate. Posterior end of penis (vergic sac) showing penis papilla. 14. Fig. 7-9. etc. Copied from Connolly. Animal without its shell. H. 11. Preputium cut open to show pilasters and end of verge. 3. u. intestine. X 5. 5. Fig. (Bourg. Fig. Proc. caecum. 17. Fig. X 8. Posterior end of penis (verge) with sheath split open. anus. Reproductive organs (genitalia). LXXXVI. 9. Royal Soc.sac). with the folded branchial tube. 12. 3. 13. 15. Pulmonary Fig.378 The Molluscan Family Planorbidae PLATE 71 Isidora and Afroplanorbis 1-6.. pjeifjeri (Krauss)).. Planorbis pfeiffcri Krauss {^= Ajroplonorbif< 8. orifice. s. X 6. orifice. Phil. 1. Fig. seen from the left side. this specimen exhibiting a different condition from that seen in fig. XII.)). Includes 8. X 12. Lobes beneath the pulmonary pseudobranch and rectum. 2. uterus. Nat. showing the junction of the vas deferens with the penis (verge). Fig. F. 18. Fig. 53. 19. 12. Acad. Trans. 1. X 11. Fig. salivary glands. 4. X 6. Figures are great Iv enlarged . on the right of it. Fig. Buccal mass. 8. Male organ (penial complex) showing preputium and verge.

Illustrations of the Anatomy and of Shells 379 .

ps. from above. Acrorbis petricola 4. Portion of genitalia. rp. and from above. one-half row on X 1000. flagellum (anhangsdrlise) spermatheca (bursa copulatrix) c. from above. t. Animal removed from shell to show organization. k. oesophagus. tentacle. 9. Digestive system. . o. for Zoologi. here including the whole penial complex. r. v. j. od. a blind sac. 8.. kidney (niere) the ovisperm duct) p. a. ganglia in nerve collar. prostate. . front view. jaw. pseudobranch (kiemen falte) I. 5. preputium and vergic sac) . caecum. salivary glands (speichel driisen) radula sac (radula tasche) u. foot of animal. stomach. go. penial retractors. . Head and Buccal sac. 380 The Molluscan Family Planorbidae PLATE 72 Acrorbis petricola Odhner Copied from Odhner's paper in Arkiv. heart in pericardium. rectum. vas deferens. Xo. . s. liver. 14. 7. Penial complex. 10. uterus. preputium (penissack. f. ovotestis (gonad). fore view. h. left side. oviduct (in the figure this refers to n. d. Shell from below. buccal sac. 1-3. intestine. m. Band 29B. ph. . 6. Description of symbols used on this plate: b. g. Radula. . Odhner. two plates redviced to one plate in this volume.

Illustrations of the Anatomy and of Shells 381 1 mm 2 '®(^ O V 10 PLATE 72 .

Bf. Pr. . spermatheca (poche copulatrice) penial). verge (tube musculaires) Pc. vergic sac (poche du penis). Transverse section of vas deferens (canal deferent). ovotestis (glande hermaphrodite). horizontal line Vertical line indicates measurements in millimeters. Explanation of symbols used on the plates: buccal sac.. Al. C '^ . . vagina. Cd. Sp. p. . General view of genitalia (in French works the figures of both the animal and the shell are in reverse as compared with the position of American figures). retractor compare the insertions of this figure with muscle (muscle fetracteur) the figures on plate 22 of this work. . Od. Pp. albumen gland. Ut. C. 288 1. oviduct. P. . uterus. ovi- sperm duct (canal hermaphrodite) G. nature of verge. preputium (gaine) . in microns. pilasters (piliers nidamental gland (glande uterines) Pe. junction of preputium with vergic sac (col). 382 The MoUuscan Family Planorbidae PLATE * 73 Indoplanorbis exustus (Deshayes) Genitalia cojiied from Larambergiie. Penial complex (appareil copulateur) with vergic sac cut open to show length and 2. prostate. Va. 3. vas deferens (canal deferent) G ^. Gu. sperm duct (spermiducte) . 1939b. R. .

Illustrations of the Anatomy and of Shells 383 PLATE 73 .

the open to show portion of verge not everted. Specimens anesthetized. verge (tube penial). 1939b. animal acting as male.384 The MoUuscan Family Planorhidae PLATE 74 Indoplanorbis exiistus (Deshayes) Genitalia copied from Larambergue. 9 Vertical line indicates size in millimeters . vas deferens. Indi\'idual with verge latter cut and preputium protruded from male genital opening. 293 6. prejiutium (gaine) Pe. Pp. Two individuals of Indoplanorbis in coitus showing the extension of the complex from the body 8. Schematic figure of penial complex retracted within the body of the animal. . Schematic figure of penial complex everted during copulation. vergic sac (poche du i)enis). p. penial of the 7. Explanation of symbols on the plate: Cd. G.

Illustrations of the Anatomy and of Shells 385 6 7 PLATE 74 .

5. verge (penis). testis C Al. Cd. oviduct. Pp. atrium. vergic sac (poche du penis). 93 4. 386 The Molluscan Family Planorhidae PLATE 75 Bulinus contortus (Michaud) Genitalia copied from Larambergue. uterus. C. (glande hermaphrodite). Bb. sperm duct. vagina. G. . prostate. 6. junction of preputium with vergic sac (col). Carrefour of the genitalia. retractor muscle. 1939a. Penial complex cut open to show characteristics. . Ut. p. General dissection of genitalia and fore part of body. Va. albumen gland. buccal sac. vas deferens (canal deferent). R. C '^ ovisperm duct (canal hermaphrodite). duct of albumen gland. pilasters (piliers musculaires) Pc. preputium (gaine) G ^. P. Vertical line indicates size in millimeters . ovo- Od. Pr.. Explanation of symbols on the plate: A. . spermatheca (poche copulatrice) Pe. Al. Sp.

*«r.Illustrations of the Anatomy and of Shells 387 .-' PLATE 75 .

Resultate d. Wissens. V.. flagellum. 3.. op. 6. Apertural lamellae. Greatly enlarged. 12. 6. Vorticifex tryoni (Meek). Forbes and Hanley. 6. X6. 9. fig. Drepanotrema cultratum (Orbigny). 5. Greatly enlarged. fig. cit. Planorbula armigcra (Say). 7. About X5. cit. 17. B. vide Pilsbry. Planorbis krambergeri Halavats.. Lamellae on outer lip. 3b. Op.. Planorbida icheatleyi (Lea). XVII. Lamellae on outer lip.).S. Lorenthay. I. plate Vorticijex binneyi (Meek). figs. 8. Armiger crista (Linn. plate 126. 13. Polypylis hemisphacrida (Benson). Lamellae on outer Greatly enlarged. 5-7. Mus. Odontogyrorbis krambcrgeri (Halavats). Amer. Erforsch. figs. vd. Art. 3. figs.. Pompholopsis whitei Call. p. 2. lij). Greatly enlarged. 8. plate 2. figs. Geol. pr. Baker). 5. Segmentina nitida (Miiller). plate 31.. plate 3. 3a. Colombia. Conch. Hist. 5-7. figs. Nat. Costorbis strauchianits (Clessin). Figures 11 to 13 are natural size . U. About X4. IV. Cab. fig. vas deferens. E (after H. White. Penial complex. fl. X3. Enlarged. Moll. From Bolivar. 1934. 7. 11. 5.388 The Molluscan Family Planorbidae PLATE 76 Shells of Planorbidae 1. Brit. Proc. 9. Pilsbry. 4. 3c. Tropicorbis obstnictus (Morelet). des Balatonsee. Apertural lamellae. 10. 147. penial retractor.

Illustratiois of the Auatotny and of Shells 389 1 4!^:.^ -"^mmS^ o 11 12 10 .

390 The Molluscan Family Pla7iorbidae PLATE 77 Shells of Planorbidae 1-3. Wheatland Center Road. Drewnica..). Anisus leucostoinus Titicaca. X2. Xl^/. Baker coll. Poland.. northeast of Warsaw. 13-15.. Baker coll. X2. Baker coll. (Linn. Poland.Z.). 40466. southeast of Warsaw. 3972. Taphius andecolus (Orb. Australorbis glabrotus (Say). 40595. Pond at Scarborough. 7-9. England.. Baker 19. of Warsaw. Poland. X4.). Pond in Poland. 4-6. .I..I. 29-31. M. Anisus vortex (Linn. Anisus spirorbis (Linn. 10-12. U. jiark in Krolikarnia. Stream in meadow in Jab-Tonna. X2. 3969. (Morelet). A pond 25 km. a suburb of Warsaw.. .B. northea. Mexico. X4. 3971. 65218. Small pond in Bielany Park. 26-28. Tropicorbis pallidus (C. Monroe Co. Gyraulus albus (Miiller)... U. 20-22. 18 km. Baker coll.).C. Blue Pond. 23-25.L. Lake (Millet). Jamaica. XP/o.I. 4 km.. Kingston. X4. 1234. Planorbis planorbis (Linn. Puerto Rico. Peru. X4. Baker coll. X4. Gyraulus parvus (Say). Tropicorbis orbiculus Valles.Adams). Bathyomphalus contortus Poland.. 3968.). NewYork. 3970. X4. U. San Juan. U. 41369.. 28525. 16-18. Drift at falls. Warsaw.

Illustrations of the Anatomy and of Shells 391 10 ^^&B 11 4| 16 12 23 13 17 26 14 24 27 15 25 20 19 f 28 29 21 "^ 22 31 PLATE 77 .

S. Inst. 4. Lutz. X3.. Baker coll.. 121167. 10. fig. Hongkong. Animal from above. Carinif ex neu'berryi {Leu).S. 121025. fig. F. California. X3. California.. Painters Pond. Canton. X3. (Lea). Isabela.. California. Laguna del Bili. 20.N. LT. Upper and under side of animal. Figures 5. (Pfr. 4. Figures greatly enlarged. X3. Painters.. U.S.N. 6. Canoe Creek. 14-16. 3. Antigua. Carinifex 9256. 5. X3. newberryi (Lea).S. California. Pingiella peipinensis (Ping and Yen). Parapholyx 121133. Lea coll. X3.). China.. U.). Drepanotrema cultration (Orb. Baker coll. U. Parapholyx effusa (Lea). 24.2. 2751. X3. China. 3. Types. M. Animal from above.C. Canoe Creek.. . 6. 56405. plate 16.X.. 11-13. X3.. Figures Baker 3975. fig. c. mislabeled Planorbis nigricans by Lutz.Z.. c. 7. M. 25-26.. Peiping.N.M. Cuba. lot. coll. 272282.M.N. 8-10.N . Tropicorbis nigrilabris (Lutz).392 The Molluscan Family Pla)iorbidae PLATE 78 Shells of Planorbidae 1. X3. 5-7.S. Alacranes Mts. coll. Lea coll. Oswaldo Cruz. 2Z.. Tropicorbis nigrilabris (Lutz). mcJlcus (Lutz). fig. C. U. Carinifex newberryi minor Cooper. Baker coll. Mem. Helicorbis U7nbilicalis (Benson). Puerto Rico. a. Lea U. lighter in pigmentation than From Lutz. Baker. Drepanotrema hoffmani 3976.. Lea coll.M.. China. effasa 22. California. Sacramento River.M. Tropicorbis fig. From Lutz. c.. Probably type Sacramento River. 17-19.. Clear Lake. X3. Type.M. 1-4 from 2756.S. vol. Drepanotrema lucidum 129991.21.

Illustrations of the Anatomy and of Shells 393 ui>\^ 11 12 13 14 15 16 .

13-15. Peiping. Baker coll.. 16. Baker coll. Wrights Dike. X5. 16-18. 17. Mill Pond. Poland. U. Planorbida ivheatlcyi (Lea).. 1140. 227303.. 31. U. on 6. plate plate 8.). Xl%. Mcnetus opercidaris (Gould). X4. coll.. Thief River Falls. PerriniUa corddlcriann (Hannibal). Minnesota.. Hill near HawBelmont Stage Road. 34.M. Baker 2229. Hip pe litis fontanus Painswich.. X4. Lake Beds. Brazil. 19-21. Nevada. Truckee thorne. Para. Strand.. 10-12.) England.Z.. X4. Baker 3978. Polypylis largiUiarti ^ hemisphaerula (Pils.). Peru. X4. Swamp on a meadow Warsaw. 72832. 348535. Platytaphius heteropleurus & Van.N. Calcutta. 1231. X4. Drepanolrcma anatinum fig. 1140. New York. Baker coll.. San Francisco. Alabama. 35. Duxbury. Damsells X6. Indoplanorbis exustus Figures 33. Hannibal. 3974. fig. X2.394 The Molluscan Family Planorbidae PLATE 79 Shells of Planorbidae 1-3. (Phil. Soc. California. Bombay.N. 18. Small brook west bank Caj^uga Lake. China. Lidia. Massachusetts. Planorbida armigera (Say).. (Gould). 34. 33-35. Baker coll. 27093.). 22-24. (Orb. dilatatus Mcnetus coll. Baker 5973. near Cayuga. Proc.C. 7-9. Lake Titicaca..). (Benson). Promenetus exacuous (Say). {Ijightioot)=^ cornplanat us (Linn. 3972.S. 25-27. coll. Figures 16. Baker coll. 28-30. Mai.S.. (Desh. fig. in Czerniakow. West coll. Segmentina nitida (Miiller). Para. U. a suburb of 4-Q. Boligee..32. Brazil M. X.M. India. X4. . About natural size.I. Nevada. Baker 1256. fig. X4.

Illustrations of the Anatoiny and of Shells 395 10 11 12 13 16 19 22 €1 14 17 15 20 23 ^BP^ 18 21 Bp^ 24 33 25 28 31 26 29 34 32 27 30 35 PLATE 79 .

River. Braddocks Bay. 1962. Helisoma duryi seminole Pilsbry. trivolvis 13-15. Physoid shell. 237L X2. 4-6. Like Say's X2. Dade Co. Lake Eustis. South Street Brook. X2. = Planorbis II/2 bicarinatus Sowb. York. Florida. X2. 9. 41489. head Baker coll. 27112..L. St. Strongly 7-8.). Lake U. Typically planorboid U. miles above Plum Point. Everglades.L. U. Miami River. 16-18. Lake Pinsett. shell. Helisoma Ontario. Marciuette Co. Helisoma multivolvis (Case). From Delaware River figures.396 The MoUuscan Family Planorbidae PLATE 80 Shells of Planorbidae 1-3. France. New 19-21. U. Michigan. 25740. New Jersey. . near Rochester. U.. carinated form. Howe Rhone Lake. Helisoma scalare (Jay).. Baker 11-12. Florida. Planorbarius corneus (Linn. Helisotna anceps (Menke) U...L. Z25738. Auburn. coll.L.L. Z25934. X2. XlV->. of New York. Baker 22-24. bicarinatus Say. (Say). Xli/2. Helisoma campanulatum (Say). Thousand Island Park. Helisoma anceps (Menke). 29789. Xli/1'.L. New York. coll. 3977. XIYj. and P. Florida. Lawrence River. Helisoma duryi nurmale Pilsbry.

IUustratio)is of the AiiatoDiy and of Shells 397 .

Pilsbry and Bequaert. 4. Line near figures 10.N. 12. Choanomphalus plate 4. 5. Choanomphalus schrenckii Dybowski. p. 9. Vicinity of Warner Lake. 215381. Xli/4. 118. flattened spire to long. . cit.. Pilsbry and Bequaert.. Op. fig. Proc.S. 1. figs. 11. Belgian Congo.. inaacki Ger. cit. Bull. Afroplanorbis (Martens). 86. Nat. p. Anisopsis loryi (Coquand). op.. p. 1-3. figure.). Poecilospira trochiformis (Stahl). Pliocene period. Showing extreme range M. cit.. Oregon Pub. . plate 4. 28. lower figures. 8-10 indicates natural size. 2. Africa. fig. paratypes from Lake Edward. Jour. a. 28b. figs. de Conch. taf. Op..H. conical spire. 122. N. 7a. plate 21. dome Kettleman Line indicates Hills. Amer. 1. taf. Germany. sudanicus fig. 7. XXVH.stfeldt. Biomphalaria smithi Preston. fig.-Conch. c. 7. North fornia.. Paraplanorbis condoni (Hanna). LIII. fig. Calib. X2. Lake Edward. plate 4. specimen from Kabare. Univ. mm. op. Africa. Xli^.Z. 28a. plate 1. cit. Anisopsis calculus (Sandberger) (stark vergrossert). Pliocene.C. About X7. Africa. 8. Sci.. Vorwelt.. Acad. 10. Xfi/^. No. Hanna. PiLsbry. fig. Upper figures. eastern Oregon.398 The Molluscan Family Planorbidae PLATE 81 Shells of Planorbidae 1. of variation Steinheiin. I. Mus. Planorhijex vanvlecki (Arnold). 6.. 9. cit. Pilsbry and Bequaert. figs. Ajroplnnorbis adoivensis (Bourg. 6. M. 1 fig. Op. Crosse and Fischer. Reduced one-half from Pilsbry "s 7. Bahr-el-Ghazal. Land und Siiss. Choanomphnlus valvatoidcs Dybowski.. 4. U. Phil. 8. 9. King Co. 3. 121. from short.

Illustrations of the Anatomy and of Shells 399 m^ 10 28* ^ 26^ 11 PLATE 81 .

32-34. type locality. B999. Idaho. 15. Oregon. Milwaukee.400 The Molluscan Family Planorhidae PLATE 82 Shells of Planorbidae 1. U.A. Z19354. Priests Lake. (F. 3-5... Portland. . 81798. East Lake Okoboji. Frontenac.S. Helisoma anceps (Menke). 27-29..M. 18. Portland. Figures are enlarged approximately two diameters coll. Baker.2. Walker sembling Conrad's Planorbis antrosa. 31. B2851. 30. Maryland. U.M. 6-8.M. 24-25. From marl bed.S. Oregon. Wisconsin. Helisoma anceps (Menke). Baker. Idaho.. U. C. paratype.M. 14. Helisoma anceps (Menke).. Helisoma anceps niinnesotoise B1002. Shimek U.M. Dove Co. 26. Michigan. Hemphill Helisoma anceps striatum (F.N. Helisoma anceps corrugatum (Currier). re- resembling sayi. Wiscon.N. Re- Helisoma anceps (Menke)...19. Coeur d'Alene Mts. U. Washburn 23. B1819. 6875. Minnesota. North Carolina.. 20-22. U. Ashhmd. 9-n. Virginia. Helisoma anceps (Menke).I. Bonnar Co. Alpena. Baker).C. 476580.M. Resembling Rackett's BIOOL coll. Northern part of Minnesota. Alabama. Long Lake. Co. 23143. U. Baker).. 16. 81780. Potomac River.. Bay View. 364718. Resembling Menke's at figure. 23-25. sembling unicarinatum .I. Types.. C. Fort Washington.. Shell Lake.^in. C.. C. Spring Grove. Helisoma anceps shellense F. Iowa. Helisoma anceps sayi F. Helisoma ancejhs (Menke).S. Blue Creek. Helisoma anceps (Menke). 6740L 12-15. Helisoma anceps (Menke). Z13474. C. 17. Helisoma anceps (Menke). B985. coll. Below dam figm'e of Helix angulata. Lake Junalaska. U..

lUustratious of the Auatomy and of Shells 401 mm J^ ^m ^W ^^^ ^F •t 10 11 ^ 12 13 *? "^W^ 17 14 15 16 C) 18 19 20 /" 21 22 (• 23 '«* J 25 ^ 24 26 •.«>. (' 27 28 29 30 31 32 33 34 PLATE 82 .

Baker.. Aroostook Co. Variety? Mouth LT.S. Helisoma anceps unicarinatum (Hald. U.N.N. Helisoma anceps aroostookcnse (Pilsbiy).402 The Molluscan Family Plauorbidae PLATE 83 Shells of Planorbidae 1-3. TJ.. Variety? Potomac River. Baker. U. U. Helisoma anceps sayi F.M.S. Helisoma anceps politum F. 109888. 23-25. Type locality. paratypes.).L. Iowa.. River Station. Little 10-12. Alabama. 26. Riverton. 18. C. LT. U.. Helisoma anceps sayi F.N.. Canada. 367426. paratypes. U. Shimek ing toward anceps. West Virginia... Delaware River. above New Jersey. Figures are enlarged approximately two diameters . 16-18. Z25938. Type 4-6. 475945.. Blackstonc River. West Lake Okoboji.M. Helisoma anceps (Menke). Falls. Brook at Great fig.M. of Holotype. Honeywell Ontario. Holotype. Yaqui River.S.I.S. 364718. 27-29. C. Virginia. Helisoma anceps (Menke). Vary- 7-9. C. Helisoma anceps unicarinatum (Haldeman).. U. Z21015. C.S. Cherokee River. Baker. coll. 53677.S.. Plum Point. Woodland.M. Z25736. Tomahawk Lake. Carleton Co. locality.N. Helisoma anceps unicarinatum (Hald. Baker.I. Creek. R. Z41055. figs. Latchford coll.. 17. 227858. U. Oneida Co. 16. 19. Helisoma anceps bartschi F. Northwest Mexico. above Courtland.M..). 20-22.I. Wisconsin. 19-22. Maine. figs. Figure 13 resembles Haldeman's angisiomum.N.M. 13-15. fig.N . F.

Illustrations of the Anatomy and of Shells 403 10 11 12 13 14 W 4^1 ^^ '"^ 15 16 17 18 ( 19 20 21 22 ^% 25 23 26 24 / f 27 ^^ 28 29 PLATE 83 .

. U. Immature shell. 22-24. X4. Michigan. 15-17. XS^/^... Michigan. C.404 The MoUuscan Family Plauorbidae PLATE 84 Shells of Plauorbidae 1. figs.M. HeUsoma eucosmius vauyhani figs. 11-12 paratypes. HeUsoma rmceps royalense (Walker). XZYzXS^/^. C. voix Co. XSi. Georgian Bay. B995. Lake Charlevoix.N.. Helisoma anceps royalense (Walker). Helisoma eucosmius (Bartsch). Ontario. U. Types. C.. Immature Helisoma anceps portagense (F. Bull. Helisoma eucosmius vaughani U. X4. Ontario. 3. 1-3. Z30844... Toad Island. Helisoma anceps idahoense F. Royale. Helisoma anceps percarinatum (Walker). Paratype. (Bartsch). Baker. Cotype. Burke's 33. 6-8. Immature shell. 21. U.. Place. Isle NE of Jordan. 4-6.N.N. U. Baker. 125719. plate 57.I. shell. U. All figures are enlarged about twice excepting those otherwise listed . 25. plate 57. 5. 61589. B1863. 10-12. Holotype. Helisoma anceps rushi F.M. Helisoma anceps (Menke)..S. Immature XS^/^.M. U. 13. (Bartsch). shell. Z30853. Bamiji Lake.N. Types. Joseph Lake..M. 33. St. Paratype.S.. Ontario. XSV-i- Bull. (Bartsch). Sand Point. Z25239. North Carolina. Charle- 9. shell. lOholotype. Helisoma anceps royalense (Walker).I. Walker.S. eucosmius U. Baker). 193890.14. 4. Helisoma anceps jordanense (Winslow).I. Immature Helisoma anceps sayi F. XSVj.S. C. Idaho. Helisoma anceps royalense (Walker). 26-28. Pend (Jreille River. Louisiana. Baker. 18-20. Topotypes. Helisoma Greenfield Pond.M. B. 2.

-- ^ .fllusf rat ions of the Anatomy and of Shells 405 Y.

. Baker. Helisoma anceps cahni F. C. Aroostook Co. Crystal Lake. Baker). locality. 12 ridged shell. Chemung Co. 14-16. C.. 30. trivolvis Immature shells. U.L. Helisoma Z29134. Quebec.. New York. Portage Lake.. Meach Lake. Canada. New York.L. Wisconsin. Benzie Co. C. Types. Z34976. U. Michigan. Helisoma anceps portagense Meach Lake. Baker). Elmira. LT. Z26276. U.I. Michigan. Helisoma anceps porlagenac (F. (Say). 25-29. Helisoma anceps latchfordi (Pilsbry). Braddock's Bay. increa. Maine.L. 5-7. Big Muskallonge Lake.. Helisoma trivolvis (Say). near 20-24. (F. B988. 17-19. Canada.. U. 8-10. Helisoma anceps percarinatum (Walker). Helisoma trivolvis (Say). U. Type 4. Z37203. X3. Vilas Co. Quebec. Z21124. Showing regular in growth of shell. Type 11-13. B986.406 The Molluscan Family Planorhidae PLATE 85 Shells of Planorbidae 1-3. trivolvis (Say). All figures excepting 17-19 are enlarged about 1% diameters . Helisoma anceps percarhuititm (Walker).. locality.I. Helisoma Z29789. locality. B3686. Douglas Lake. Type Collected by Latchford.I.

Illustrations of the Anatomy and of Shells 407 ) 13 t 17 18 23 24 25 26 .

Illinois. Helisoma Z30538. II.) B930. X7. 19. New chautauquensc F. ChautauYork. holotype). on base. Hclisoma trivolvis (Say). 20. Illinois. Baker.). Type locality. X7. chautaucptoisc F. C. chautauqucnse F. Baker.l. 16.4. 23.). Baker. C. paratype. 17. Z32365.. trivolvis lentum (Say). B3915.1^. Urbana. Chautauqua Lake. megastoma DeKay. Alabama. near Urbana.I. Chautauciiia Assembly. Z27842. X7.. Enlargement of whorls on base.I. All figures. U. Fork. Enlargement of whorls on base. 5-8. Z32472. Louisiana. 9. Type locality. Hclisoma ciua trivolvis Lake. Baker).I. Indian River. Z 11292. Florida. B1085. C. 4. 21.L. U. Hclisoma trivolvis intcrtextum (Sowb. 12.L. B3914. New York. Z23780. 2.10. trivolvis Helisoma Paratypes. Hclisoma trivolvis intertcxtum (Sowb. C. B3917. C. Hclisoma Hclisoma Hclisoma Helisoma trivolvis (Say). 15. are enlarged about I73 diameters . Chaniiuugn Co. F. Enlargement of whorls 22. U. Baker. C.ficlisotna trivolvis intcrtextum (Sowerby). Salt Fork. New York. Florida.I. Heliaoma { trivolvis (Say). X7.. Orleans. New Type locality. {^ Planorbis pseudotrivolvis F. Baker. Michigan.I.) 2. U. Z32367. \j. Hclisoma trivolvis (Say). near =^ Planurbis psiudotrivolvis F. Manatee Rivor.nlargement of whorls on base. U.408 The Molluscan Family Plaiwrbidae PLATE 86 Shells of Planorbidae I. Salt (^ Planorhis... Manatee River. Holotype. 3. excepting those indicated otherwise. Z23780. Cheboygan Co. {^= Planorbis pseudo- trivolvis F. U. U. B3916. trivolvis lentum (Say).. trivolvis (Say). Shoal Creek. 18. 14..).. Hclisoma trivolvis Chavitauqua Lake. trivolvis intcrtextum (Sowb.

Illustrations oj the Anatomy and of Shells 409 .

holotype. HeUsoma trivolvis marshalli F. B3918. Immature. Hclisoma Helisomn trivolvis fallax U. C.. trivolvis fallax (Haldeman). New Jersey. All figiu'es are enlarged about two diameters . trivoJvis fallax (Haldeman). HeUsoma Z41108. U.I.C.. Holston River. Aquarium specimen. Baker. Massachusetts. C. Baker. (Haldeman).9. Virginia.. Virginia. Variation Z41444. Fresh Pond. 23. U.I. Maine.M..24. Xeponset. trivolvis fallax (Haldeman). Smith Co. Z41443. U. D. C.. Milton Mills.S. 16. 18-20. Holston 17.S. Immature. LT.M.M. Massachusetts. HcUsotna B904.. U. 8. 5-7. Port Elizabeth..N .410 The MoUuscan Family Plmiorhidae PLATE 87 Shells of Planorbidae 1-4.S.I. holotype. 17-20. lentum (Say). Virginia. 124989. 22. paratypes... paratypes. Milton. U. Baker. Green Lodge. HeUsoma trivolvis marshnlli F. Baker. 26449. Baker. HeUsoma in color. 14-15. Holston River. 13. HeUsoma Hclisoma U.M. trivolvis holstonense F. C.N. 11-12. Z41444.S. Louisiana.X. HeUsoma trivolvis holstonense F. immature. 33657L 10. Washington. Cambridge. C. 504153. trivolvis 21. New Orleans.N. 11-15. Massachusetts. trivolvis holstonense F.I. River.

IUustratio)hs of the Aiuitony and of Shells 411 r\ 10 11 12 13 14 15 17 18 19 ^ 16 20 6i i» 21 22 23 24 PLATE 87 .

I. B3715. Ontario. C. Canada. 15. Isle.. paratype. trivolvis 7. In beach pool. U. Z41268. Z32361.. C.I. U. locality. 14. B2986. C. Z13677a. Type 11-13. Wisconsin. lectotype. H rlisoma U. Basswood River Rapids. Helisoma pilsbryi Tomahawk Lake. Canada. B843. near Montreal. macrostomum (Whiteaves). 5-6. Helisoma pilsbryi injracarinatnm F.I. Baker. but typical. Baker). Z26268. Z1279L 4-6. Helisoma trivolvis macrostommn (Whiteave. 4. B3870. Helisoma trivolvis trivolvis (Say). holotype. (F.I. Baker. Large form. paratypes. Rainy River District. 3869. Wisconsin.412 The Molluscan Family Planorhidae PLATP: 88 Shells of Planorbidae 1. Michigan.. National Museum of Canada.. 14.2. Helisoma Winnebago Lake. C. Ontario. Baker. All figures enlarged about 1V> diameters . pilsbryi infracarit^atum F. Presqiie macrostomum.s). 8-10.I. Rideau River. Canada.. near Ottawa. trivolvis trivolvis (Say). Mile End Gate. Helisoma U. Basswood Lake. 15. Helisoma pilsbryi infracarinatum F. U. Oneida County. Bayfield. 16. Wisconsin. near Oshkosh. Transition form toward 3.

lUustrations of the Auatomij and of Shells



The Molluscan Family Plwwrhidae



Shells of Planorbidae





Austin, Texas.

Figure 3 varies toward hiter-

textum. U.S.N.M., 252256.

Helisoma lentum

trivolvis intertcxtum (Sowb.).
(fig. 6).

Fort Smith, Arkansas. Variation toward
179 of Pln-

U.S.N.M., 120965.



trivolvis intertextum (Sowb.).

South CaroHna? Binney's


norbis glabratus (not of Say). U.S.N.M., 29219.


trivolvis intertextum (Sowb.). Leon, Texas.

U.S.N .M., 28210.


Helisoma trivolvis reticulatum (Dall) {^intertextum Say). borough Co., Florida. Types. U.S.N.M., 75421.
U.S.N.M., 9010.

Lake, Hils-


Helisoma truncatum (Miles). Saginaw Bay, Michigan.





Pknwrbis bellus Lea (^= Helisoma


lentum (Say)


Type. Tennessee. Lea

U.S.X.M., 121178.
Isle la Crosse, English River, Canada. immature. U.S.N.M., 29231 (old 9272).


Helisoma kennicotti F. C. Baker. Lake


16, 17,


13, 14,


Helisoma subcrenatum (Cpr.). Reservoir near Flowcree, Montana. B3792. Helisoma subcrenatum (Cpr.). Wainwright Park, Alberta.




Helisoma subcrenatum {C]n\). Fort Yukon, Alaska. U.S.N .M., 218908.

22. 23. 24.

Helisoma subcrenatum disjecturn (Cooper). Boise, Idaho. Helisoma subcrenatum plexatum
Helisoma occidentale (Cooper). Klamath



(Ing.). Sagnache, Colorado.
Falls, (3regon.

U.S.N.M., 535328.

U.S.N.M., 219750.

Helisoma occidentale (Cooper). Tule Lake, California. U.S.N.M., 160839a.

26. 27.




Daggett, California.

U.S.N.M., 349083.

Helisoma amnion (Gould). Spring Valley, San Mateo Co., California. U.M., 84105.
(Gould). Clear Lake, California. U.M., 81743.



Aquarium specimens raised by Dr. E. G. Berry. Note resemblance to Helisoma duryi seminole. B3796.
trivolvis (Say).

Figures 18-29, immature.
All figures are enlarged a trifle less than

two diameters

lUustratious of the

Anatomy and

of Shells





The Molluscan Family Plauorbidae



Shells of Planorbidae

Helisoma (runcatum (Miles). Saginaw Bay, Michigan. Type Helisoma truncatuui (Miles).
Spirit Lake, Iowa.

locality. U.I., Z32515.


U.S.N.M., 476566.

Helisoma truncatum (Miles).

Spirit Lake, Iowa.

U.S.N.M., 505847.
Co., Wisconsin.


Heluoma truncatum

(Miles). Sturgeon Bay,



11,12. 13-15. 16-18.

Helisoma truncatum (Miles). Winnebago Lake, Wisconsin.



Helisoma subcrenatum disjectum (Cooper). Tuolumne, California. B1090. Helisoma subcrenatum disjectum (Coojier). Boise, Idaho.



Helisoma subcrenatum horni (Tryon). Paul Lake, British Columbia. B3148. Helisoma subcrenatum variety? Quamichan Lake, British Cohmibia. B1973.
All figures are enlarged a little less than


two diameters

Illustratio)is of the

Anatonnj and of Shells















The Molluscan Family Planorbidae



Shells of Planorbidae

Helisoma winslowi

(F. C. Baker). Little

Arbor Vitae Lake, Vilas Co., Wisconsin.
U.L, Z18637, Z19398.
Co., Wisconsin. B.







pilsbryi (F. C. Baker).

Chetek Lake, Barron


Helisoma pilsbryi preblei F. C. Baker. Knee Lake, Manitoba. Figure
paratypes. U.S.N.M., 180279.



Helisoma subcrenatum (Carpenter). Idaho. Shimek
Alberta. Pathologic specimen.

U.S.N.M., 504310.

Helisoma subcrenatum (Cpr.). Third Vermilion Lake, below warm sulphur cave,
U.S.N.M., 272105.


Helisoma subcrenatum (Cpr.). Lake La Hoche, British Columbia. U.S.N.M., 43346.

Helisoma subcrenatum (Cpr.). Locomotive Springs, Kelton, Utah. U.S.N.M., 308926.

16.18,19. Helisoma subcrenatum

Utah Lake,

2 miles south of Lehi, Utah.


Helisoma subcrenatum variety? Same natum. B1894.

locality as above.

Variation toward subcre-

All figures, excepting 4, are enlarged about V/j diameters

lUustrations of the Anatouiy and of Shells













The MoUuscan Family Plaiwrbidae



Shells of Planorbidae

Helisoma subcrenatum (Cpr.). Newton, Utah. B3046.

Helisoma suhcrenatum






U.S.N .M.,


Helisoma subcrenatum (Cpr.). Cleary, Alaska.

Fos.sil pits.

U.S.N .M., 381941.

Helisoma subcrenatum (Cpr.). Great Slave Lake. Kennicott Helisoma subcrenatum (Cpr.). Fort Simpson, Mackenzie
cott Coll., U.S.N.M., 28378.

U.S.N.M., 9275.


Helisoma subcrenatum (Cpr.). Wainwright Park, Alberta. B3919.


Canada. Kenni-


Helisoma subcrenatum (Cpr.). Creek one mile west
matm-e. B3793.


Devon, Montana. Im-


Helisoma subcrenatum horni (Tryon). Fort Yukon, Alaska. U.S.N.M., 218908.
crenatum. U.S.N.M., 180280.

Helisoma subcrenatum horni (Tryon). Dall Ri\er, Alaska. Vaiying toward sub-


Helisoma subcrenatum plexatum (IngersoU). St. Mary Lake, Mineral Co., Colorado. Figure 17, lectotype, U.S.N .M., 420210; figs. 18-22, paratypes. U.S.N .M.,
(Lig.). Sagnache, Colorado.


Helisoma subcrenatum plexaium

Lnmature. L^.S.N.M.,

All figures are enlarged about



Illustrations of the

Anatomy and

of Shells



The Molluscan Family Planorhidae



Shells of Planorbidae

Helisoma subcrenatum plexatum




Montana. U.S.X.M.,


Helisoma occidentale (Cooper). Tule Lake, California. L'.S.X.M., 160S39a.
Helisoma occidentale (Coojier). "Wa-^hington Territory.'




Helisoma occidentale (Cooper). Klamath



U.S.X.M.. 219750.


Helisoma occidentale (Cooper). Upper Klamath Lake,
Oregon. L'.M., 62741.


of reserva-




Lower end Upi)er Klamath Lake, Oregon.


Helisoma occidentale (Cooper). Slough

Wacos Bay, Klamath Lake, Oregon.


Helisoma occidentcde (Cooper). Uiqier Klamath Lake. Oregon. B3011.
of reserva-

Helisoma occidentale (Cooper). Upper Klamath Lake, south boundary tion. U.M., 62741. Helisoma occidentale depression F. C. Baker.
Helisovia occidentale depressum F. C. Baker.


Lower Klamath Lake, Oregon. Lower Klamath Lake, Oregon.

17, 18.

Paratypes. B3240.

Helisoma binneyi (Tr3'on). Portland, Oregon. Stanford Univ., B3220.

21,22. Helisoma binneyi (Tryon).

Lake Whatcom, Washington. B853.

Helisoma binneyi (Tryon). Holders Lake, Xanaimo, British Cohunbia. U.AL.
All figures are enlarged about



lUustrations of the Aiiatomij and of Shells


Seattle. 22-24. California.M. California. 19. Stockton. 5.N. Helisoma ammoii (Gould). California. Oregon. California...N. Columbia River.X. Immature. 25. Exped. B1888.. 14. 15. (Gould). U. U..M.424 The Molluscan Faniilij Plaiwrbidae PLATE 94 Shells of Planorbidae 1-5. 84105. Baker.S.26. plate 8. I'. Portland. Helisoma hinneyi (Tryon). Immature. fig. San Joachin River. Back view U. U. California. Seattle.21. 35676. U. Helisoma antmon (Gould). B3246.. fig. Immature. Helisoma hinneyi randolphi F. Back view of shell.S. Immature. C.. San Mateo Co. 174051. Helisoma am>no)i U.. immature. 20. TT.S.S. Stockton.M. California. 28. Z18459. Spring Valley.. Oregon. 1.. 5530. Figure 2-4. (Gould). 8119.. 504360. half grown. holotype.M... San Joachin Co.N. Binney's 191 of Planorhis corpulentum Say. Washington. 16..I. Hilisotna hinneyi (Tryon).M. Baker. 81746.. 27.. Oregon. 9-11. U. Helisoma Helisoma ammon ammon (Gould).S.X.S. parat. LT. Near Cerritos. Oregon. Tyi)e. C. figure of Planorhis 18. 81749. Helisoma hinneyi (Tryon).M. Helisoma hinneyi randolphi F. 6-8.ypes. Helisoma hinneyi (Tryon)..M.L.M. Helisoma a)nmon (Gould).13. 81571.. Astoria. 17. ammon (Gould). corpnlentum U. Helisoma bi)tiieyi (Tryon). San Diego. Monterey Co. Z35295. Oregon. About U. 81748. 474779. U. Helisoma amnion (Gould). Gould's Say. California. U. Los Angeles Co. Astoria.. All figures are enlarged about II/2 diameters .M. of Helisoma •shell. San Joachin Co. Green Lake. Washington. San Benito Co. 130.. Monterey.M.M. 12. Hollister. California. Ex.

Illustrations of the Anatomy and of Shells 425 .

.M. B3288. Clear Lake. 6-9. California. U.. California? 23-25. 47616.. 12-14. Immature form. Helisoma traskii (Lea). Tulare Co.11..M. Helisoma traskii (Lea). Lea Coll.21.. Helisoma B9124. 10. U. California. Kern Lake. Tulare Co. of Lea. Watsonville. More depres. Mountain Lake near San Francisco. 17-19. 121000. Type by Binney.. ammon ammon (Gould). 15. LT. .M. LT. Stages U. B3066. All figures are enlarged about P/i' diameters . figured 20. California. California. 16. Z35624. which drained Kern Lake.S. Helisoma 81754.M. 188. Point Happy. Helisoma 81747.M. Cooper Coll. Helisoma amnion (Gould). Bakersfield. draining Buena Vista Lake. California. 22. Helisoma amnion (Gould). Indio. ammon (Gould). Bucna Vista Canal. (Gould). California. Clear Lake..N. Helisoma avimon (Gould).. Santa Cruz Co.N. Helisoma Helisoma traskii Kern Co.I....S. of growth... 81752. (Lea). California.M. California. 81743. Immature.426 The Molluscan Family Planorbidae PLATE 95 Shells of Planorbidae 1-5. traskii (Lea). U.sed than Lea's type. U. Kern Lake. California.

Illustrations oj the Anatomy and oj Shells 427 Obilito 1 6 .

Z. 46502. fig. U. figs. paratypes. Type locality. 92410.M. Sixteen miles northwest of 9-12. Utah. 193321. Coll. 35679. 25.M. magnificum (Pilsbry). U. 31-34 are about natural size.C. Figure 13. North figs.. 13-17. Helisoma Greenfield figs. 19-22.. Wyoming. Anticosti Island. Tennessee. figs.. holotype. 28. plate 57. .M.N. C. 48285. near Wilmington... U. (Tryon). Typo locality. 14-17. Helisoma subcrenatum perdisjectum F. 31-34. Nat. English Bay. the angistomus form of Haldeman. 33. 23-30. M.. holotyi)e.. 6-8.S. Helisoma oregoneiise (Tiyon). 27.428 The Molluscan Family Planorbidae PLATE 96 Shells of Planorbidae 1-3. C.C.Z.Z. B1096. Baker.C. 29. Nevada. Helisoma oregonense (Tryon). 7-9. 18-22. S. paratypes. Figures 24.5. normal anceps. Helisoma anceps (Menke). U. Baker. Gerbach.. Helisoma oregonense B3920. 26. of Oregon. 30207a. immature. Carolina. Mus. 18. Fig. sixty miles 4. Helisoma anceps anticostianum F. (Tryon). 23. in salt spring. Stearns Helisoma oregonense M. Pueblo Valley on boundary west of east boundary. Pond. Bartsch's Proc. Tooele Co. Chevanus Pond. Figures 1-30 are enlarged about two diameters.N. north of Knoxville. Nevada.S. Yellowstone Lake. Variation. M. 30.

Illustrations of the Anatomy and of Shells 429 te#^ ""9 CP 10 ^ 11 12 13 14 15 117 18 1^ \ ^r3 24 16 21 22 23 *( 25 27 26 28 '-•< ^.*- 29 30 31 .^ \^ M^V 33 32 34 96 PLATE .

18-25. Helisoma tenue bourcardi (C. F.N.N.S. Platpa near City U. U. Lake Patzcuaro. Mexico.S. F.M. Helisoma tenue (Philippi). Mexico. Mexico. 22009.S. U. 13-17. of Mexico.S. Helisoma tenue craggeratum (von Martens). and 26453. Helisoma tenue (Philippi).M.X.430 The Molluscan Family Planorbidae PLATE 97 Shells of Planorbidae 1-4. 12. 11.S... U.M. Coll. Toluca. Sources of Rio Lerma.N . 160175. Valley of Toluca. Vera Cruz.S. near Tampico. Lanos River.N. 8506.S. Mexico. 7.s).. near City of Mexico. Helisoma tenue (Philippi).. Helisorna tenue (Philippi)..C.). Helisoma tennc applanatum (von Marten.Z. 10585. U. Vera Cruz.. 467433. 467462. Mexico. City of Mexico. Mexico..M. Helisoma tenue bourcardi (C. U. U.S. about P/o diameters . Strebel Helisoma tenue (Philijipi). Tj'pe locality.S..N. Helisoma tenue (Philippi). Mexico. U.M. 6.N. Lake Texcoco. Mexico. Laguna de Tamos.). and City of Mexico.M. 16252. 5. 9. U..M. Mexico. All figures are enlarged of Michoa- 26-28. Orizaba.M. 467437.N. Type locality.M.8.N. 10. Chichuachua.M.S. State can. Mexico. M.. 185942. Helisoma tenue (Philippi). U.N.

lUustratiotis of the Anatotny and of Shells 431 PLATE 97 .

Santa Cruz River. Sonora. 9-13.S. 10538. C. In 14-. Helisoma tenue ealijorniensc F. 134552. Helisoma tenue calijorniense F. Santa Cruz Mts.. Helisoma tenue caUforniense F. Yaqui River. C. Guadeloupe Creek. 2-6.M. Albuquerque.. Mazatlan.. Strebel Coll. U. 26.N.S.. B3227.N. U. Bixby. Baker.N. Tucson.N.S.. Immature. stream bed. 47572. Santa Cruz. 4 miles from Los U. California. 19-21. 17. 25. 121192.M. California. Sonora. diameters . Rio Grande.S. Helisoma tenue calijorniense F. B3262. 32111. New Mexico. Figures 25 and 26 show differences in spire whorls and sculpture of the two ^'arieties All figures are enlarged about V/^. 15.N. Helisoma tenue applanatum (von Martens). Baker. applanatuni (von Martens). Helisoma tenue calijorniense F. California. C. Baker. U. U. 467429. 22.8. Helisoma tenue 7. Gatos.24.M. California.. Texas. U. Soroyta Creek. U. Crystal Creek.. Santa Cruz. (von Martens). 75437. Baker. M. 47571. Baker.S.N.S. California. Orizaba.M.. 18. Helisoma tenue sinuosum (Bonnet).N. M. El Paso.S. altitude. Lea Coll.S. Helisoma tenue applanatam U. Paratypes. C.S.. Mexico. Los Angeles Co. Helisoma tenue sinuosum U. Baker. San Jose. Arizona. San Jose.M.N. 130229.N.S..M. California. near Cimlad Obregon. Mexico. 23.M. 16. 130237. Helisoma tenue sinuosum (Bonnet).N. U.N. 130239.S..M. C. (Bonnet). Helisoma tenue calijorniensc F. Mexico.. C. U.. Mexico. Immature.M. Helisoma tenue sinuosum (Bonnet). 5000 ft.432 The Molluscan Family Planorhidae PLATE 98 Shells of Planorbidae 1.

Illustrations of the Anatomy and of Shells 433 .

Hclisoma tenue strebelia7ium B3921. Immature. Immature. Hclisoma tenue strebclianum (F.S.S. Imma- U.S. Mexico. 162654.M. Hclisoma tenue strebelianum (F. and C). Mexico.M.S. Alta Mira. and C). 27-30.M. sinuosiini River. Immature. 9.N. 4-8. Tamaulipus. 130237. Los Angeles Co. (F.. 32-36.. Arizona. San Luis Potosi. U.M. Nicaragua. 26. Tucson. California. 174048... Santa Cruz River. Mexico. Baker.. C. San Luis north of 20-24. 109916. Naranjo. U..N.M. U. LT.. Guatemala. Hclisoma ienue applanatutn (\on Martens).S. (F. El Paso. Yaqui Immature.N. Immature.M. Near Vallecitos. 251815. Hvlixoma tenue ture. Hclisoma caribacum (Orb. Tamaulijius.M.. Helisoma caribaeu}7i guatemalense (Clessin). near Ciudad Obregon. 5 miles north of 16-19.N. (Juatemala.M. Hclisoma tenue sinuosum (Bonnet). Hclisoma tenue strcbelianuia U. U. Mexico. Lower California. Mexico.S. 83490. Bixby. Reservoirs 4 miles 25. 83485. All figures are enlarged about 1^/4 diameters . U.M. 11-15.N. Hclifioma tome applanatum (von Martens).10. Penasco..N.. 31.. LT.3.434 The Molluscan Family Planorbidae PLATE 99 Shells of Planorbidae 1. (Bonnet). 251807.S.M. 83487. Varying toward Hclisoma cari- bacum guatcmalense. and C). 2.). 130239.N. Hclisoma caribacum guatcmalcnsc (Clessin). Rio Grande. Texas. Potosi. Guatemala City. 467429.. Helisoma tenue calif orniense F. U.S.M. LT. Coy River.N. and C).

Illustrations oj the Anatomy and of Shells 435 .

.). Ilclisoma 3. Mexico.30. U. U.. hi Helisoma caribaeum (Orb.S. (Tristram).. B3075.). Mazatlan. 25-27.M.. 18-20. Punta d? Helisoma caribaeum (Orb. Nicaragua. U.X. Mexico.S.S.S. U. 360391.M.N.33. U. Vera Cruz. 47517. 210882. Somorrostro. 360392.N. Havana.). San Salvador. M. San Salvador. unjldii (Tristram). Helisoma wyldii (Tristram).. caribaeum (Orb. U..).M.S. Helisoma caribaeum (Orb.5.4. U.X.S.436 The Molluscan Family Plauorbidae PLATE 100 Shells of Planorbidae 1. Cuba.X.S.. 6-10. about \y> diameters . B2284. Helisoma lumens (Cpr..N. U. Mexico. Helisoma tumens (Cpr.M. 121023. Mexico. Junta. Panama. 360390. Helisoma caribaeum (Orb. 46967. ivijidii Guatemala. U.S. Helisoma Helisoma Helisoma unjldii saluini (Tristram).2. Helisomn tumens (Cpr. 83491. Cuba.N.M. 29... 28.). All figures are enlarged 32. 14-17.N. 31. Cuba. 11204. 83492. Mazatlan.N.S. I'.). Lake Olomega. M. Lake Metapan. Mazatlan.M. San Salvador. 354295. San Carlos. Helisoma wyldii (Tristram). Lake Guija.). 21-24.. U. 11-13. 34-37.M.M.). M.

Illustrations of the Anatomy and of Shells 437 18 27 PLATE 100 .

74762. Jamaica.M. Catalina. Parent from type 8.N. Helisoma caribacum (Orb.M.N..M. 13.M. 11. Cuba. Helisoma costaricense (Preston). locality. 74762. 15. U.S. Figured type. Costa Rica. Catalina. Cotype. Utah. Helisoma foveale (Menke).S. (Cpr..6.M. U..N. teniic applanatum (von Martens).S. C.S.. excepting 15 and 16.N.N. M.4.S. 3. of Elbow or Lindbergh Lake. are enlarged about U/2 diameters .N. 2. Montana. Jamaica. 90479.S. 202522. 83491.N.. Helisoma foveale (Menke). 10. Helisoma subcremttum variety? Utah Lake. Guanacaste. Kingston. Foot Mission Mts. Costa Rica.S. To show flat inner whorls.M. U.Z. Kingston. 14.... U. Baker. U.C. bom in capti\-ity. 16. Jamaica.N.S. Oregon. Jamaica. To show rounded inner whorls. Helisoma magnificum (Pilsbry).S. (Preston). Young. X6. Helisoma jovealc (Menke). Helisoma suhcrcnatum. M. 120986. U.. Guanacaste.). 454058. Helisoma jovealc (Menke). All figures. U.N.M.438 The Molluscan Family Planorhidae PLATE 101 Shells of Planorbidae 1. 7. U. Helisoma 10538. From Nuttall in Lea Coll.9. 5. 29332.12.). U.M. U.M.. Helisoma anceps idahoense F. Helisoma costaricense B2132.

Illustrations oj the Anatomy and of Shells 439 •»*. -.^ M Ul^ 16 15 PLATE 101 ...

En- larged.. near Leesburg. Acad. Phil. \k 193. Hdisoma tenue chapaleme (Pilsbry). 13.14. U. near Guadalajara. 117952. B2965. Helisoma contrerasi (Pilsbry). Florida...N. 2. 19. B1106. Acad. fig. Florida. are enlarged about two diameters .440 The Molluscan Family Planorbidae PLATE 102 Shells of Planorbidae 1. 1. seal are (Jay).N. 193. Oaxaca. Everglades..20. i>. All figures. Sci. Mexico. Everglades. B1438. fig. Mexico. Lake B2969. Florida.S..4.12. Helisoma scalare (Jay). Lake Chaiiala. Hclimma Helisoma ten tie ehapalense (Pilsbry).S. Caloosatchee Pliocene formation. Griffin. 5-10. Hclisoma contrerasi (Pilsbiy). B3002. 2.M. Florida. 1920. B2962. excepting 1 22. Helisoma scalare (Jay). Proc. Helisoma scalare (Jay). Head of Miami River. 225863. Enlarged. Sumter Co. Helisoma duryi seminole Pilsbry. 3. Sci. Helisoma duryi seminole Pilsbry. 15-18. Proc. Florida. U. 1920. Everglades.M. 21. and 2. Phil.23. Dade Co. Florida. 11..

Illustrations of the Anatomy and of Shells 441 £ #^% .

. M. Helisoma duryi semiiiole Pilsbry. 68236. Florida. 3-5. Near West Palm Beach. 71599.. Lake Apopka. Florida. Helisoma duryi seminole Pilsbry. Survey. Everglades. Z32368.. Lake Co. Florida.. show the great variation in form of spire and shape of shell of the dnryi complex. near More Haven. Lake Co. Florida. B2962.Z. Florida. 71598.9. . 82856.C..Z. 31.Z.. Helisoma duryi seminole Pilsbry.Z.Z. B2960. M. U.C.S.. 43311. Z32364. Florida.L. Lee Co. M. Helisoma duryi (Wetherby). U. Helisoma duryi (Wetherby). Florida.8. 19. Miami. Okechobee Lake.442 The Molluscan Family Plauorbidae PLATE 103 Shells of Planorbidae 1.M. Lake Eustis.. Helisoma duryi seminole Pilsbry.L. Paratypes. 13.. Helisoma duryi soninole Pilsbry. especially the form seminole. Helisoma duryi semiiiole Pilsbry.. M. 6. Helisoma duryi seminole Pilsbry.C. Florida. Florida.2.N. These figures 20-30. Lake Apopka. Helisoma duryi scminole Pilsbry.7. M. 10. Sumter Co. 11. 12.C. Florida. 153412.C. diameters All figures are enlarged about P/i. Lake Eustis. 14-18. L'.

Ilhisf rations of the Anatomy and of Shells 443 V K J^*' #^* \^10 .

Boynton.C. M. Helisoma duryi normale Pilsbry. 18-22.M..Z. \'ariation toward variety normale. HcUsoma Palm Beach Co. Florida. Near West Palm Beach. Heliscmici duryi (Wetherby). 168921. All figures are enlarged about 1V> diameters .. Florida. 76659. Kissimmee River. Florida. Ever- Z34637. U. M. Florida.444 The Molluscau Family Planorbidae PLATE 104 Shells of Planorbidae 1-3. duryi (Wetherby). of 17. M.. U. Variation toward variety normale. Florida.Z.. 45103. 8-16.. 4-7. 87505. One mile north glades. of Oil well.Z.C. Helisoma duryi normale Pilsbry. Magnolia. Helisoma duryi normale Pilsbry.S.N.C. Canal west Malleated base.. Pinecrest.. Palm Beach Co.I.

lUustratioiis of the Anatomy and of Shells 445 15 (^ 16 10 r^ C 11 17 12 18 19 13 PLATE 104 .

M. near Miami. Dade Paratypes.446 The Molluscan Family Plauorhidae PLATE 105 Shells of Planorbidac 1-3. AH figures are enlarged about lY^ diameters . Florida. Beecher Spring.M. 167040. East shore Lake Okeechobee.Z. Florida.S. Meyers. Helisoma duryi intercalarc form alternalum Pilsbry.C. Co. Helisoma duryi eudiscus Pilsbry. 46572... Helisoma duryi nurtnale Pilsbry. Varying toward intercalare. Lake Co..ssup.Z. Type M.^bry).. Helisoma duryi eudiscus Pilsbry. Helisoma duryi prcglabratum (Marshall). 10. Helisoma duryi iuterccdare (Pilsbry). Florida. Silver Springs. Florida. 4-8. Everglades 40 miles from Ft. U. 12.M.X. 153079. 20-27. . M. Florida. Florida. Wakulhi.13. B3074. 14-16. Head of Miami River. Blue Creek. Marion Co. Palm Beach Co.. locality.Z. Je. M..C. Type locality... Helisoma duryi intercalarc Co.C. Wakullii Co.11. Helisoma duryi intercalarc (Pilsbry).C. (Pil.N. Palm Beach 9.. 83729. U. B3073. 71020.S. Florida. 17-19.. Lake Florida. Canal.Z.

Illustrations of the Anatomy and of Shells 447 .

Helisoma corpulentum (Say). Canada. corpulentum corpulentum midticostatum F.. 4. Ontaria.N.S. Helisoma ivhiteavesi F. Paratypes. Ontario. 331971. B1107. Z32311. Helisoma whiteavesi F. Helisoma ivhiteavesi F.S. C.Z.X. U.L. Youno. Puerto Rico. Ontario. 9-11. Florida.M. 99301. Baker.S. Baker.. U. Canada.M. Canada. 18. U.. Rainy Ri\er District. Immature. 8.448 The Molluscan Family Plauorhidae PLATE 106 Shells of Planorbidae 1-3. Baker.C. C. Ontario. Helisoma disstoni (Dall). 99182. 535394.. Florida. St.X.. M.L... Z40570. Helisoma Helisoma Z32626. U. (Say). Lac des Mille Lacs. Lac des Mille Lacs.L.-'> All figures are enlarged about diameters . C.M. Lac des Mille Lacs. 22.'^. Canada. Young.. Lucia Co. Began as a corpulentum and on last whorl changed form of shell toward that of multicostatuyn. 361737. Helisoma disstoni (Dall). U. 21. P. U. Lac la Croix. Baker. U. Z32312. Ontario. Florida. 12-15.N. Helisoma connnti Dull. Kettle Fall. Vero. Z32313. Palm Beach Co.M. Holotype.I. Helisoma conanti Dall. Canada. Ontario. 5-7. 19.L. 16. Canal. Paratype. Helisoma species? Asecibo. U. 17. Florida. U. Caloosahatchee River.20.. Helisoma duryi seminole Pilsbiy..S. Calocsahatchee River. Florida.I. Lake Kahnipiminanikok. C. Canada. U. Z32628.

Illustrations of the Anatomy and of Shells 449 € 12 ^^ 15 .

Baker. Canada. Canada. Paratype. Rainy Lake.L. Young. C. Baker. 16-18. Minnesota. Holotype. Helisovia corpulentuvi vermilionense F. Lac la 7. Canada. immature. B3014. of the Helisoma corpulentum (Say). C. Paratypes. Z32307. Lac la Croix. Kahnipiminanikok Lake. U. Minnesota. 20. Knife Lake.L. Baker. Young. Z32296.L. Minnesota. Z32308. U. U. Helisoma corpulent lun (Say). 6. Helisoma corpulentum vennilioncnse F. Vermilion Lake.10. Bemidji Lake. U.L.L. Z32518. Baker. 21. Lake Kahnipiminanikok. Vermilion Lake. Young. Croix. Z32475. Canada. C. Canada. Canada. Helisoma whiteavesi F. U. Helisoma corpulenlum (Say). Helisoma corpulentum multicostatum F. Helisoma coripulentum vermilionense F. 8. 9. Lnmature. U. la Croix.L. Minnesota. 11. corpulenlum multicostatiim F. All figures are enlarged about P/^ diameters .L. Louis Co. Canada. Baker. Vermilion Lake.L. Helisoma corpulentuni vermilionense F. Kahnipiminanikok Lake. Resembles Say's figure. Lac Ontario. 12. C. Holotype.L. 15. Z41286. C. A type locality. U. Ontario. A type locality. Rabbit Point. U. Z32626. Vermilion Lake. Ontario. Minnesota. Helisoma corpulentum multicostatum F. Ontario. Helisoma corpulentum multicostatum F. C. Ontario. U. Minnesota. 5. Z32628.. 19. Z32627. west of Sabaskong Bay. Helisoma Ontario.450 The MoUuscan Family Plauorbidae PLATE 107 Shells of Planorbidae 1-4. C. Canada. C. 13.I. St. Hclisomo corpuloition (Say).. B3021. Helisoma corpulentum multicostatum F. Z29828.L. Lake Woods. C. BUOl. Paratypes. Baker. U.. U. Baker. Lac des Mille Lacs. Helisoma cor]>uletdum (Say). Canada. Ontario. Baker. Z32312. Z32626. Baker. 14. C. Baker. Beltrami Co. U.L.

niusf rations of the Anntomy and of Shellt 451 ^W^^is^f^^ r 5 .

B1043. 7-9. locality. Baker. 20-25. U. C. Oakland Co.n. Variation Marl Lake. Cayuga Lake. Hdisoma campanulatum B3860. Baker.L. Lermond. campanulatum (Say). P1068a. 19. Joliet. ferrissi F.N. Ontario. Z20908. 26-28. Helkoma campanulatum type locality. Canada. F. Probable \0. C. Hdisoma campanulatum gan. Baker. Z19877. Holotype. Crooked Lake. (Say).L. White Lake.L. Pl068b. P1068. IlHnois. ferrissi F. Ottawa.N. 18. Baker. (Say). Same locality as above. Michigan. C. U. Para- U. Baker. B3858. Literior Charity Lsland. 120985. Hdisoma campanulatum size. New York.S. (Say)..L. jcrrissi F. 15-17. Hdisoma campanulatum North end Canandaigua Lake.. Odcn. ferrissi F. Michi- 29-31. Hdisoma campanulatum land. C. 32-34. M.L. Michigan. C. in U. U.S. Same locality as above. Bonne Bay. C.M. Helhoma cainpanulatum 212700. (Say). 12-14. Hdisoma campanulatum types. Fair Grounds. Paratype. U. New York.. New York.452 The Molluscan Family Plauorhidae PLATE 108 Shells of Planorbidae 1-6. ferrissi F. Baker. Type North end Cayuga Lake. Will Co. Hdkoma B3859.. Hdisoma campanulatum fcrrmi Hdisoma campanulatum U. Newfound- All figures are enlarged about two diameters .

Illustrations of the Anatomy and of Shells 453 % 32 33 34 PLATE 108 .

.454 The Molluscan Family Planorhidae PLATE 109 Shells of Planorbidae 1-6. Lake. Ottawa.. Canada. 346627. Exaggerated campanulate lip. midtirolvis 26-28. 17-20. wisconsinense (Winslow). Howe Lake. Wisconsin. Helisoma campanulatum smithii (F. C. Baker. Helisoma multivolvis (Case). C. Holotype. B1719. 14... Howe Lake. Michigan. B1056. Michigan. 29. Helisoma campanulatum smithii type. Type locality. Marquette Co. Canada. HoloDouglas Lake. 346627. 8. form. 12. C. C. Kent Co.S.11. Michigan. B3922. Small Helisoma Z23192. U. Baker).. Douglas Lake. Z18421. Helisoma campanulatum (Say). (F. Rideau River. Helisoma campanulatum B1718. Marquette Co. Wisconsin. Helisoma campanulatum wisconsinense (Winslow). 10. Tomahawk Lake. Douglas Lake. Baker). M. Wisconsin.. U. Michigan. Baker). Tomahawk Tomahawk Tomahawk 9.I.S. Wiscon. B3861.S. Helisoma campanulatum wisconsinense (Winslow).sin.. Helisoma campanulatum rideauense F. (F. C. B1728. Ottawa. Baker). 21. Michigan.N. U. Douglas Lake. Helisoma cantpanulatum wisconsinense (Winslow). Pine Island Lake. Lake. Michigan. All figures are enlarged about iy> diameters . Helisoma campanulatum wisconsinense (Winslow).M. Baker. Type locality. C. Helisoma campanulatuui smithii types. Little 7.16. B1054.13.I. (Case). Helisoma campanulatum rideauense F. Helisoma campanulatum smithii (F. U.. Rideau River. Arbor Vitae Lake. B3857. B1725. Wisconsin. Paratypes. 272274. Para- 15. 25. Co.N. Michigan. Vilas Lake.N. 22-24.M. U..

1^ J n 'S^ 26 I (S 28 27 29 PLATE 109 .niustrations of the Anatomy and of Shells 455 *^.

Baker.. Holotype. 8-10. B1056. Small form. Z41303.456 The MoUuscan Family Planorhidae PLATE 110 Shells of Planorbidac 1-4. C.L. 13. Marlboro. Holotype. U. Ontario. Sturgeon Lake. Variation in size.. Baker. Jersey.S. Canada. 5-7.N. C. C. C. U. Helisoma campanulatum smithii form. U. Doughis Lake. Baker. Helisoma campamilatum canadense F. Lake. Bamiji Lake.L.. New 22-26. 15-17. Z40064. Tomahawk C. AVhite Pond. 1. U. (F. Baker.M. Brent Lake.N. Z30721. 18-20. Z29858. Helisoma campanulatum near variety smithii Michigan. All figures are enlarged the about two diameters . Z41297. 21.L. 3. Bamiji Lake. Helisoma campanulatum canadense F.I. Paratypes. C.M. Ontario. Marlboro.L. C. Wisconsin. U. U. Canada. U.. Cameron Lake. (F. Variation in size. 121195. 12. U. Helisoma campanulatum canadense F. Ontario. 14. Immature. Helisoma campanulatuin canadense F. Z32331. Ontario. Canada. Canada. Z30721. Variation in size. U. Baker). C. Small 11. Helisoma cainpanulatum wiscon-'iinensc (Winslow). Z13957. Lake near Lake Mercutio. New 27-35. Baker.L. Helisoma campanulatum pleistocenicum F. 2.I. Jersey. Canada.S. Michigan. Douglas Lake. Ontario. Baker. LT. 121195. of Helisoma campanulatum collinsi F. Baker. Helisoma campanulatum canadense F. White Pond. Baker). Helisoma campanulatum pleistocenicum F.L. Canada. Lake Woods. Paratypes. C. C. 4. Baker.

.\h lUuslratnn.. oj the A„at..^ 32 33 34 35 PLATE 110 .and 0/ Sh.3^S 11 10 12 13 14 15 16 17 18 ^*^ 20 19 21 22 #^ 23 e 24 25 26 27 # 28 _ '^i.u 457 cm r V-^/ .l 29 C^ 30 31 '/ ^^.w.

Baker.M. C. X31/2. Helisoma scalare (Jay). Marl bed 10-15 Variation toward seminole form. Paratypes. Everglades.. U. Canada. 18. Michigan.N.S. Canada. Helisoma carnpanidalum michiganense F. holotype. Michigan.S. 13.. 23-31.M.. dalli F. Lake Apopka. Florid. 15. excepting 21 and 22. Emmet Co. Hclisoma campanulatum michiganense F. Hclisoma duryi seminole Pilsbry.S. Z41297.S. Hclisoma camptuiulatuin rudentis (Dall). U. 566570. Helisoma campanulatniii colUnsi F. U.458 The Molluscan Family Planorhidae PLATE 111 Shells of Planorbidae 1-5. are enlarged about 1% diameters .M. Baker. Cameron Lake.M. U.9. B3868. 12.N. 162724. Para- 17-19. All figures. 365574. i/(7i.N. Marl Lake. Young. 7. paratype. 21.22. Lake Woods. Roscommon Co.. 19. Canada. Helisoma cainpaitidatum michiganense F. dalli F.16.. of the 20.. U. Michigan. Michigan.. 162724.I. 32. Helisoma campanulaUim collinsi F. Helisoma campanulatum types. C. U. 173102. Paratype. C. Michigan. LT. U. Canada. 6. Baker. Co. 6. C. Lake of the Woods. Showing advanced lower part of aperture in contrast with the receding aperture of scalare (fig.som« campauulatum ))iichiganeyise F. B1809a. 12. Knee Lake.N.M.S. Baker. C.N.N.i. Cameron Lake.. holotype. 173102. Baker. U. C.M. 17. Baker. A small specimen. 8. C.M. Florida.S.S. I.. Roscommon Marl Lake. Z41297. 348566.. ft. Anticosti Island.13. 14. Ontario. U. Manitoba. C.S. Abnormal.. under peat.N.N. 173102. Crooked Lake. holo- paratype. Helisoma campanulatinn type. 31). Canada. Marl Lake. Marl Lake.M. 11. B1809. 10.. paratypes. Baker.7.. Helisoma catnpanulatum (Say). Anticosti Island. Baker.

.Illustrations oj the Anatutny and of Shells 459 £b| 4 i\ 9 € 10 12 11 13 /-^•.i^ 9% 4 4 c#rf 22 23 24 ^^^"^ 27 28 29 30 31 32 PLATE 111 .

Helisoma duryi seminole PiLsbry.. Helisoma duryi seminole Pilsbry. scalare-Wke forms.. 120331.M. Carinijex occidentalis Hanna.S. Lasen Co.. U. 251516. Range of variation. 9-11.N.M... California. 120333. California.S. Carinifex newberryi (Lea). 12-14. 7..N.S. California.8. M. U. Lake Apopka.N. 16-21.M. Hanna. Carinifex occidentalis Owens Lake.S..M. 506868. Florida.M. 15. Eagle Lake.S. 506870. Eagle Lake. U. U. Carinifex occidentalis Hanna. U. U. California.N. Eagle Lake.S.N.N.460 The MoUuscan Family Plauorhidae PLATE 112 Shells of Planorbidae 1-6. Florida. Lake Apopka. Figures 12-21 show range of variation All figures are enlarged about 1% times . 120331.

rf^*^.1 11 ust rat ions of the Anatomy and of Shells 461 2 h \ 10 .a 12 11 n ^ 14 18 ^m^ 15 16 17 ^---s 19 20 21 112 PLATE .

Type locality..S. Carinifex Canoe Creek. 5. 170772. California. U. California.N. 7. Carinifex newberryi malleata Pilsbry.462 The Molluscan Family Planorbidae PLATE 113 Shells of Planorbidae 1-4. U.N. Carinifex newbcrryi (Lea). Carinifex newberryi malleata Pilsbry. U. U.S. Canoe Creek. California. 349132.. 2954. Carinifex newberryi (Lea). 29. Binney's fig.M. California.11. U.N. Pitt 30. LT. 120. California. 56405.S. B3877.8.S. probablj^ types.M.. 9256. Nevada.S. U. 24-28.6. 9342. See 9. 12-15. Bear Lake. Canoe Creek. Carinifex newberryi (Lea). 75.S. California. Utah.. Carinifex newberryi malleata Pilsbry.. Canoe Creek. See 9254. Lea Collection.M. newberryi (Lea). U.N.M.N.M.N. 16-23...M. 9341. 122. River. All figures are enlarged about 1% diameters .S. 10.. Ccirinifex Binney.M. California.S. Pitt River. Carinifex newbernji (Lea). newberryi (Lea). Binney fig. Similar in form to malleata.N . p. Said to be types.S.N.. Canoe Creek. Walker Lake. Pitt River.M. U. California. 120991. Lakota Resort.M.N. Carinifex newberryi (Lea).

niust rations of the Anatomy and of Shells 463 t^ e 10 n 14 12 13 '^' 1^ 15 16 n \% 19 0' 20 21 22 r / 24 23 25 ^^^ ^fl 27 28 29 30 PLATE 113 .

Lower Klamatli Lake. South end Upper Khimath Lake.N.S. Carinijex 5. Carinijex neieherryi (Lea). Lower Klamath Lake. ponsonhyi E. 2. 18. 2. 17-24 are enlarged 3 diameters. U. li/-> diameters.. Proc. type. Smith. U. Immature.11. Falls. Klamath Falls. A..N.N. U.N.N. newherryi subrotuncla Pilsbry. ponsonhiji E. Immature. B3924. Carinijex figs. 86. Carinijex ponsonhyi E. Eagle Lake. B3924. A.N.. A.6. A.. Smith. Oregon.M.S. 19-24. Sci. Carinijex ponsonbiji E. A. 334387. 4.N.. California. U.N. 380807. 5. Carinijex ponsonhyi E. 510024.15. Canoe Creek. Oregon.. 12. Oregon. U. 16. Carinijex ponsonhyi E. Smith. 219749. 2a. Smith. U. 7-9. Oregon. 14. Phil. plate 11. IT . U. 17.M..N. 3. Klamath Lower Klamath Lake. Upper Klamath Lake. South end Upi)er Klamath Lake.S. paratype.S. 121025.M. A. U. Inunature. Acad.464 The MoUuscan Family Plauorhidae PLATE 114 Shells of Planorbidae 1. California. 337022.S. Oregon.S. A. California. Carinijex occidentaUs Hanna. Oregon. Immature. Carinijex ponsonhyi E. California. 120333. Carinijex Lower Klamath Lake. 2b. Klamath Falls. Figures 1-13. From Lea coll.S. A. Lower Klamath Lake. Carinijex newberryi (Lea).N. Clear Lake. Smith. 25-28 are enlarged about figs. 56405. M. 9256. 25-28. Smith. U.M. Oregon. Smith.M. Oregon.S. immature.S....M.S. 3. . Smith. 13. 334389. 10. Nat.. M. Carinijex newberryi minor Cooper. M. Canoe Creek. Carinijex 'ponsonhyi E.M.

IUustratio)is oj the Anatoiny and of Shells 465 W ^w 16 ^^^ 14 %^^ 17 15 18 1i^% ^ 21 25 .

M. dalli F. U.N. B3869. 5-7. 13 miles north of Klamath Falls. Baker. Dalles of the Columbia River. From type lot. Baker. Jackson's Lake.N.. 23. locality... C. Figures figs. 406024. M. Guatemala. Holotype. Klamath Falls.M. Parapholyx effusa dalli F.S. From type with diagonal grooves. Lake Miragoane.state. 18. 28-30. 1-7. U. Helisoma caloderma (Pilsbry). 8. 10-13.N.N. Parapholyx klamathensis F. 406024. U. Klamath Falls. Baker..M. Henderson.S. 2-4. lot. 510045. Oregon. Haiti.M. 219748. From type lot. C. Wj-oming.S. Helisoma caribacum (Orb.M. U.S. Klamath Baker.S. Parapholyx effusa 219747.N. slightly co. Same locality as above.Z.. Small form. Henderson. 24-26. Type 27. Guatemala. Porapholyx klamathensis F. 21-23. Helisoma eyerdami Clench and Aguayo. Paratypes. Parapholyx klamathensis F. 9. U. U..N. 8-23 are enlarged about two diameters. 20. Wyoming. C. No. Falls.S.C. 219747. Holotype. East side Upper Klamath Lake.). Oregon. Paratypes. Same locality as above. 406024. Z28577.S.S. From type U. Baker.S. Carinijex jacksonensis J. U. C. Parapholyx klamathensis F. .S. Baker..N. Parapholyx effusa costata (Hemphill)... From type lot.N. 406024.M.I.N. Jocolo. lot. C.N. Oregon.M. Carinijex jacksonensis J. Parapholyx klamathensis F. 14. 2 miles southeast of Miragoane.M. 47520.. 24-30 are enlarged about four diameters. Oregon. 83888. Immature.. Cotypes. U. 15-17. Same locality as above. C. Baker.466 The Molluscan Family Planorhidae PLATE 115 Shells of Planorbidae 1. Parapholyx klamathensis F. 19. C.M. U. costate. B3925. Esmeralda. Baker. Jackson's Lake.. Same locality as above. U. Paratypes.M. Oregon. 406024. C.

lllusiratioDs of the Anatomy niid of Shells 467 ^9 ' 10 ^ i^ \ ij^ 12 13 15 16 17 14 21 18 19 20 i4i^ 24 25 26 m 23 ^.<fe 22 W 27 28 29 30 PLATE 115 .

White Pine. 19. U. Type locality.. U. California. 9-11. T.M. of the Parapholyx effusa costata (Hemphill).S. U. Sacramento River.M.26.N. Eagle Lake.M.N. 4-6. 3. 56409.vpe type lot.S. Oregon.S. River. All figm'es are enlarged about three diameters . Dalles of the Columbia Oregon.M.. 47652..M. Nevada. M. U.S.N. Probably U.N. Henderson.8. Sacramento River.N.S.M. U. Type locality.S.N. Nevada. Oregon.. California.N. J. Smooth form..N. 36617. Parapholyx solida (Dall). localit\'. 24839.M. 21.M. locality. White Pine.N.2. Oregon. Crater Lake. Oregon.S.20.N. Parapholyx solida (Dall).S. U. 520085. 121167. Oregon. 23. Dalles of the Columbia River. U.S. White Pine. Type localit.S. Parapholyx effusa (Lea). Type 12. 75438. Nevada.13. 36615. Dalles of the Columbia River. 18. Nevada. 24. 56409. Parapholyx solida (Dall). slightly costate. U.M. 22.y.. Columbia River.. 15-17. Parapholyx effusa (Lea). cffusa (Lea). Oregon City. Dalles Smooth form.M.S. Parapholyx effusa neritoides (Hemphill). Lectotype. White Pine.. 7.. 14. U. 47519.S.25. White Pine. Nevada.468 The Molluscan Family Planorbidae PLATE 116 Shells of Planorbidae 1.S. Parapholyx effusa costata (Hemphill). 27-30. U.. 37518. Paratypes. Parapholyx effusa neritoidcs (Hemphill).M. U..24. Parapholyx 121133.N. Type locality..N. Parapholyx effusa diagonalis B3926. Parapholyx mailliardi Hanna.N.. Parapholyx solida (BaW). California. U. Parapholyx solida (Dall).M. 56410.

Illustrations of the Anatomy and of Shells 469 fr # ^* " w jP •^ 13 C" 10 11 -r*^^ 14 12 y ^^^ 16 17 18 ( 19 20 21 22 ^B^^^^MMf' 23 24 25 26 ^^^^^ 27 28 29 30 PLATE 116 .

Nevada.N.M.A. Pyramid Lake.M..M. 75440. Parapholyx Figm-e 1.N. Nevada.470 The Molluscan Family Planorhidae PLATE 117 Shells of Planorbidae 1-6. Nevada. 2-5.S.M. Parapholyx nevadensis form. holotype.... Parapholyx nevadeyisis Henderson.N. Nevada. paratypes. U.S. 7-14. Scalariform specimen. 15.N. 75439. J. All figures are enlarged about 3y> diameters .S. U.. Eagle Lake. Parapholyx nevadensis J. U.N. 27-30. costate specimen. From type lot. Parapholyx mailliardi 23040. 63493. pusilla F. Parapholyx nevadensis J. U. 16. Six miles west of 6.. Nevada. Pyramid Lake. \'ariation in Hanna. 126765. Pyramid Lake. 308925.M. C. Pyramid Lake. J.S. Baker.S. Henderson. 17-26. C. Henderson. Pyramid Lake. LlS. Henderson. California. neritoid to bulimoid.

Illustrations oj the Anatomy and of Shells 471 ^^ ^«^ -• 0W 9 10 11 ««iP^P^>^F ^9 v^ 21 22 ^ 23 26 ^(r 24 ^ff^ 25 27 -^ \ 28 29 ^- 30 platp: 117 .

Manitoba.M. 27-30.I. Baker. Planorbula armigera indianensis F. Ahibama.I. Planorbula armigcra (Say).N.. Wayne Michigan. Stevenson. Figures 1-8 show the folds within the aperture of each species 8. Michigan. 12. 31.M. C. 527656.. C.. Mt.S.N.M. Medford. East Nebraska.S. Z27094a. Planorbula smithi F. U.N. U. 33.S. Indiana. about four diameters .S. East Nebraska. Iowa. Indiana. Z27086.N . U.. B1195. B1195. Planorbula armigera (Say). Planorbula smithi F. U. U. F.472 The MoUuscan Family Planorbidae PLATE 118 Shells of Planorbidae 1.M. 3. 321196. Planorbula jenksii (H. Planorbula armigera palustris F. Figures 31-33 show condition of crest behind aperture All figures are enlarged U. 20-22. Planorbula smithi F. U. Mason City.M. Planorbula jenksii (H. Rhode Island.N.N .. Indiana. Douglas Lake. Stevenson.S. 63393. (Walker).S. Planorbula crassilabris locality...S.N. U.. Planorbula armigera variety? 17-19. Specimen with double parietal lamella.N.. Type B1200. White Co. High Bluff. Planorbula jenksii (H.M. U. 11. U. East Nebraska.N. 5.S.M. Porte Co. Planorbula armigera Camp Colfax. Brookston... 7. Planorbula crassilabris (Walker). 251213. B1194. 26. 6. 9.. Iowa. Stevenson. 41411.S. Brookston. 4. La variety? Camp Colfax. Carpenter)... C. U. Planorbula crassilabris (Walker). Baker. Carr's Slough. Nayat. 23-25. 2. Baker.N.I. Hamtramck. Planorbula wheatleyi (Lea). U. 321196. Z27086. Co. Z2891. U. Alabama. C. U..M. Baker.S. U. C. Carpenter).M. Planorbula crassilabris (Walker).M. Boligee. 526054. 32.M. U.S. Rhode Ishind.M. Alabama. 321196.I. 526054. B1194. F.. Carpenter). Indiana. Pathologic. Baker. 10. 526054.N. Figures 9-12 show the form of the parietal lamella in three species 13-15. F. 527656.. Bohgee. East Nebraska. Alabama.. U. 526054. Vernon. 16. Massachusetts. Planorbula armigera variety? Big slough near Mason City. Planorbula armigera (Say). Z27094b.. Nayat. Planorbula armigera (Say).S.N. Planorbula campestris (Dawson). Indiana.I. Alabama.. Planorbula ivhcatleyi (Lea).

Illustrations of the Anatomy and of Shells 473 10 11 12 13 16 20 23 17 14 21 24 ^^ 18 15 22 25 ^im 19 27 29 26 28 31 32 33 30 PLATE 118 .

16. 23-25. C. U. 16-25 1% .. 63393. U. Manitoba.8. Zapata Swamp.N. 2 miles southwest holotype. 321196. Indiana. 11. Type. C. Catoma Creek. Pkmorbula wheatlcyi (Lea). Stevenson. figs. 4-6...M. Holotype. C. layers of peat.S. M. Planorbula jenksii (H. 22. Lake La Hache. Catoma. Alabama.S. Helisoma caribaeum cubcnse F. 10. High BlutT.S.N. Medford. Uniontown Road.M.15. Massachusetts.S. U. 2. F. Co.C.. U. Mt.S.S. Co.S. Dist. 13. U. holotype. Baker..474 The Molluscan Family Planorhidae PLATE 119 Shells of Planorbidae 1-3.. 1. Planorbula smithi F. Caribou Columbia.M. Planorbula armigera of iiidianeyisis F.X. Stevenson.M. Alabama. paratypes.. Baker.N.M. High Bluff. 7.N.. P2833. Helisoma columbiense F.I. 14. Carpenter). 3. Baker. holotype. Paratypes. paratypes. 10. Alabama... Vernon. plate ii. Cuba. Manitoba. Planorbula smithi F. 41411.M. Montgomery 29292. Baker.N. British 22-25. Planorbula irhcatlcyi (Lea). Planorbula christyl (Dall). 321196.N..M.S. Planorbula christyl (Dall) {^ campestris Dawson). C. 12. U. 404719.N.Z. U. diameters. figs. Figures 1-15 are enlarged about are enlarged about 3^/4 diameters. 1(5-21. M. 120929. Paratypes. U. 17-21. Figured type. 9. 93710. Marl between U.. Baker. Alaska Moll. C. 11.. Montgomery Alabama. 63393.. paratypes.

Illustrations of the Anatoiity and of Shells 475 .

F. = 13. Bellingham.N.S.N. Baker. Salem. Fidalgo Island.. Puget Sound..12. Lake Washington.. 19-21. C. 9118). Menetus cooperi midtilineatus (Vanatta). San Francisco.S. 27. U. (Vanatta). Menetus cooperi Menetus cooperi Hemphill coll.M. 29280a. Menetus cooperi 169070.M.N. Parapholyx solida optima (Pilsbry). Washing- ton. 23-26. Menetus opercularis (Gould). M.S. Menetus cooperi type of Whidby's Island. Washington.S. Lake Union. rnultilineatus (Vanatta).N. 29-31. C. Baker { Sound. F..S. U. U. 152185.S.S.S. C. C. Seattle. C. Whidby's type of Puget Sound. U. M. Menetus cooperi F. 216936. Whidby's Island. 32. Very flat spire. planulatus.N. 47630. C.N.M.N. F.. Oregon. Baker. Baker. Figures 1-31 are enlarged about four diameters.S. IT . 29280. Co- planulatus. Menetus cooperi 525100. 346742.M. C. Olga.S. Island.. 22.M. Mountain Lake.M.. Binney's 209 (No. Menetus cooperi F.476 The Molluscan Family Plauorbidac PLATE 120 Shells of Planorbidae 1-10. U.S. U. Washington. U.. Type locality.N. Freeport. Menetus cooperi Pass Lake.P. A. 11. 32 enlarged two diameters. 15-18. Washington. U. M. 47627.. Cotypes.N. callioglyptus Washington. Baker. 253617. is fig. California. Baker. Washington. Type Lake Bigler. 14. Nevada. U. Paratype. 28.M.. Washington. M. Holofig. . Menetus cooperi planulatus Cooper). Baker. B3927. Puget F..N. 125363.N. Washington. locality. Washington.M. F.

Illustrations of the Anatonnj and of Shells 477 \ o hi\\ 11 10 12 ^ 17 21 13 14 15 16 18 19 20 22 23 24 /^ ^^^i 25 26 9"' 29 30 ^ 27 31 120 28 ^^ 32 PLATE .

Massachusetts. near St. 214590. Type coll. paratypes. 85913. Terminal Island. 99302.M.25. 24. (PiLsbry)..M. 29. . Harbert. Menetus ceniervillemis (Tryon). California.S. 43-44.M. (Pilsbry). Menetus Menetus alahaniensis U. Menetus dilatatus buehanensis Cincinnati.M.S. M.. coll. U. C.N. U.S. 88030. dilatatus buehanensis (Lea). dilatatus Mt. 85913. (Pilsbry).M. Michigan. Alabama. Jackson Co. Holotype. Woodville. Massachusetts.. B168. 190021. Alabama.20. Baker.S. Ohio.N. Woodville. U.. 120981. Portland.M.S. Novo.N. holotype. Menetus labiatus F. M.478 The Molluscan Family Planorhidae PLATE 121 Shells of Planorbidae 1-5. U.M. 26. 42-44.. Part of tyjie lot. U..S.M. District of Columbia..M.S.. 42.M.M.27. 473961. 473961.N. Oregon. Meyietus cooperi crassilabris F. Paratypes. Menetus dilatatus buehanensis (Lea). 37.N . Los Angeles Co. Type locality. portlanclensis F. U. U. U. San Bernardino California.S. Seattle.. 47639. 21. Florida. Pleasant. Menetus alabamensis avus Hammock Johns River. crassilabris F. Woodville. 121002.N. Cincinnati. C. Terminal Island. Woodville.S.N... U. Lea (Lea). Ohio. 101698.N.M. Three and one-half miles southwest of Cambridge. Virginia. coll. Menetus labiatus F. holo- 13-18. Menetus alabamensis Menetus alabamensis U.S. 40. 12. 19. California.N.. C. dilatatus Medford. 31. Virginia. C.N. Mt.M. Half mile west of Luray. Menetus brogniartianus (Lea). lectotype. Hingham.28. Baker.N.S. U.. Lea paratypes.M. 9. U. 22.S.S. 19.. U.. Tanner Creek. Menetus ccntcrvillcnsk (Tryon). (Pilsbry).M. Baker. Maryland.41. Menetus dilatatus pennsylvanicus PiLsbry.30... U. 33.N. Paratype.. 190021. 175089. 227832. Type locality.. 10..36. Alabama. B3928. 32. U.s. C.. 41725. U. Near Luray..35. 11. buehanensis (Lea). Menetus dilatatus (Gould). Berrien Co.N. Oregon. 38. 34. locality. paratype. 184182. Bear Lake. U.S. Menetus type.N. Alabama. 20. 23. 120948. 39.M. 12-18. Los Angeles Co. Varia- tion toward dUatatus...N. Menetus Menetus buchanenMs (Lea). Lea Types. California.M. Washington. Pathologic. Menetus cooperi Portland. Menetus dilatatus (Gould). Baker.N. U. 129005.S. Baker. 6-8. Menetus alabamensis (Pilsbry).S.

Illustrations of the Auatoiiiy ruid of Shells 479 ^^^^^# 10 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 ^' 35 ' ^i 38 36 37 ^^ 39 42 40 41 43 44 PLATE 121 .

N. Promenetus exacuous (Say).. Menetus cooperi F. Promenetus exacuous mcgas (Dall). holotype. 3-6. 13. Hospital Creek.2. 24.M. Promenetus exacuous (Say). Oregon. Baker. Colorado.M. Baker. 23-25. San Mateo Co. (Vanatta).S. C. U. 19. California. of Menetus cooperi F. Orcas Island. U. Menetus cooperi F. 14. Vermont. West Lake Okoboji... Swamp. Two miles north 531171. 9-12. Menetus cooperi planospirus F. Figure 5 shows well the very flat spire. C. Petaluma. 3. C. 10114. Baker.M. Blue Lake. U. Baker.C.. Colorado. Types. paratypes. U. Figure 5. California. 336591. of Menetus coloradoensis F. B3929. 20-22. Eldora. Washington. near holotype. head Eldora Lake. paratypes.S. Grand Coulee.. Figure 25. All figures are enlarged about four diameters . 475975. B3930. 526.N.C. 4. 6.N. Menetus cooperi multilincatus (Vanatta). British Columbia. M. Salem..S. Iowa.N. New York. Manitoba. Promenetus exncuous (Say). U. Birtle.480 The Molluscan Family Planorhidae PLATE 122 Shells of Planorhidae 1. Promenetus exacuous variety? Gresham Lake. 23. Menetus cooperi calUoghjptus B2030. C. Boulder Co. between Gresham and Ward.S.Z. Baker. Washington. Chimney Point. Type crassilabris locality. B3935. 8. 525095. 31583. Puget Sound. C. U. 7.. Lake Champlain. Quamichan Lake. 15-18..N. 26-28. U. New York.C.M. Colma. 63391.S.M.

Illustrations oj the Anatomy and of Shells 481 PLATE 122 .

Sumter Co.N. Florida. Figures 2 to 10 are enlarged about two diameters. Vol. 12. Illinois. 23-25. Baker.. 211002. Cotypes. 3. 27. locotypes. 14 miles north of Klamath Falls. San Mateo Co. holotype.S. four diameters. U. half mile south of Fountain..28. M. Types. 531029. Phil. 14-16. 22. fig. 2. figs. Promenetus hudsonicus (Pilsbry).C.. Meredosia. C. C.N. Johns River. 27 to 30. Types. near Hamilton. figs.. near Sedalia. Temporary pond in Flat B3923. U. f^g. Figure 20. Oregon. Creek bottom. 37-39.N. B2018. B2101. U. near Sedalia. Proc. C. near Hamilton. U.). Palatka. 2-10. 669. in Flat II.35. Devonshire Swam]). Vol.C. 62. M.13. Wisconsin.). Bermuda. Parapholyx klamathensis sinitsini F. Colma. 31-33. Proc.. L"'^pper Dells. (Ckll. Zi. New York. 29.. California. South end Lake Panasofflee. 36.M. Phil. Kilbourn. but enlargement of shells is about . Creek bottom. Menetus sampsuni (Ancey). Promenetus imus (Vanatta). figs. Sci. Menetus uUgmosu-s (Vanatta). 37 to 39.M. Menetus cooperi F.S. 1. Figure 2. beach pond. one-half size of original figures. 31 to 33.N. Ditch south of New Philadelphia.482 The Molluscan Family Planorbidae PLATE 123 Shells of Planorbidae I. Bermuda. Menetus sampsoni (Ancey).S. paratypes.. M. Pro7nenetus umbilicateUus Plummers Point. Lake Butte des Morts. p. 21.M.Z.S. holotype. Menetus sampsoni (Ancey). Promenetus umbilicateUus Promenetus rubellus B160. 3. B166. Baker. 4900. B2102. (Pilsbry). p. Menetus dilatatus floridensis F. 669. Acad. 4. five diameters. Missouri. paratypes. Wisconsin. Bermuda. (Sterki).. (Ckll.. in 17-19.M. crassilabris 26. Acad. Pembroke Marsh. Pond Type locality. 11 to 26. Figures 5-10. Pond Part of type lot. Ohio. Baker. U. 531064. Sci. 108996.30. east side Canandaigua Lake. 34 to 36..N.S. B2904. Menetus alabatnensis Illinois. St. (Pilsbry). 37585. Four miles southeast of Canandaigua. Menetus uUginosus (Vanatta). Bercley Spring. 20-22.Z.. 525095. six diameters. Menetus alabamensis avus Florida. Missouri. 62.

^ * 5 ^6 ## 9 13 11 10 14 17 20 27 23 ^EL 18 ^is0 21 ^BR 24 28 15 ^B^ 19 22 25 29 16 32 35 30 » 26 33 36 38 41^ 31 ) 34 37 39 PLATE 123 .•«a>...Illustrations of the Anatonnj and of Shells 483 •i*^« 2 3 %r< .

.C. Baron Hills. Buena Vista. 21-28.Z...S. Bolivia.Z. probably part of original lot of haldemani. 29.).. 14-20.S.N ..) M. (Orb. Brazil.C.M. 72832. Tuccas. {^ Planorhis haldemani C. Venezuela. Adams. 348535.. 87164. Drepanotrema parapseide 87196. Dept.M. Peru.M. Juninaguas. (Orb. 7. Dept. U.484 The Molluscan Family Plauorbidae PLATE 124 Shells of Planorbidae 1. Drepanotrema anatinum (Orb.). Juninaguas.).).). Peru.). 83729. Type locality of Planorbis esperanzensis Tryon.M.N.M.. Baker.N. Brazil.). Drepanotrema anatinum 381510.S. Cuba. LT. 8-10.S. B.) Lea Collection. Loreto.Z. Trelawny. Para.. Jamaica. lot. Jamaica. Loreto.. U. Drepanotrema lucidum 381510. two miles SE Miragoane. M. 3.Z.C. M. Drepanotrema ahenum H.N. 4-6. B. 94758. U. with All figures are enlarged about foiu.). Drepanotrema anatinum (Orb. showing age U.M. {^^ Planorbis haldemani C. Lake Miragoane.31. 11-13. of Drepanotrema anatinum M. Esperanza. Santa Cruz. Adams. Drepanotrema anatimim Drepanotrema anaiinmn Haiti. 30.32. Type variation in shell. Drepanotrema anatinum (Orb.C. B. (Orb.2. U.. (Orb. 65839. (Orb.).diameters . (Pfr. 85875. Para.

Illustrations of the Aiiatonnj and of Shells 485 10 n 9^9 ^4 15 12 13 16 »»a. PLATE 124 .

Large variety with closely coiled whorls.Z. del Bili. Drepanotrema Incidum 376088. (Pfeiffer).. 8. Bahamas. 107179.N. M. U. Drepanotrema melleum (Lutz). 9. (Pfeiffer). Rio de Janeiro. 423966.. 402809. Drepanotrema lucidum shells.. 2.N. Haiti. Lagumi Louis. Cuba. Half mile NE of Orange Creek. {^^ Pkntorbis redfieldi C. U.Z.M.7. Alacranes Mts.M. U. Drepanotrema lucidum (Pfeiffer). 10. 18-21..N. Drepanotrema lucidum (Pfeiffer).M.6. 535918.) Lea Collection. Immature IT . Jamaica.C. M. 11-17. 129991. St. Brazil. Jamaica. Croix. U. St. Drepanotrcma lucidum (Pfeiffer). B.M..S. Dejit.. du Sud.N. Cat Island.S.X..486 The Molluscan Family Plauorbidae PLATE 125 Shells of Planorbidae 1. Possibly part of original lot of redfieldi. St. M.S. Elizabeth. Drepanotrema lucidum (Pfeiffer).3. All figures are enlarged about four diameters .4. Maggoty. 94757..S. 5. Jealousy Stream.S. Adams.C.

Illustrations of the Anatomy and of Shells 487 ^1 .

N. Tj-pe Isabela. Puerto Rico. 20705. Paratypes. Isabela. C. F.). Near Maldonado.S. All figures are enlarged about four diameters . Drepanotrcma castcuuonitcns locality. Drepanotrema hojfmanni F. Lower CaliU. Argentina.).M. M. Lima. C. B3953.. Age 13.. B3952. C. 362857. Baker. 126 Planorbidae and Van. locality.S.M.488 The MoUuscan Family Planorhidae PLATE Sliells of 1-5. Drepanotrema ciiltratum anitense (Cooper). U. fornia.N.S. 6. 7-9. Peru. (Pils. 151668.. F. Baker.). Figure 14 shows 14-16.S. U. U. Uiuguaj'. Type an immature specimen. Puerto Haberton. 129292. Puerto Rico. 420573. Drepanotrcma hofftnanni Holotype. Puerto Rico. Drepanotrema kermatoide (Orb.N. Santa Rita. Baker.N.N. Drepanotrema hoffmanni variation.M.. Near San Juan. Drepanotrema kermatoide (Orb. LT. 10-12..M. 17-19.S. Laguna.

'''V^iSV^'^ajj^^ V 10 13 11 14 15 17 /' 12 16 mJ 18 19 PLATE 126 .lUustratiois oj the Anatomy and of Shells 489 / \ i ^^^^h^p^" ^tSg^'^^ .

U.).S.S. Antigua. All figures are enlarged about four diameters Brazil.X. . {^ Planorbis macnabianus C. B1258. Drepanotrema cultratum 272282.M. Drepanotrema depressissimum (Moricand). Jamaica. Matanzas.C. U.N.. Paratypes. Guatemala. 4-6. Painters.S. 129995. 94756.S. 17-19. Painters Pond.. Drepanotrema cimcx (Moricand).490 The Molluscan Famih/ Planorbidae PLATE 127 Shells of Planorbidae 1-3. probably part of original lot. Drepanotrema cimex (Moricand). Drepanotrema cimex pistiae 13-15. H. 362133.M. U. i-aihvay near 9-12.. Adams. M. Drepanotrema cimex (Moricand). (Orb. Rincon del Valle. Cuba. Baker.Z. M. U. 7.N. 31432. IQ.8. Pools alono.. Hunts Bay. B.) Lea Collection.C. Drepanotrema cultratum duenafsianum (Tristram). near Caracas. St. Lake Amatitlan. Andrew.N. B.Z..M. Jamaica.. Venezuela. Valle de Yumuri. 378054. M. Bahia.

Illustrations of the Anatotny and of Shells 491 .

Lake Amatitlan. Lower peninsularis California. 129291. are enlarged about four diameters. Locotypes. Tampico. San Luis U. Tampico. Texas. Texas. 26-30. U.M. Guatemala. .S. Trail. B3967.. Tropicorbis obstructns (Morelet). Brownsville. Canal Zone. 11-16. 7-10.N. Mexico.. M. Locotypes. 31-33. figs. M. Pilsbry. Drepanotrona cimex (Moricand).N. 65716. Drepanotrema Anita.2. Mexico. (Cooper).. Guyoz. 6.S. Pilsbry. Drcpanotrema cultratum labrosum Drepanotrema cultratum B1192. Z28550.492 The Molluscan Family Planorhidae PLATE 128 Shells of Planorbidac 1. Potosi Co. B2124. B1183. 31 to 33 .M. Drepanotrema sumichrasti (Crosse and Fischer). Pasture west of San Dieguito. Anapolis. {^Drepanotrema anathuon) Santa U. Brazil.Z. Figures 1 to 30 are enlarged about five diameters. 273933. 23-25. U.C. Tropicorbis obstructns (Morelet).. labrosum Pilsbry. 3-5. B2125. 20-22. Drepanotrema cimex (Moricand). Mexico. Gatun Lake. Drcpanotrema cultratum panuco Drepanotrema cultratum panuco Pilsbry. Brownsville.L. 121018.S.N.. Brazil. Barro 17-19. Barbovu' Lathrop Colorado Island.

Illustrations of the Anatomy and of Shells 493 11 12 m 27 28 i^**^. 22 # 25 ^^ 33 PLATE 128 . 26 13 ^1 14 ^ # 16 10 15 29 30 © 31 17 20 23 18 c 24 il V 19 it.

Tropicorbis orbiculus (Morelet). El Abra. Texas.S.N. Titicaca. Vera Cruz.S.. Tropicorbis orbiculus (Morelet).M.C.N. Mexico. Topo- Agassiz Expedition. C. Catanas Lake. Mexico. New name. Tropicorbis orbicidus dunkcri F. B2113. 32-36. Mexico. 5-7. 16. below Z28528.. 75102.Z. 210881. Edentate. Tropicorbis obstructus (Morelet).C. 8. Tropicorbis orbiculus (Morelet).N .M. Tropicorbis orbiculus dunkcri F. Tropicorbis obstructus donbilli (Tristram). Dry pool near Tampico. Near San Miguel. 210881. Baker.9. Same locality as above. U. Co. 10-15. Cozumel Island. Mexico. U.. {^^ Planorbis licbmanni Dunker). 17-22. 4. 75102. Edentate. Victoria Brownsville.. Quintana Roo.. Chamiai Creek. M. Mexico. Lake types. Valles. Tropicorbis orbiculus (Morelet). All figures are enlarged about two diameters . Los Canoas. B1191.I. U. obstructus donbilli 23-25. 26-31. 37-42. Drepanotrema heteropleurus Pilsbry and Vanatta.Z.494 The Molluscan Family Planorbidae PLATE 129 Shells of Planorbidae 1-3.. C. Mexico. M. Texas. Baker. Same as above.. 464970. Tropicorbis (Tristram). Tropicorbis obstructus (Morelet). B1187.M. Peru. B1184. U. Guadalupe River.S.

* v^^ n ( 18 12 3 7 19 V f) 13 20 37 26 ^(^^. 14 15 38 27 ^^ 28 39 ')> "> # 22 16 21 32 40 29 33 36 23 41 34 30 24 ^^ v5^ 3^ 35 25 PLATE 129 .llhstrations of the Anatomy and of Shells 495 _« /§* 10 17 .

.. All figures are enlarged about two diameters . 59646.C. New Braunfels.C. 33. Yucatan. philippiamis (Dunker).C. Cuba. A small \-ariety. Tropicorbis philippianm (Dunker). (Pfciffer).Z. 26477. Tropicorbis havaiicnsis maya (Morelet). Tropicorbis havanensis (Pfeiffer). 84038. 13-17. gracilentus 4-12. 27-28.M. 13622.M. Tropicorbis gracilcntus (Gould). 84039. M.Z. 35-37. Rio Grande do Norte. Tropicorbis havancnsis (Pfeiffer). Type locality.C. Asuncion.34. Binney's type. Merida.Z.N. p. Type locality. fig. 62490. San Marcos.S. Tropicorbis })crc(jrinus (Orbigny). Ceara Miriin. Tropicorbis coiled. Colorado Desert. 64963. Tropicorbis centimeimlis (Lutz). Figure 31. M.C.Z. Texas.Z.. 75376. Tropicorbis (Gould). Havana.. U.Z. M. 183.. 108. Brazil. Tropicorbis hovancnsis (PfeifTer). U. in drift adjoining fish hatchery. M. Holo- U. Texas. New Orleans. 21-25. B3954. Brazil. 29-32. Tropicorbis 18-20.M. B3298. Louisiana. 92825.. Santa Clara Prov. 38-41.. M. Guayos..496 Tiie Molluscan Family Planorbidae PLATE 130 Shells of Planorbidae 1-3.C. Rio Grande de Sul. Cuba. Argentina... loosely M. Espcranza dc Santa Fe. 26. Paraguay. Figure 10 shows closely coiled whorls like those of Tropicorbis orbiculus.

Illustratious of the Anatomy and of Shells 497 4. 6 9 10 11 14 15 16 17 j/- — 19 21 22 18 20 (^ 2i 5 25 26 24 27 28 32 29 30 31 36 IZ 34 35 'v 7 37 41 G* 38 40 PLATE 130 ..

Valparaiso. Tropicorbis obstructus donbilli Dr\' pool in Tampico. montanus (Orbigny). 74655. B127. Panama. Adams). four diameters.Z.I. U. (Pilsbry). 36702.S.. 32-35. Tropicorbis decipiens (C.Z. 5-7.N.498 The Molluscan Family Planorbidae PLATE 131 Shells of Planorbidae 1^. Panama City. 11. Tropicorbis isthmicus (Pilsbiy).C.26..stram). U. Rockport.C. Tropicorbis isthmicus Type locality. U. Z40460.L. 36 to 40. Puerto Rico. 88181. Texas. Tropicorbis decipiens (C.. B. B. figs. M. 12-14. (Tri. Miraflores Lake. B3966. 36-40. 88180. Tanja (Oroya). Panama. Peru. 162827. 27-31. Taphius subpronus (von Martens). LT. U. Tropicorbis chilensis (Clessin). Puoa.M. Z40575.M. Costa Rica. Mexico.M. 8-10. . 25.. Peru.. Taphius andecolus (Orbigny). M. Taphius andecolus Brownsville.. Adams). Figures 1 to 35 are enlarged about two diameters. Terrialba. B1189. Tropicorbis obstructus donbilli (Tristram). Jamaica. 15-24. Chile.

lUustratiois of the Anatomy and of Shells 499 9t "^ '^>- %P 10 /^ 11 12 13 14 15 K) 17 18 #1 19 • # ^^ 24 28 20 21 22 il 25 26 30 31 27 ) 29 32 ic 34 33 35 37 39 fgp' 36 38 40 131 PLATE .

89990. B. . 22.5. 33-35.M. Panama Canal.Z. M. Tropicorbis riisei {'Dunker) Lares.30.14. kuhnianus (Dunker). Bejuma. Adams). 4534. Nicaragua.23. 74654. U.Z. Venezuela. Venezuela. 6-9. U. U. Venezuela. {^stramineus). 15-21. Tropicorbis fieldi (Tryon). Tropicorbis stramineus (Dunker).I. Type locality.N. Z40466. Gatun.. Tropicorbis stramineus (Dunker).. Tropicorbis fieldi (Tryon).C.I.S. Tropicorbis Adams). Z40463. 29. Merida. Near Granada... 21187. M. Tropicorbis ineridaensis (Preston). Jamaica. Mexico. Tropicorbis gundlachi (Dunker) U. 89991. 31.. 24-28. Tropicorbis bourcardianus (Preston).C. Tucaras... King. U. 65720. Tropicorbis pallidus (C. 13. 4..ston. 202523. Tropicorbis paUidus (C. Paratypes. B3955. 10-12.M.Z.32.C. Ceara. Venezuela. B.Z. Municipio de Aquraz. All figures are enlarged about three diameters . M. Z40461. Ex Dunker. Lagoa Giboia... M. Brazil. Trinidad.M. Puerto Rico.500 The Molluscan Family Planorbidae PLATE 132 Shells of Planorbidae 1-3.C. VI.

Illustrations of the Anatomy and of Shells 501 .

U. Tropicorbis dcclivis (Tate). Uruguay.M. Valles. Tropicorbis nigrilabris (Lutz).Z. Tropicorbis Z40465. Brazil... U. 34. Falls of the Valles River. 18-21.Z. Pool at Piste. 10-13.. 4-9...S.M.N. 22. Antigua. Montevideo. Tropicorbis petenensis Morelet. Tropicorbis Bahia. Tropicorbis heloicus (Orbigny). Tehaimtepec. U. 25-27. schrammi (Crosse). 102211.. Martinique. 152677. Rio de Janeiro.502 The Molluscan Family Planorhidae PLATE 133 Shells of Planorhidae 1-3. 28-33. M.M. U. schrammi (Crosse). B2061. U.Z. 535720. Guatemala. B2117. locality.S. Reservoir north of Guatemala City.M. Tepic.N.S. Tropicorbis obstructus anodontus (Pilsbry).N.C. Brazil.24.C. 59754. Tropicorbis tcpiceiisis (Martens). Tropicorbis janeirensis (Clessin). 24006. 23. All figures are enlarged about three diameters .S. U. M. Mexico. Nicaragua. Type 535713. Yucatan.. Type locality. Tropicorbis obstructus (Morelet).M. 14-17. 36-40. 64995. Guadalajara.C.. In small creek back of the Cerro. 113883.35.N. M.I.. Mexico.S.N. Mexico. Tropicorbis obstructiis (Morelet). Type locality.

5 14 22 32 % 33 6.Illustrations of the Anatomy and of Shells 503 18 28 19 29 20 30 12 4 13 21 31 >. 16 23 • 37 36 ^ 17 25 J^ fti 26 m 35 34 27 133 • 39 ^ 40 38 PLATE .

Baker). Mexico. 14. Tropicorbis janeirensis Tamosopo. Tropicorbis obstruents (Morelet). 12.. 23. Tropicorbis albicans (Pfr.9. Mangrove Cay. 26. U.M..).L.11. 65838. Anapolis. Nicaragua. Tropicorbis paparyc7isis (F.N. U. Tropicorbis 27. Tropicorbis dentiferus (C. Mexico. Tropicorbis obstructus donbilli 24. Brazil. 59754. Tropicorbis albicans (Pfr. Jamaica. Z40465.C. U. B. Brazil.. Catamas Lake.. (Clessin). M.M. six diameters.. C.N. U. Z28545.S. paparyensis (F. St. Ometope.Z.L. Mexico. M. 65717. Pistae. Baker.Z.. Paratypes. 152677.N. along Kings Road. Baker). Baker).C. Mangrove Cay. . 270084. Tropicorbis shitneki F. Tropicorbis 29.S. I^. 20.S.). 534290.S.Z. Z40459. 65717.N. 270084. (Tristram).Z. U. 7. U. U.N..504 The Molluscan Family Planorbidae PLATE 134 Shells of Planorbidae 1-3. Tropicorbis nigrilabris (Lutz)..L. U. Baker. Nicaragua..M. M. Bahamas. schrammi (Crosse) . 65717. Mexico. Z40464.C. Baker. Yucatan. 21. C. 10.). M. Lisbon Andros Island. Goyaz.S. Antigua. C.M. 219696. U. U. Brazil.S. Figures 22 to 29 illustrate lamellae on parietal wall Figures to 11 are enlarged about four diameters. Tamosopo. Bahia.N . Figures 18 to 21 illustrate form of lamellae in aperture 22. Tropicorbis shimeki F. Z40459. M. 15-18.I.N. Tropicorbis schrammi (Crosse). Brazil. Holotype. El Abra.M. 8.. Ometope. 13. Andros. U. Tropicorbis obstructus donbilli (Tristram). 4-6. Adams). 25. Cuba. Catherine. Bog Walk. 534290.. Anapolis.C. B1218. 19. Coatzocoalcos. Tropicorbis paparyensis (F. 1 12 to 29. Tropicorbis obstructus (Morelet). shimeki F. Z28545. U. M.. Cuba.I. Anapolis. Tropicorbis albicans (Pfr.. figs.Z. Martinique.. Tropicorbis 28.S. Point..C. Tropicorbis albicans (Pfr. Brazil.M. Bahamas.). 535713.I.

Illustrations of the Anatomy and of Shells 505 12 V. 5 13 (^ i 14 ii'Ty^f^j 10 15 24 ? 25 11 16 V IS 26 19 17 27 20 22 28 21 ^ PLATE 134 29 .

.N. 24-27. Tropicorbis meridaensis (Preston). Venezuela. Peru.. Soacha.M. U. Tropicorbis peregrinus (Orbigny). 9-11..Z. Tropicorbis chilensis (Clessin).N.S.N. U.E. 380999.M. Peru. U. 17-19.Z. 219755. Ipaguaza. M... All figures are enlarged about 2i/l> diameters .C..M.S. near Caracas.N. U.S. Lea Coll.. 64967. Colombia. 22.S.506 The Molluscan Family Planorhidae PLATE 135 Shells of Planorbidae 1-3... Mexico. M.W.. Small lake near Titicaca. Ibarra. M. Arroyo de Solon. Tropicorbis canonicus (Cousin). Tropicorbis gracilentus (Gould). Tropicorbis centimetralis (Lutz). From Lutz. 20.N. M. U. Tropicorbis bourcardiai^us (Preston). philippianus (Dunker).Z. Coma de 14-16.13.21. L'.S. 271591. of Yacuiba). Montevideo.N.000 feet elevation). of Santiago. Ecuador. 30 miles S.S. 'La Picata' near Bogata. Ruicon de Valle.C. Tropicorbis canonicus (Cousin). 28384.23. Bolivia 126412. M. from Paris Museum. near Bogata. 12. U. Uruguay. Tropicorbis canonicus (Cousin). 5-8.N.M.C. Mexico. U. 341049. Uruguay. 32-35.S. Tropicorbis (15 mi.M. Colombia. Manzanello. Tropicorbis trigyrus (Philippi).M. M. Lake near N. 28-31. U. 535952. 4. (14.S. Tropicorbis peregrinus (Orbigny). 535956.N.S.N. 362132.

.^ 28 13 29 24 20 ^k 25 21 17 30 IP) 18 22 26 r 27 31 19 23 34 32 33 35 135 PLATE .Illustrations oj the Anatomy and of Shells 507 14 10 15 11 ^^r i 16 12 ^^^^ f.

M. Gyraulus deflectus (Say). Gyraulus deflectus (Say). Gyrmdus hirsidus (Gould). Baker borealis Coll. 180292. Miller Bay. Shumigins Zool. Gyraidus hirsutus (Gould). U. M.C. 32-37. Massachusetts.. Charlotte. Immature specimen with Baker Coll. Baker Coll. 11. Lake Maxinkuckee. Lynnfield.. 2768. Baker 127. Group.N.. Indiana. Round Pond. U. Named All specimens are enlarged four diameters . Gyraulus deflectus (Say). Baker Coll.. Lynnfield.. Hirsute and less numerous rows than in albus. Massachusetts. From Mus.S.. Gyraulus deflectus obliciuus (DeKay). Miller Bay.S. 34915.N. Harbor Island. Gyrardms Coll.508 The Molluscan Family Planorbidae PLATE 136 Shells of Planorbidae 1-3. Massachusetts. 15248.. near Scarborough.N.Z. 34915. Wisconsin.C. Gyraidus albus (Miiller). Lynnfield.. 2036.N. Coll. 24. by Westerlund. Base of shell showing 12.. 34915.C. Harbor Island. 426560. 127. Baker coll. 3 shows fine spiral sculpture. Siberia. Popoff Island. shell showing larger 13-15.Z. U.S. near Rochester. 4-7. U.17. Scaly Beck. Lynnfield. Gyraulus deflectus (Say). Gyraulus borealis (We.S. Immature. hairy periostracum. (Say). Lake Maxinkuckee. Lake Winnebago.S.M. 38. epidermis.. Baker 18-20. Russia. New York. No. Chena. Showing hirsute periostracum. Wisconsin. 127. Massachusetts. 2036. England. (Westerhmd). 39. Lake Maxinkuckee. Tanana River. 1120. Gyraulus deflectus (Say). Gyraulus 193289. Gyraulus hirsutus (Gould). Gyraulus hirsutus (Gould). M.Z.. Indiana.M. New York. 16. L^S.N. Saigan.26. Lake George. Gyraulus deflectus 426560.sterlund). Lake George. 8-10. Okoboji Lake... rounded whorls.M. Alaska. Alaska. 27-31. 3. Iowa. Specimen with very acute periphery and hairy periostracum. Gyraulms borealis (Westerlund). fig.. 525319. Gyraulus deflectus obliquus (DeKay). (Say). Showing hirsute 23.. 22.M. New York.M. Lake Winnebago. 21. Indiana.M. U. deflectus 25.X.

Illustrations of the Aimtomy and of Shells 509 13 24 25 14 26 15 27 16 # 12 21 :# 10 28 17 J 18 ^ 29 30 19 22 31 20 11 23 34 m 32 33 S^ 35 # 37 36 38 39 PLATE 136 .

Z.17. Lake Valencia.S. de Cobrera. Australorbis guadcloupensis (Sowb.X. immunis (Lutz).. 271598.C.C. U. L'.S. Brazil..S. 111649. From Liitz. Guadeloupe. Peru. 28. L'. M.15. (Fossil. Taphius montanus (Orb.30. Venezuela.. B3382. U. 25-27. Taphius pronus (Martens). 29.M. Brazil. 74578. Lake Titicaca. Australorbis inimunis (Lutz). 16.36.22-24:. 74578. Duli.M. Briies. 151238.).. 18. Australorbis immunis (Lutz). Peru.. Taphius pronus (Martens). Taphius proiius (Martens). .).M. M. 535957.C. Taphhis andecolus (Orb.X.S. Salle Coll. 110 ft. Australorbis 31-34. Rio de Janeiro. Lake Titicaca..X. Venezuela. 14.X. M. Immature. C.). 251596. (From Lutz) Rio de Janeiro..M.). U. Brazil.X. 19-21. 35.510 The Molluscan Family Plauorbidae PLATE 137 Shells of Planorbidae 1-13. 535953..S.M. Rio de Janeiro.Z. Duli.Z. Penninsula Lake Valencia.

Illustrations of the Anatomy and of Shells 511 ^W 6 .^>> 11 9 J^ .

5-7. Australorbis lugubris (Wagner).Z. U. {nigricans. 336132. Brazil. San Domingo. Montevideo.S... 74586. 26. (Wagner).N. .C. M.M. 21-23. Victoria.Z.S. 121006. U.L..N. ITruguay.Z. Near Caracas. 13-15. M.C. Brazil. Bahia. 126406. Puerto Rico. 17. Australorbis bahiensis (Dunker).Z. Venezuela. 426247. Brazil. 535955.S.. Australorbis lugubris (Wagner). U. M. Lea Coll. Z40479.M. 74584.19. Campos.. Australorbis lugubris (Wagner). (Wagner). Australorbis bahiensis (Dunker).N. 74585. Australorbis bahiensis (Dunker). Australorbis bahiensis (Dunker).Z. Brazil. Caracas. Australorbis bahiensis (Dunker).4.12. 74587. U. Isle 11.512 The Molluscan Family Plauorbidae PLATE 138 Shells of Planorbidae 1.M. 124592. Brazil. Campos.. 24.Z. 16. from Lutz). Lea Coll. Webb Coll.M. M.. 8.C.N. Australorbis blauneri (Germain). U.27. Australorbis lugubris 10.C. Brazil. 20.. U. Venezuela.25.. Brazil.M. M. Vieques.2.. U. Between Caracas and Puerto Cabello. 74588.9..S..M.18. M. Australorbis lugubris Salle Coll.S. 3.N. Australorbis bahiensis (Dunker). 74588. San Domingo.C.C. Venezuela. Australorbis bahiensis (Dunker).S.N.

Illustrations of the Anatomy and of Shells 513 .

Immature.S. Australorbis tenagophilus (Orb..S. French Guiana. U. 8-10. F. Brazil. 3. Australorbis tenagophilus (Orb.N. San Jose. Kitts. 17.C. 102281. Near Quebradillas. Dept. 74577. 36.C. Australorbis glabratus olivaceus (Spix and Wagner).C..C.Z.N. Dept.30. 74577. Dept.C.). Von Ihering Coll.C. 29. 346818. M.Z.S.Z. Brazil. Jurical. Bequaert Coll. M.16. Australorbis paysanduensis (Marshall)..C. ^M.N. (Paratype).N.S.. Paysandu. Uruguay. M. no number. Uruguay. Austral orbis paysanduensis (Marshall).M.Z. Australorbis antiguensis (Sowb. Young. Laurent du Maroni. Australorbis tenagophilus (Orb. 380694.M.. Uruguay. 13. 32. 12.N. Australorbis glabratus olivaceus (Spix and Wagner)...S.S. 34. U.M. 122309.N. Cotypc. Kitts. U.. Arroyo de Solon. Arroyo Arroyo Jurical.). Australorbis glabratus olivaee us (Spix and Wagner) M.18. 15. Rio Couraguam. 322385.M. Duragno..). (Figured type)..). 380746. Australorbis glabratus olivaceus (Spix and Wagner). (Marshall). 19-25.514 The Molluscan Family Planorbidae PLATE 139 Shells of Planorbidae 1.. Australorbis glabratus christopherensis Pilsbry.14. Australorbis antiguensis (Sowb. St.4.). U.2. 380522. 5.M.. Australorbis glabratus rejulgens (Dunker). 31. Paysandu.M. 104573.Z. U. Australorbis tenagophilus (Orb. M.Z. St.M. Corrientes. Australorbis paysanduensis U. Januaria. 2(5-28. Felippo Coll. Santo Domingo. 380997. 74651. 11. M.).Z. Brazil. .7. West Indies.S.N.35. tenagophilus (Orb. 33.).M. Brazil. Uruguay.. U. M. 83890.Z. San Jose.C. Brazil. 103302. Immature. Uruguay. U.s-^ra/or6i!'s Montevideo.. Uruguay. 6. St. M.).. Pilsbry Coll.N. Puerto Rico. Paysandu. Uruguay.. Januaria. 380693. Australorbis tenagophilus (Orb.S.

Illustrations of the Anatomy and of Shells 515 PLATE 139 .

.N. Guadeloupe Immature.S. U. Baker. Puerto Rico. F. M. U. C. 15-17. LT. Near Maracay.S. 14. M. Christopher. U. Holotype. Florida. . 18. 23.. P6778. Kansas. 64969.S.NM. St. C. C. Menetus alabameiisis avus Coll. Australorbis glabratus (Say). Baker.M.I.. U.8. Franklin Estate.. Australorbis glabratus (Say). 24. Immature.Z. Castres.S.S. Gill Coll. 9. Puerto Rico. Australorbis glabratus (Say). Dall 26..I.C.N.S.M.Z. Pilsbrj*. St.516 The Molluscan Family Planorbidae PLATE 140 Shells of Planorbidae 1-3. Dall Coll. Australorbis glabratus (Say).N. Immature. Henderson Coll. Guadeloupe Bequaert. 22.. Antigua. Australorbis glabratus christopherensis Marie U.. U. 390137. La Juria. Paratypes.C.S. 25. Island. 535961. San Juan. Dall Coll. 13. Menetus kansasensis F.M. U. Menetus kansasensis Kansas. Florida.Z. clay). 121204.I. C. U. Galante.. Menetus kansasensis F. Caloosahatchie River. Menetus dalli F. Aiistralorbis glabratus (Say). 349069. (90 feet deep in clay). 112556. W.S.. H. Island. Australorbis glabratus christopherensis Pilsbry. 27. .S.C. Baker. Menetus Baker. Arugua State. Th.. Australorbis glabratus (Say). 28. Bequaert..S.29.N. Paratype. 19. Tortuoguera.M. 535961. 10-12. H. 454061. U.I. Bequaert.M.M.N.N. Lucia. 5. Venezuela.. H. Brazil. C.Z.6...20. 4.. Pliocene. U. U. Baker. 112556. W. 112556. U.M. Florida. 65218.N. W. Aiistralorbis glabratus olivaceus (Spix and Wagner).. Pliocene. 21. Australorbis glabratus (Say) Young. W. Lea Coll. dalli F. 72552. (Pilsbry).N. (90 feet deep in Meade P6778.C. Caloosahatchie River. M. no number. County.N. Puerto Rico. Paratypes. M.M. Aiistralorbis glabratus olivaceus (Spix and Wagner). Holotype. P6778. Meade County. 7.. Caloosahatchie River. 390136.M.

Illustrations of the Anatomij and of Shells 517 25 28 PLATE 140 .

. Young.s.518 The Molluscan Family Planorhidae PLATE 141 Shells of Planorbidae 1...' (Britain in Callao. Z40593. n. (Cousin). Brazil. 29. Ecuador.N.. Helisoma equatorium (Cousin).30. 20702. Peru. 25. 2-4.L. U. Helisoma equatorium (Cousin). Milagis.I. error). Rio Lope Castro. Parana. 12.. W..N.M. Remate.N.M.S. St. Quito. U. Clarendon. Tropicorbis petencnsis Morelet.P. {= Gyraulus). Type.S.^i Miller {= moricandi Beck. (Beach) Eben.M.. U. Helisoma affine.S. U. U. U. . Ecuador. Helisoma peruvianum (Brod.N.M.. U.S. 65265.M. 20. Hondm'as. Tropicorbis denticns (Morelet). U. M.. Moll. Guatemala.S. 26-28. .N. 365157. Plate 7.s. 1.32. 8-11. Andrew. Planorbis boetzke. Schmidt Coll. L. Becjuacrt. U.22).Z. 19. A.C.N. 123897.. Helisoma equatorium U.S. Helisoma fovcalc (Menke). 2i. U. Planorbis turgidus Jeffr. Blatt. n.M. 13-18. Plate 7. Planorbis pedrinus Miller 35. Belize.M.M. Frankfield.S. 126411.. 347900.) 33. 378040.S. 31. 22. Laguna de Peten. 400632. Brazil. Blatt.). 'Georgia.N.). Jamaica. 5-7. Jamaica. (= Tropicorbis) Moll. 1. 64965.S. (Labelled habiensis).21. Helisoma equatorium (Cousin). Helisoma jovcale (Menke). Z40572.. Helisoma peruvianum (Brod. M. 378049. (Like Clessin's figures).N.N. Lea Coll.S.N.34. Tropicorbis pcdrinus (Miller). Hents Bay. 382775. Peru. Ecuador.

Illustrations of the Anatomy and of Shells 519 PLATE 141 .




230 memorial to Cristonrmigcr 78 crop 9 crystalline style 10 219 Bnthyoniphaln. 48. 154 carrefour 6 Castalia 19 central tooth 11 cephalic artery 13 cercariae 21 cerebral ganglia 14 acknowledgments ix Acrorbis 47.-i 46. n. sp. internal 4 Andrews 25 Anisopsis 48. 61 Australorbis 21. n. Helisoma caribaeum. abdominal ganglia 14 accessory cusps 11 callus 15 Campelonn 29 Carinifex 47. 89 blood flukes 21 blood sinuses 13 Boycott 29 buccal ganglia 14 buccal sac 9 Bulinidae 196 Bulinus 21. n. 49. 121 affinities uncertain 192 Ajroplanorbis 21. 226 Menetus. 29. Helisoma cainpanulotum. 46. var. 48. Hdinonin n7iceps. crassilabris. 219 aorta 13 Armiger 46. synonymy. and names of authorities where these api^ear in descriptions and comparisons would extend the size of the index unreasonably without adding value to the reader. 30. 222 costae 16 Costorbis 48. consequently these names are not repeated in the index. var. n. sp. Similarly. Helisoma anceps. 94 Anisus 9. 62 bibliography' 204 biographical sketch xvii cubense. 27. n. Helisoma. 50. 55 anticostianum. 48. var. distribution. n. A list of the more important synonyms is conveniently available in the text in a chronological list following the heading for each group. sp. 86 Aguayo 22 albumen gland 6 anatomical kej's 46 anatomy. 49. 48. var. plate. Chadwick 29 Choanomphalus ' 49. To avoid confusion of numerals for page. 48. For each scientific name the chief reference to the full taxonomic and morphological treatment is cited in boldface type. 196 dalU. n. 46. Menetus 31 coopcri. Frank Collins biographical contributions xxxv book notices and reviews xxxv xvii scientific jniblications xxv barli^chi. 46. references to these details are not cited in the index. a separate index to the illustrations has been prepared wherein all figures are listed under the respective generic and specific names used in the explanation of the plates. Parapholyx dextral 3 231 227 ejfusa. development 25 diaphragm 7 digestive sj'stem 9 523 . Crabb 7. 90 autofecundation 29 Baker. 230 Colton 29 new columbicnsc. and figure numbers. and lists of valid species are treated in full for each genus and subgenus. n. n. morphological terms.INDEX TO TEXT Since in the text of this volume all anatomical features. Menetus. var. hence these items are likewise omitted. 75 arteries 13 arterioles 13 Atropoarmiger 78 auricle 13 history of 41 45 Clonorchis 21 color of shell 16 coloradocnsis. 222 cusps. var. the inclusion of all taxonomic names. accessory 11 central 11 Biomphalaria 48. 192 circulatorv system 13 Clapp 7 classification.

23. n. migration 40 migratory birds 40 Miller 22 48. n. 47. n. 48. cooperi crassilabris. n. 232 labiatus. 177 male organs 4 mantle collar 3 marginal teeth marshalli. var. n. var. 48. 18. 221 campamdatum dalli. 226 var. n. 47 hermaphrodite duct 5 . n. 220 anceps politum. sp. n. n. to subfamilies 46 kidnev 12 Krull 31 labiatns. var. sp. \ar. 230 cooperi planospirus. n. 49. var. n. 28 Lutz 24 Lymnaea 22 Lj'mnaeidae.story of classification 41 Hoffman 22. 187 miracidium 21 moriihology 3 . 219 anceps idahoense. 224 subcrenatum perdisjunctum. n. 227 caribaeum cubense. 22. n. 220 Indoplanorbis 196 intermediate teeth 11 internal anatomy 4 intestine 9 Intha Fasciola 22 Fasciolopsis 22 47. 114 48. 221 binneyi randolphi.1 524 The Molluscan Family Planorhidae Heterodiscus Hippeutis 47. 223 pilsbryi preblei. 49. n. var. var. for other animals 19 habits 19 foot 3 foreword to Part II 215 kennicotti. length of 20 lines of gi'owth 16 lingual ribbon 11 liver 9 liver fluke 22 locomotion 19 Lowrance 25. 49. 60. 54. 31 Jankowski jaw 10 dilatatus. figured specimens. 231 dilatatus floridensis. 23. n. var. ^•ar. 29 Lateorbis. memorial to Baker xvii Menetus 17. var. 46. var. sp. 48. 28 ganglia 14 genera recognized 48 wrongly referred 202 genus new in this volume Pingielln 109 geographical distribution 38 geological history 37 gizzard 9 groups of uncertain affinities 192 growth lines 16 Gyraulus 17. 232 portlandensis. 49. var. 49. 222 kennicotti. 106 47. 230 campanidatum pleistocenicum. Helisoma. 233 n. 65 haemocyanin 13 haemoglobin 13 haemol^•ml1h 13 Larambergue 5. 23 Holmes 25 hosts 21 eupyrene spermatozoa 7 excitatory oi'gan 8 eyes 3 idahoense. sp. 118 Menetus. 123 key to genera of. subgen. n. 112 69 Faust 21. n. sp. n. 225 Helisomatinae 5. 24. n. 223 keys to groups 46 to genera 46 et seq. repositories of. 226 Mattox Menetus n. ix. subgen. n. sp. 182 color adoensis. var. var. Fossaria 22 Fossulorbis 47. 49. 232 lacunae 13 Lankester 25. n. mesometamorphosis 1 Metagonimus 21 Micromenetus. 22. 123 anceps anticostianum. 12. environmental conditions 17 error in generic reference 202 49. 216 floridcnsis. n. Helisoma anceps. 224 trivolvis marshalli. var. sp. campanidatum rideauense. var. 219 Helisoma 29 trivolvis. 231 dalli. sp. monograph of. 19 Haldcmanina head 3 Helisoma 49. 222 columbiense n. var. n. 11 Helicorbis 22. 100 hi. dispersal of species 39 distribution 37 88 Drcpanotrema 21. Menetus 232 flukes 21 food. 49. 47. n. 16. ecology 17 egg gland 6 embryological development 25 et seq. 225 Martins 24 anceps bartschi. n. 85 lateral teeth 11 length of life 20 life.

Helisoma n. 21. Pseudosuccinea 22 publications of F. P3'loris 9 14 n. n. 14. 221 Polypylis 22. var. var. 50. effusa dalli. 11. 49. 22. 48 Micromenetus 187 new subgeneric name Syrioplonorbis 88 new varieties in this \oIume anticostianutii Helisoma anceps 219 anccps bartschi 219 anceps idahoense 220 anceps politum 221 binneyi randolpki 221 campanulalum dalli 226 Planorbijex 49. receptacukun seminis 6 rectum 9 var. 49. rediae 21 renal organ 12 rejiosit