Malacologia 16264 Inst

HARVARD UNIVERSITY
LIBRARY
OF THE
Museum of Comparative Zoology

VOL 1 1962-1964
/"2
MALACOLOGIA
International Journal of Malacology
Revista Internacional de Malacologia
Journal International de Malacologie
Internationale Malakologische Zeitschrift

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/I
DATES OF PUBLICATION
At least 50 copies of MALACOLOGIA were mailed to subscribers (including a free

copy to the Library of Congress, Washington, D. C.) on the following dates:
Vol. 1, No, 1 November 14, 1962
Vol. 1, No. 2 August 7, 1963
Vol. 1, No. 3 June 1, 1964
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MALACOLOGIA VOL. , 1
CONTENTS
J. B. BURCH
Cytotaxonomic studies of freshwater limpets (Gastropoda: Basommatophora)
I,The European Lake Limpet, Acroloxus lacustris 55
Cytological studies of Planorbidae (Gastropoda: Basommatophora).

I.The African subgenus Bulinus s.s 387
J. B. BURCH and M. PATTERSON

Cytotaxonomic studied of freshwater limpets (Gastropoda: Basommatophora)
II.The New Zealand River limpet, Latia neritoides 313
J. B. BURCH, J. E. WILLIAMS, Y. HISHINUMA and R. NATARAJAN

Chromosomes of some Japanese freshwater snails
(Basommatophora: Branchiopulmonata) 403
T. C. CHENG
The effects of Echinoparyphium larvae on the structure of and
glycogen deposition in the hepatopancreas of Helisoma trivolvis in
parasite larvae 291
A. M. CVANCARA
Clines in three species of Lampsilis (Pelecypoda: Unionidae) 215
H. W. HARRY
A critical catalogue of the nominal genera and species of neotropical
Planorbidae 33
The anatomy of Chilina fluctuosa Gray reexamined, with prolegomena

on the phylogeny of the higher limnic Basommatophora (Gastropoda:
Pulmonata) 355
H. W. HARRY and D. V. ALDRICH

The distress syndrome in Taphius glabratus (Say) as a reaction to
toxic concentrations of inorganic ions 283
W. H. HEARD
Distribution of Sphaeriidae (Pelecypoda) in Michigan, U. S. A 139
L. HUBRICHT
The bidentate species of Ventridens (Stylommatophora:
Zonitidae) 417
J. LEVER, J. C. JAGER and A. WESTERVE LD

A new anaesthetization technique for fresh water snails, tested
on Lymnaea stagnalis 331
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MALACOLOGIA, VOL. 1
A. R. MEAD
A flatworm predator of the Giant African Snail Achatina fúlica
in Hawaii 305
J. K. NEEL and W. R. ALLEN

The mussel fauna of the upper Cumberland Basin before its
impoundment 427
J. J. PARODIZ and A. A. BONETTO

Taxonomy and Zoogeographie relationships of the South American
Naiades (Pelecypoda: Unionacea and Mutelacea) 179
R. POHLO
Ontogenetic changes of form and mode of life in Tresus nuttalli
(Bivalvia: Mactridae) 321
I. STIGLINGH, J. A. VAN EEDEN and P. A. J. RYKE

Contributions to the morphology of Bulinus tropicus
(Gastropoda: Basommatophora: Planorbidae) 73
D. W. TAYLOR and N. F. SOHL

An outline of gastropod classification . 7
D. W. TAYLOR, H. J. WALTER and J. B. BURCH

Freshwater snails of the subgenus Hinkleyia (Lymnaeidae: Stagnicola)
from the western United States 237
H. VAN DER SCHALIE

Mussel distribution in relation to former stream confluence in
northern Michigan, U. S. A 227
H. J. WALTER
Punctation of the embryonic shell of Bulininae (Planorbidae) and
some other Basommatophora and its possible taxonomic-phylogenetic
implications 115
G. L. WARMKE and L. R. ALMODOVAR

Some associations of marine mollusks and algae in Puerto Rico 163
M. YAGER and H. W. HARRY

The uptake of radioactive Zinc, Cadmium and Copper by the
freshwater snail, Taphius glabratus 339
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MALACOLOGIA, VOL. 1
TAXONOMIC AND NOMENCLATURAL CHANGES

(New names, emendations, and new ranks)
PELECYPODA
Mutelacea, Parodiz and Bonetto, 1963, 186
GASTROPODA
(Arranged in systematic sequence after Taylor and Sohl, p 8-13)
Architectonicacea, Taylor and Sohl, 1962, 14

Proposed as new, but antedated by Cox,
1960, Proc. Malac. Soc. London, 33:255.
Cylindrobullacea, Taylor and Sohl, 1962, 17
Acochlidioidea, Taylor and Sohl, 1962, 18
Philinoglossoidea, Taylor and Sohl, 1962, 18
Payettiidae, Taylor and Sohl, 1962, 18
Acroloxacea, Taylor and Sohl, 1962, 18
Ancylacea, Taylor and Sohl, 1962, 18
Planorbacea, Harry, 1962, 34
Taphiinae, Harry, 1962, 34
Drepanotrematinae, Harry, 1962, 38
Lymnaeacea, Taylor and Sohl, 1962, 18
Cionellacea, Taylor and Sohl, 1962, 19
Corillacea, Taylor and Sohl, 1962, 19
Strophocheilacea, Taylor and Sohl, 1962, 19
Juliacea, Taylor and Sohl, 1962, 20
monodon, (yentridens), Hubricht, 1964, 420
pilsbryi, {yentridens), Hubricht, 1964, 418
Gnathodoridoidea, Taylor and Sohl, 1962, 21
Dendronotoidea, Taylor and Sohl, 1962, 21
Arminoidea, Taylor and Sohl, 1962, 21
Euarminoidea, Taylor and Sohl, 1962, 21
EoUdoidea, Taylor and Sohl, 1962, 21
Rhodopoidea, Taylor and Sohl, 1962, 12
Veronicellacea, Taylor and Sohl, 1962, 22
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VOL. 1 NO. 1 OCTOBER 1962
ifiüS. W' "^"^' '
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. J 1962
MALACOLOGIA

ALACOLOGIA
A. GISMANN, General Editor J. B.BURCH, Mmiaging Editor
19, Road 12 Museum of Zoology
Maadi, Egypt The University of Michigan
U.A.R. Ann Arbor, Michigan, U.S.A.
EDITORIAL BOARD CONSEJO EDITORIAL

SCHRIFTLEITUNGSRAT CONSEIL DE RÉDACTION
. H. BARNARD N. A. HOLME W. L. PARAENSE

South African Museum Marine Biological Association Instituto Nacional de
Cape Town of the United Kingdom Endemias Rurais
Republic of South Africa The Laboratory, Citadel Hill Caixa Postal, 2113
Plymouth, Devon, England Belo Horizonte
CR. BOETTGER
Zoologisches Institut der Minas Gérais, '3rasil
Technischen Hochschule G. P. KANAKOFF J. J. PARODIZ
Braunschweig Los Angeles County Museum Carnegie Museum
Braunschweig 900 Exposition Boulevard Pittsburg 13, Pennsylvania,
Pockelsstrasse 10a Los Angeles 7, California, U.S.A. U.S.A.
Germany
A. M. KEEN R. D. PURCHON
A. H. CLARKE, JR. Department of Zoology
National Museum of Canada
Department of Geology
University College of Ghana
Stanford University
Ottawa, Ontario, Canada Legon, Accra, Ghana
Stanford, California, U.S.A.
C. J. DUNCAN
S. G. SEGERSTRALE
Department of Zoology
University of Liverpool
Y. KONDO Zoological Museum of
Liverpool, England
Bernice P. Bishop Museum Helsinki University
Honolulu 17, Hawaii, U.S.A. P.-Rautatiekatu 13
E. FISCHER-PIETTE Helsinki, Finland
Muséum National d'Histoire
N. MACAROVICI
naturelle
Laboratoire de Géologie
J. STUARDO
55 rue de Buff on Departamento de Zoología
Paris ye, France Université "Al. I. Cuza"
Instituto Central de Biología
laçi, Romania
A. FRANC Universidad de Concepción
Muséum National d'Histoire
Cas. 301, Concepción, Chile
naturelle D. F. McMICHAEL
55 rue de Buffon The Australian Museum VAN DER FEEN
W.S.S. - VAN
Paris V^, France College Street BENTHEM JUTTING
Sidney, Australia Zoologisch Museum
K. HATAI Amsterdam, The Netherlands
Institute of Geology and
Paleontology J. E. MORTON CM. YONGE
Faculty of Science Department of Zoology Department of Zoology
Tohoku University The University of Auckland The University
Sendai, Japan Auckland, New Zealand Glasgow, Scotland
MALACOLOGIA was established with the aid of a grant tion, Washington, D.C, U.S.A.
(NSF-G24250) from the National Science Founda-
tion, Washington, D.C, U.S.A.
MALACOLOGIA fue establecida con la ayuda de una
subvención (NSF-G24250) de la National Science
MALACOLOGIA wurde
MALACOLOGIA
unter Beihilfe einer Unterstü-
tzung (NSF-G24250) der National Science Founda-
tion, Washington, D.C, U.S.A., gegründet.
fut avec l'aide d'une subven-

établi
tion (NSF-G24250) de la National Science Founda-
Foundation, Washington, D.C, U.S.A.
.
0.10 (NSF -G24250)
,-
.
MALACOLOGIA, AN INTERNATIONAL JOURNAL OF MALACOLOGY
The founders of MALACOLOGIA hope tions obtained, establishment of a self-

that thisnew journal will fill the needs of supporting journal seemed practical.
those workers in different countries who The funds for the establishment of
have found it difficult to get long articles MALACOLOGIA, and for its maintenance
or monographs published. They also aim during the first years of operation, were
to create an international friendly inter- obtained from a grant made by the Na-
est among all those who favour exchange tional Science Foundation, Washington,
of moUuscan information. A definite need D, C, U. S. A. Since such grants are
for a journal of high quality, that would usually awarded only to bona fide non-
be devoted to the multiple aspects of the profit organizations, the Institute of Mala-
study of mollusks, and would publish cology, which meets the requirements,
promptly long manuscripts has been re- was legally established. The founding
cognized for quite some time in the United group of the Institute of Malacology is
States, especially by those active at the responsible for selecting editors and for
major malacological research institutions making such occasional policy decisions
A small group of malacologists, while as may be necessary for the operation
attending the annual meeting of the Ameri- and publishing of MALACOLOGIA. Pre-
can Malacological Union in June 1961, sent members of the Institute of Mala-
discussed the feasibility of promoting such cology are listed on the inside front cover
a journal on an international basis. It of this issue.
seemed to them that such a journal would The purposes and goals of MALACO-
provide not only an outlet to all those LOGIA may be summarized as follows:
requiring it, but would also promote an 1) To publish promptly contributions too
exchange of ideas between the continents long for most of the present malacologi-
and assist in the speedier integration of cal journals. 2) To maintain scholarly
knowledge by providing multilingual publi- standards and continuity of publication as
cation and abstracting. well as editing by an editorial board.
It seemed prudent to test the extent of The present Editorial Board will be in-
possible support for such a journal. Ac- creased by the addition of a sufficient
cordingly, a questionnaire was submitted number of editors to ensure coverage of
to 192 malacologists and other zoologists major areas of specialization and of all
workingwith mollusks at home and abroad. countries. To make sure of originality
The group to which it was sent is broadly of research and technical competence,
representative of malacology all over the each manuscript will be reviewed by two
world; it is formed of the members of or more editors. 3) To assemble in one
various zoological societies, of scientists publication papers that otherwise might
who are carrying on research, who ac- be scattered in a number of different
tively publish and who are associated with scientific journals, in the hope of has-
institutions granting advanced degrees. tening a fruitful exchange of information
The respond ^ was encouraging: 80% of and ideas within the field of malacology.
the questioruaires were returned; 92% of 4) To promote cooperation and to stimu-
those who replied favoured establishment late research in malacology through the
of a ne- journal as proposed, and many medium of an international journal.
favoured it strongly; 6% were undecided
and 2% opposed the establishment of such Elmer G. Berry Robert Robertson
.ajournai. Many replies went beyond J. B.Burch Allyn G. Smith
merely filling out the questionnaire and Melbourne R.Carriker Norman F. Sohl
contributed thoughtful advice. On the ba- Anne Gismann Dwight W. Taylor
sis of the estimated number of subscrip-
(1)
MALACOLOGIA, EINE INTERNATIONALE MALAKOLOGISCHE ZEITSCHRIFT
Die Gründer von MALACOLOGIA hoffen Der Widerhall war recht ermutigend:
mit dieser neuen Zeitschrift jenen For- 80% der ausgesandten Rundfragen wurden
schern in verschiedenen Ländern von beantwortet; 92% der Antworten waren
Nutzen zu sein, die bei der Veröffent- zustimmend, manche darunter sogar äus-
lichung längerer Abhandlungen oder Mono- serst bejahend; 6% der Befragten waren
graphien auf Schwierigkeiten gestossen unentschieden und 2% waren dagegen.
sind. Gleicherweise bezwecken sie eine Viele der Antworten gingen weit über den
freundliche Anteilnahme in all denjenigen Rahmen des Fragebogens hinaus und
anzuregen denen ein Meinungsaustausch brachten wohldurchdachte Ratschläge. Auf
in Sachen der Weichtierkunde zwischen Grund der voraussichtlichen Abonnenten-
den Nationen wünschenswert erscheint. zahl erschien die Gründung einer auf
Dass in den Vereinigten Staaten von eigenen Füssen stehenden Zeitschrift
Amerika zweifelsohne Nachfrage nach durchaus im Bereich des Möglichen.
einem Qualitätsblatt besteht, welches sich Die erstmaligen Mittel für die Grün-
den mannigfaltigen Gesichtspunkten der dung von MALACOLOGIA, sowie ein Be-
Weichtierforschung widmen und längere triebskostenzuschuss für die ersten Jahre,
Manuskripte raschestens veröffentlichen sind durch eine Schenkung der ''National
würde, wurde besonders von den an den Science Foundation" in Washington, D.C.
wichtigeren malakologischen Forschungs- gedeckt. Da derartige Subventionen im
instituten wirkenden Wissenschaftlern allgemeinen nur bona fide nicht ertrag-
längst erkannt. Bei der alljährlichen bringenden Organisationen gewährt wer-
Zusammenkunft der Amerikanischen Mala- den, wurde vorerst, um den bestehenden
kologischen Union, im Juni 1961, besprach Bedingungen zu entsprechen, ein "Institut
eine kleine Gruppe von Malakologen die für Malakologie" (Institute of Malacology)
Aussichten, auf internationaler Basis eine gesetzesgemäss gegründet. Seine Be-
derartige Zeitschrift zu errichten. Man- gründer haben die gegenwärtige Schrift-
war sich einig, dass eine solche nicht leitung gewählt und treffen allfällige zur
nur denjenigen, die eines Ausdrucksmittels Handhabung und Veröffentlichung von MA-
bedurften, dienlich sein würde, sondern LACOLOGIA erforderlichen Entscheidun-
dass ein Gedankenaustausch zwischen den gen. Die gegenwärtige Mitgliedschaft des
Kontinenten im allgemeinen von Vorteil Institutes ist auf der Innenseite des vor-
wäre und dass, durch Darbietung ver- deren Umschlagsblattes aufgeführt.
schiedensprachiger Veröffenlichungen MALACOLOGIAs Ziele und Bestrebun-
und vor allem Zusammenfassungen in genkönnen nun wie folgt zusammengefasst
mehreren Sprachen, der Integration un- werden; es ist beabsichtigt: 1) Beiträge,
seres Wissens eine fruchtbringende Be- die für die meisten bestehenden mala-
schleunigung erteilt werden würde. kologischen Blätter zulange sind, schnell-
Es schien daher angebracht das Aus- stens zu veröffentlichen. 2) Mittels eines
mass der zu erwartenden Anteilnahme an auf breiter Basis gewählten Redaktion-
einer solchen Zeitschrift zu ergründen. sausschusses das wissenschaftliche Niveau
Fragebögen wurden an 192 in- und aus- der Zeitschrift, die Kontinuität ihrer Ver-
ländische Malakologen oder andere sich öffentlichungwie auch einer angemessenen
mit Weichtieren befassende Zoologen aus- Schriftleitung zu gewährleisten; er soll,
geschickt. Der befragte Kreis, welcher um den verschiedenen Spezialisierungs-
über die Welt hin mehr oder minder für gebieten und den sprachlichen und regiona-
Malakologie repräsentatif ist, besteht teils len Anforderungen gerecht zu werden, der
aus Mitgliedern verschiedener malako- bestehende Redaktions rat durch Aufnahme
logischer Vereine, teils aus rege for- einer genügenden Anzahl von Schriftleitern
schenden und veröffentlichenden Gelehrten, erweitert werden. Um sich der Ursprüng-
sowie aus solchen die an Instituten tätig lichkeit des Forschungsstoffes wie auch
sind welche höhere Diplome erteilen. der technischen Kompetenz der Arbeiten
MALACOLOGIA
zu versichern, wird jedes Manuskript soll auf diesem Gebiete, mittels einer
von zumindest zwei oder auch mehreren internationalen Zeitschrift, die Zusam-
Schriftleitern rezensiert werden. 3) Es menarbeit gefordert und die Forschung
sollen fernerhin in einer einzigen Ver- angeregt werden.
öffentlichung Arbeiten vereinigt werden
die sonst in verschiedenen wissenschaft- Eimer G. Berry Robert Robertson
lichen Zeitschriften zerstreut erscheinen J. B.Burch Allyn G. Smith
würden, wobei die Beschleunigung frucht- Melbourne R. Carriker Norman F. Sohl
barer Wechselbeziehungen im Bereiche Anne Gismann Dwight W. Taylor
der Malakologie zu erhoffen ist. 4) Es
MALACOLOGIA, UN JOURNAL INTERNATIONAL DE MALACOLOGIE
Les fondateurs de MALACOLOGIA es- pour un journal pareil. Un questionnaire

pèrent par ce nouveau journal combler fut donc soumis a 192 malacologues ou
une lacune et être utile à tous ceux qui, autres zoologues s'occupantde mollusques.
dans les divers pays, sont venus à Ren- Le groupe questionné - plus ou moins re-
contre de difficultés dans la publication présentatif en malacologie de parle mon-
de leurs articles longs ou de leurs mono- de - comprend des membres de diverses
graphies. En outre, ils envisagent créer sociétés malacologiques, des savants soit
une entente amicale parmi tous ceux qui engagés en recherches, soit publiant ac-
regardent avec faveur un échange d'in- tivement, ainsi que du personnel associé
formation dans le domaine de la malaco- à des institutions scientifiques conférant
logie. des degrés supérieurs.
Le fait qu'il existe certainement une L'écho que provoqua notre enquête fut
demande pour un journal de haute quali- encourageant: réponse fut faite à 80% des
té,dévoué aux multiples aspects de l'étude questionnaires envoyés; 92% de ces ré-
des mollusques et publiant promptement ponses étaient en faveur d'un nouveau
des manuscripts longs, a été reconnu journal tel que proposé et certaines d'en-
depuis quelque temps déjà aux États Unis tre elles l'étaient fortement; 6% des cor-
d'Amérique, particulièrement parmi le respondants se montrèrent indécis tandis
personnel des principaux centres de re- que 2% étaient opposés à l'idée. Certaines
cherches malacologiques. En Juin 1961, réponses dépassaient même de beaucoup
un petit groupe de malacologues, parti- le cadre du questionnaire et apportèrent
cipant à la réunion annuelle de l'Union des conseils réfléchis. A la base du nom-
Malacologique Américaine, discuta les bre estimé d'abonnements, l'établisse-
possibilités de fonder un journal de ce ment d'un journal subsistant par ses
genre sur une base internationale. Il propres moyens semblait faisable.
leur parut qu'un pareil journal non seule- Les fonds pour établissement premier
ment fournirait un débouché à ceux qui de MALACOLOGIA ainsi que pour les
en auraient besoin, mais servirait aussi frais d'entretien pendant les quelques
à avancer un échange d'idées entre les premières années proviennent d'une do-
continents et, en fournissant une publi- nation de la "National Science Founda-
cation et surtout une résumation multi- tion" a Washington, D. C. Parceque,
linguale, favoriserait en même temps une d'habitude, de telles subventions ne sont
intégration plus rapide de notre savoir. accordées qu'à des organisations de bonne
Il semblait alors opportun de mettre à foinon-commercielles, et pour faire face
l'épreuve l'étendue du support à attendre aux exigences de la situation, "l'Institut
MALACOLOGIA
de Malacologie" (Institute of Malacology) cialisation et aux exigences des diverses
fut d'abord légalement établi. C'est le langues et régions. Afin d'assurer l'ori-
groupe fondateur de cet institut qui est ginalité des recherches faites et de la
en ce moment responsable du choix des compétence technique de l'oeuvre, chaque
décisions nécessaires à l'opération et à manuscript sera passé en revue par aux
la publication de MALACOLOGIA. Les moins deux rédacteurs. 3) D'assembler
membres actuels de l'institut sont cités en une unique publication des traveaux
sur la face intérieure frontale de la cou- qui, autrement, seraient dispersés parmi
verture. un nombre de journeaux scientifiques di-
Les intentions et les buts de MALA- vers dans l'espoir d'accélérer ainsi
COLOGIA peuvent être résumés comme un échange fractueux d'informations et
suit: 1) De publier promptement des con- d'idées sur le terrain de la malacologie.
tributions trop longues pour la plupart 4) De stimuler la cooperation et les re-
des journaux malacologiques contempo- cherches en malacologie par la voie d'un
rains. 2) De maintenir un niveau sci- journal international.
entifique élevé ainsi qu'une continuité et
de publication et de rédaction par un con- Elmer G. Berry Robert Robertson
seil de rédaction, dont le nombre de mem- J. B.Burch Allyn G. Smith
bres sera suffisamment augmenté pour Melbourne R. Carriker Norman F. Sohl
faire justice aux aires majeures de spé- Anne Gismann Dwight W. Taylor
MALACOLOGIA, UNA REVISTA INTERNACIONAL DE MALACOLOGIA
Los fundadores de MALACOLOGIA abri- medio para dar salida a los trabajos de
gan eldeseo de que esta nueva publica- aquellos que los requieren, sino también
ción pueda satisfacer la necesidad de los favorecer un intercambio de ideas entre
investigadores de diferentes países que los continentes y ayudar la integración
encuentran dificultad en dar a conocer más rápida del conocimiento, con la pro-
artículos y monografías de cierta exten- visión de publicaciones y resúmenes mul-
sion. Tiene, ademas, el propósito de tilingues.
crear un amistoso interés internacional Pareció prudente comenzar por probar
entre quienes favorecen al intercambio la posibilidad de soportepara tal revista.
de información sobre moluscos entre los Un cuestionario se sometió a la crítica
países. La necesidad de una revista al- de 192 malacólogos, y otros zoólogos que
tamente calificada, dedicada a los múlti- trabajan sobre moluscos en el país y en
ples aspectos del estudio délos moluscos, el extrangero, un grupo que es amplia-
con publicación inmediata de largos manu- mente representativo de la malacologia
scritos, viénese reconociendo desde hace en todo el mundo, miembros de diversas
tiempo en los Estados Unidos, especial- sociedades zoológicas, científicos que in-
mente por aquellos que actúan en insti- vestigan y publican activamente.
tuciones de investigación malacológica La acogida fue promisoria: 80% do los
mayor. Un pequeño grupo de malacólo- cuestionarios fueron devueltos: 92% re-
gos, asistentes al mitin anual de la Union spondieron favorablemente; 6% indecisos,
Malacológica Americana en Junio de 1961, y solo 2% se opusieron a la publicación
discutieron la practicabilidad de promover de la revista. Muchas respuestas so-
la publicación de una revista sobre base brepasaron el cuestionario, contribuyendo
internacional. Los iniciadores estimaron con sugestiones y consejos. En base al
que tal revista sería, no solamente un estimado número de subscripciones ob-
MALACOLOGIA
tenidas, la revista parecía automática- standards de escuela, continuidad de pub-
mente soportada. licación, y editoriales. El consejo edi-
Los fondos para el establecimiento de torial será aumentado con un número su-
MALACOLOGIA, y su mantenimiento du- ficiente de editores para cubrir las areas
rante los tres primeros años, se obtu- mayores de especialisación en todos los
vieron por subsidio de la National Science países. A fin de asegurar la originalidad
Foundation, de Washington, D.C., Estados de investigación y competencia técnica,
Unidos. Desde que estos subsidios son cada manuscrito sera revisado por dos o
generalmente concedidos a organizaciones más editores. 3) Agrupar en una publi-
no gananciales de bueno fe, el Instituto de cación, travajos que de otra mènera pue-
Malacología, que legalmente llena esos den dispersarse en numerosas y diferentes
requerimientos, fué establecido. El grupo revistas científicas, con el deseo de ap-
fundador del Instituto se responsabiliza resurar un intercambio fructífero de in-
por la selección de editores, y sobre la formación e ideas dentro del campo inala-
decisión sobre las reglas para la opera- cológico. 4) Promover la cooperación y
ción y publicación de MALACOLOGIA, La estimular estudios en malacología, medi-
lista de miembros del Instituto va en la ante un órgano internacional.
cubierta frontal interna de este ejemplar.
Les propósitos de MALACOLOGIA son Elmer G. Berry Robert Robertson
los siguientes: 1) Publicar con prontitud J. .
Burch AUyn G Smith
.
artículos y monografías que resultan muy Melbourne R. Carriker Norman F. Sohl

extensos para la mayoría de las revistas Anne Gismann Dwight W. Taylor
actualmente en publicación. 2) Mantener
*
MALACOLOGIA, 1962, 1(1): 7-32
AN OUTLINE OF GASTROPOD CLASSIFICATION^'^

D. W. Taylor and N. F. Sohl
. S. Geological Survey, Washington, D.
ABSTRACT
This outline is a compilation of familial and super- familial classification,
primarily from the "Handbuch der Paläozoologie" (Wenz and Zilch) and the
"Treatise on Invertebrate Paleontology" (Knight and others). These summaries
have been supplemented, especially in the shell-less groups, to make the classi-
fication as nearly consistent as practicable. Numbers of genera and subgenera
are listed for each family, but genera described after publication of the principal
sources have been included only rarely, so that the relative size of the groups
is indicated only in a general way. Annotations include references to the more
important taxonomic groups and explanations of the ways in which divergent
classifications have been reconciled.
Gastropods (7324 genera and subgenera) are divided into the two subclasses
Streptoneura and Euthyneura. Although the Streptoneura (4218 genera and sub-
genera) are larger, they are divided into only 3 orders. The Euthjoieura (3106
genera and subgenera) are more diverse structurally and are divided into 14 orders.
Several recent publications have sum- are taken from the sources mentioned
marized the classification of large parts above, with modifications as indicated.
of the class Gastropoda. The Treatise Genera described since these works were
on Invertebrate Paleontology (Knight and published have rarely been included. This
others, 1960) covers living and fossil classification is therefore out of date to
Archaeogastropoda and other Paleozoic varying degrees, and includes many of
gastropods. Zilch (1959-60) has dealt with the weaknesses of the general works
living and fossil shelled Euthyneura. Wenz quoted.
(1938-44) is the most recent comprehen- In some cases, especially among the
sive source on living and fossil post- Neogastropoda, there has been rather uni-
Paleozoic mesogastropods and neogastro- form disagreement as to Wenz' s familial
pods. classification. In as much as no recent
We have compiled an outline of gastro- monographic treatment exists for these
pod classification from these sources and groups we have retained Wenz' s classifi-
supplemented it by the works of others as cation but have listed the common alter-
indicated in the notes. This classification natives in parentheses.
extends only to the family level. We have The relative size of the orders based
been conservative in recognizing families on numbers of their genera and subgenera
and superfamilies proposed in sources is shown in figures 1 and 2 Our classi-
.
other than these basic works. Other gen- fication, including both living and fossil
eral and recent works which we have con- groups, is the basis of figure 1. Thiele' s
sidered are those by Korobkov (1955), (1929-35) classification, including living
Pchelintsev and Korobkov (1960), and forms only, is the basis of figure 2 . The
Termier and Termier (1952). only obvious difference between the graphs
In the following outline of classification is that in figure 2 the Stylommatophora
the number of genera and subgenera is have increased, primarily at the expense
listed for each family. These numbers of the Archaeogastropoda.
^Publication authorized by the Director, U. S. Geological Survey

^Separates of this paper may be obtained from the Managing Editor of MALACOLOGIA at cost
price.
(7)
TAYLOR AND SOHL
FIG. 1. Relative size of subclasses and orders FIG. 2. Relative size of subclasses and orders
of gastropods, recent and fossil. Arcs are proof recent gastropods. Arcs are proportional to
portional to numbers of genera given in this numbers of genera in Thiele' s (1929-35) classi-
paper. 4, Parasita and Entomotaeniata. 5, fication. Numbers as in Fig. 1.
Cephalaspidea, Acochlidioidea, and Philinoglos-
soidea, 6, Thecosomata. 9, Sacoglossa. 10,
ber of genera and subgenera for each
Anaspidea. 11, Gymnosomata. 12, Notaspidea.
14, Soleolifera, family. The right-hand column indicates
totals for categories above family.
In general we believe that the families
and superfamilies throughout the gastro- Class GASTROPODA 7324
pods correspond to about the same degree Subclass STREPTONEURA 4218
of morphological difference, with the ex-
1. Order Archaeogastropoda
ception of the Archaeogastropoda, where
many of the families and superfamilies
have relatively few genera. This order
has been more finely divided than others.
In accepting the subdivision of the gas-
tropods into Streptoneura and Euthyneura,
we have abandoned the familiar three-
fold division into Prosobranchia, Opistho-
branchia, and Pulmonata. We have fol-
lowed Boettger's(1955) andZilch's (1959-
60) fusion of opisthobranchs and pul-
monates into the Euthyneura of Spengel.
Streptoneura is preferable to Prosobran-
chia for the remaining gastropods because
of the similar derivation of the name.
CLASSIFICATION
Asterisks denote groups that are known

only as fossils.
The left-hand column indicates the num-
OUTLINE OF GASTROPOD CLASSIFICATION
*Euomphalopteridae 3 *Plethospiridae 8
*Portlockiellidae 4 *Superfamily Clisospiracea
*Catantostomatidae 1 *Clisospiridae 5
*Porcelliidae 3 *Superfamily Pseudophoracea 15
*Rhaphischismatidae 1 *Planitrochidae 6
*Phyinatopleuridae 10 *Pseudophoridae 9
*Polytremariidae 2 * Superfamily Craspedosto-
*Laubellidae 1 m atacea 16
*Schizogoniidae 2 *Craspedostomatidae 8
*Zygitidae 1 *Codonocheilidae 6
*Kittlidiscidae 1 *Crossostomatidae 2
*Temnotropidae 1 *Superfamily Palaeotrochacea
Pleurotomariidae 11 *Palaeotrochidae 4
*Trochotomidae 3 *Paraturbinidae 3
Scissurellidae 5 *Superfamily Amberleyacea 28
Haliotidae 11 *Platyacridae 5
*Superfamily Trochonematacea 7 *Cirridae 8
*Trochonematidae 7 *Amberleyidae 10
Superfamily Fissurellacea 56 *Nododelphinulidae 5
Fissurellidae 56 2. Order Mesogastropoda 1969
Superfamily Patellacea 55 Superfamily Cyclophoracea
*Metoptomatidae 3 (note 6)
*Symmetrocapulidae 2 146
Cyclophoridae 61
Acmaeidae 29 Maizaniidae 1
Patellidae 16 Poteriidae
Lepetidae 5 Pupinidae 26
Superfamily Cocculinacea 10 Cochlostomatidae 39
Cocculinidae . 4 Superfamily Viviparacea
Lepetellidae 6 (note 6)
47
*Superfamily Platyceratacea 26 Viviparidae 30
*Holopeidae 13 Ampullariidae (Pilidae) 17
*Platyceratidae 13 Superfamily Valvatacea
* Superfamily Microdom atacea 10 Valvatidae 14
14
*Microdomatidae 5 Superfamily Littorinacea
*Elasmonematldae 158
5 Lacunidae 28
*Superfamily Anomphalacea 9 Littorinidae 30
*Anomphalidae 9 *Purpurinidae 18
*Superfamily Oriostomatacea 9 Pomatiasidae 16
*Oriostomatidae 3 Chondropomidae 66
*Tubinidae 6 Superfamily Rissoacea
Superfamily Trochacea 399 (note 7)
Trochidae 231
407
Hydrobiidae (note 8) 103
*Ataphridae 7 Truncatellidae 16
Stomatellidae 12 Hydrococcidae 1
Turbinidae 91 Stenothyridae 3
Skeneidae (note 3) 46 Bithyniidae (Bulimidae)
Phasianellidae (note 4) 9 (note 9) 16
*Velainellidae 1 Iravadiidae 2
Orbitestellidae 2 Micromelaniidae 47
Superfamily Neritacea 162 Rissoidae (note 9A) 104
*Plagiothyridae 3 Assimineidae (Syncera-
Neritopsidae 17 tidae)(note 8) 37
*Dawsonellidae 1 Aciculidae (note 10) 8
Neritidae 67 Vitrinellidae (Adeorbidae,
Helicinidae 60 Tornidae) (note 11) 55
*Grangerellidae (note 5) 3 Skeneopsidae (note 12) 3
*Deianiridae 1 Omalogyridae (note 12) 3
Phenacolepadidae 5 ? Trachysmidae 1
Hydrocenidae 4 -Pissoellidae (note 12) 4
Titiscaniidae 1 Cingulopsidae (notes 12,
* Superfamily Murchi- 13) 1
soniacea 35 ? Choristidae 2
*Murchisoniidae 27 ? Trochaclisidae 1
10 TAYLOR AND SOHL
*Superfamily Subulitacea
OUTLINE OF GASTROPOD CLASSIFICATION 11
*Superfamily Nerineacea
12 TAYLOR AND SOHL
Superfamily Bulimulacea 223 Pneumodermatidae 4
Bulimulidae 94 Cliopsidae 1
* ? Anadromidae 9 Notobranchaeidae 3
Odontostomidae 16 Clionidae 4
Orthalicidae 17 Thliptodontidae 2
Amphibulimidae (note 43) 8 15. Order Notaspidea (note 50) 18

Urocoptidae 79 Superfamily Umbraculacea 6
Infraorder Aulacopoda 610 Umbraculidae 6

Superfamily Endodontacea 137 Superfamily Pleurobranchacea 12
Endodontidae 107 Pleurobranchidae 12
Otoconchidae 1 16. Order Nudibranchia (note 51) 213
Arionidae 25 Suborder Doridoidea (note 52) 104
Philomycidae (note 46) 3 Infraorder Gnathodoridoidea 2
? Thyrophorellidae 1 Bathydorididae
Superfamily Zonitacea 165 Doridoxidae
Vitrinidae 14 Infraorder Cryptobranchia
Zonitidae 94 Hexabranchidae
Parmacellidae 5 Chrom odor ididae
Milacidae 4 Dorididae 52
Limacidae 26 Halgerdidae
T rigonochlamydidae 7
? Systrophiidae (note 45)
Superfamily Ariophantacea
Trochomorphidae
Euconulidae
Helicarionidae
Ariophantidae
Urocyclidae
Superfamily Testacellacea
Testacellidae
Infraorder Holopoda
Superfamily Polygyracea
(note 47)
? Thysanophoridae
? Ammonitellidae
Polygyridae
Superfamily Oleacinacea
Oleacinidae (note 47)
Sagdidae (note 47)
Superfamily Helicacea
Oreohelicidae (note 47)
Camaenidae (note 47)
Bradybaenidae
Helminthoglyptidae
Helicidae
12. Order Sacoglossa (note 48)
Superfamily Oxynoacea
Arthessidae (note 31)
Oxjrnoidae
Superfamily Elysiacea
Elysiidae
Caliphyllidae
Limapontiidae
Stiligeridae
Oleidae
Superfamily Juliacea
Juliidae
13. Order Anaspidea
Superfamily Aplysiacea
Akeratidae (note 29)
Aplysiidae
14. Order Gymnosomata (note 49)
Laginiopsidae
Anopsiidae
Infraorder Acleioprocta
.
14 TAYLOR AND SOHL

9A. Fretter and Graham (1962, p. 622, dae (Euthyneura)
642) advocated separation of Barleeia The Skeneopsidae, Omalogyridae, Ris-
Clark, 1855, in an independent family. soellidae and Cingulopsidae were listed
It is not certain whether they would in- as Rissoacea incertae sedis by Fretter
clude all of the Rissoidae Barleeinae of and Graham (1962, p. 622-624, 639-640,
Wenz's classification. 642) They show some euthyneuran char-
.
The name Acmeidae used by Wenz

10. acters and perhaps even their ordinal po-
has been changed to Aciculidae in accord sition will be changed.
with Opinion 344 of the International Com- 13. Fretter and Patil (1958) estab-
mission on Zoological Nomenclature. lished the family Cingulopsidae for their
11. Vitrinellidae has been used as the new genus Cingulopsis.
name for this family more often than 14. Pchelintsev (in Pchelintsev and
either Adeorbidae or Tornidae. The scope Korobkov, 1960) has established a super-
of the family is that of Pilsbry and Olsson family Pseudomelaniacea which we ac-
(1945, 1952) and Pilsbry (1953). Many cept with minor modifications. Meeko-
of the genera were included erroneously spira Ulrich and Scofield, 1897 (including
by Wenz in the archaeogastropods as Ske- Cambodgia Mansuy, 1914) and Girtyspira
neidae and Cyclostrematidae. The num- Knight, 1936, of Pchelintsev's Pseudo-
ber of vitrinellid genera is based on melaniidae are included in the Meekospi-
Wenz's Tornidae, on some of his Skenei- ridae of the superfamily Subulitacea fol-
dae and Cyclostrematidae that are not in- lowing Knight and others (1960). Traja-
cluded in Knight and others (1960), and nella Popovici-Hatzeg, 1899, and Paosia
on new genera described by Pilsbry and Bbhm, 1894, which form Pchelintsev's
Olsson. Trajanellidae, have been retained in the
Genera of Wenz's Skeneidae listed here Pseudomelaniidae as in Wenz (1938-44).
as Vitrinellidae are Calceolata Iredale, We accept the Glauconiidae of Pchelintsev
1918; Caporbis Bartsch, 1915, Callom- without modification. Thus the following
phala A. Adams and Angas, 1864; Didia- genera of Wenz' s classification are brought
nema Woodring, 1928; Idioraphe Pilsbry, together: Pseiuloglauconia H. Douville'
1922; Leucodiscus Cossmann, 1918; Leu- 1921, from the Cerithiidae; Glaucoma
corhynchia Crosse, 1867; Megatyloma Giebel, 1852 {inclnáingG y mneyitome Coss-
Cossmann, 1^,
seiidorotella P.Fischer, man, 1909), from the Thiaridae (see also
1857; Rostellorbis Cossmann, 1888; So- note 17); and Pseiuiomesalia H. Douville'
lariorbis Conrad, 1865; Starkeyna Iredale, 1917, from the Vermetidae (see also note
1930; and Teinostoma H. and A. Adams, 16).
1853. Genera from Wenz's Cyclostre- 15 The relationships between Archi-
.
matidae listed here as Vitrinellidae are tectonicidae and Mathildidae have been
Cithna A. Adams, 1863; Eladio rbi s Ire- pointed out by Thiele (1928), who empha-
dale, 1915; and Sczss/Zaöra Bartsch, 1907. sized the heterostrophic protoconch and
Macromphalina Cossmann, 1888, was list- common features of the radula. These
ed erroneously by Wenz as a synonym of families were both classed by Thiele
Megalomphahis Brusina, 1871, in the Fos- among the Cerithiacea. Ovechkin and
saridae. included in the Vitrinellidae
It is Pchelinstev (in Pchelintsev and Korobkov,
after Pilsbry and Olsson (1952). To these 1960) established a superfamily Solari-
we have further added eighteen genera acea for the Solariidae (^ Architectoni-
and subgenera from the papers by Pils- cidae). Modifying the name of this super-
bry and Olsson (1945, 1952) and Pilsbry family to Architectonicacea, we also ten-
(1953). tatively add the Omalaxidae, after Wenz.
12. Fretter (1948) and Fretter and The classification of the Mathildidae in
Graham (1954) have pointed out similari- the Cerithiacea is based entirely upon
ties of Omalogyra Jeffreys, 1860, and the information provided by Thiele (1928).
Rissoella Gray, 1847, to the Pyramidelli- The family is similar to some Archi-
,
tectonicidae in operculum, radula, and dae on the basis of reproductive charac-

heterostrophic protoconch. Risbec (1955, ters. In the present state of knowledge,
p. 70) has observed that the Architectoni- it is not practicable to distribute all the
cidae have more in common with the eu- genera listed by Wenz into these three
thyneuran Pyramidellidae than the Ceri- families. Therefore, only the total num-
thiacea. Both Mathildidae and Archi- ber of genera in the three families is
tectonicidaemay prove to be primitive given.
shelled Euthyneura. 18. Risbec (1955, p. 68-69) advocated
16. Wenz (1938-44) listed 27 genera removal of the Triphoridae from the Ce-
and subgenera of Vermetidae. This as- rithiacea. "One ought to make a special
semblage is now recognized as compo- group intermediate, from certain points
site. We follow Keen (1961) in listing of view, between the Stenoglossa and the
ten genera and subgenera for the family. Mesogastropoda, -- a group closer, in
The seventeen other genera of Wenz' s my opinion, to the Columbellidae than to
Vermetidae are transferred elsewhere the Cerithiidae." [ Translated from the
(Morton, 1951, 1953,- Keen, 1961). Aga- original French] A separate superfamily
.
thirses Montfort, 1808; Anguillospira for the Triphoridae is probably justified,

Cossmann, 1912; Casimiria Cossmann, but we have not established one because
1899; Laxispira Gabb, 1877; Lilax Fin- of uncertainties about where to place it.
lay, 1927; Pro vermicular Kittl, 1899; 19. Seguenzia Jeffreys, 1876, has been
Pseudobrochidium Grupe, 1907; Pyxipoma classified in two ways. Verrill (1884)
Mörch, 18Q0;Siphomum J. E. Gray, 1850; created a separate family in what are
Stephopoma Mörch, 1860; Tenagodus Guet- now called the Mesogastropoda on the
tard, 1770; and Vermiciilaria Lamark, basis of the taenioglossate radula. Wood-
1799, are transferred to theTurritellidae. ring (1928) followed Verrill. Thiele (1925)
But, considering Siphoniiim a synonym of transferred the genus to the Trochidae
Vermiciilaria after Keen (1961), we in- in the Archaeogastropoda and this alloca-
crease Wenz' s Turritellidae by eleven tion has been maintained by Thiele (1929-
genera only. Four of Wenz' s vermetids 35), Wenz (1938-44), and Knight and others
are probably or surely annelids instead (1960). We maintain the Seguenziidae in
of mollusks: Segmentella, Spiroglyphus the Mesogastropoda because no anatomi-
Burtinella, and Cryptobia. Serpulorbis cal data have become available since Ver-
Sassi, 1827, is a valid vermetid genus, rill' s work, because the shell features do
but was listed by Wenz as a synonym of not agree well with other Trochidae, and
Lemintina Risso, 1826, which is based on because we wish to call attention to the
an annelid. Pseudomesalia Douville", 1917, dearth of information about this group.
is transferred to the Glauconiidae follow- We have placed the Seguenziidae in the
ing Pchelintsev (in Pchelintsev andKorob- Cerithiacea without conviction, and follow
kov, 1960). Dihelice Schmidt, 1906, is un- Woodring (1928) in listing it after the
recognizable and has not been counted in Triphoridae. Most recently Clarke (1961)
any family. has maintained the Seguenziidae as a sep-
17. The Thiaridae of Wenz (1938-44) arate family, placing them doubtfully next
have been diminished by transferring to the Trochidae in the Archaeogastropoda.
Glaucoma Giebel, 1852, and Gymnentome 19A. The names Scalacea and Scalidae
Cossmann, 1909, to the Glauconiidae used by Wenz (1938-44) are not based
(Pseudomelaniacea) after Pchelintsev (in upon a valid generic name. Clench and
Pchelintsev and Korobkov, 1960). The Turner (1951, p. 251) have discussed the
rest of the Thiaridae and Wenz' s Lavi- reasons why Epitonium Roding, 1798, is
geriidae (Cyclophoracea) and Anaplocami- the oldest name for the genus. Epito-
dae (Calyptraeacea) have been rearranged niidae and Epitoniacea are thus prefer-
by Morrison (1954) into the three families able names for the family and super-
Thiaridae, Pleuroceridae, and Melanopsi- family.
16 TAYLOR AND SOHL
The family Stenacmidae, included in The Buccinidae of Thiele (1929-1935)
the Amphibolacea by Zilch (1959-60), is and Wenz (1938-1944) have been divided
based upon an epitoniid according to Ro- by Powell (1951) into the three families
bertson and Oyama (1958). Buccinidae, Neptuneidae, and Buccinulidae.
20. Melanellacea and Melanellidae have In the present state of knowledge it is
been rejected in favor of Eulimacea and not practicable to distribute the genera
Eulimidae on the basis of Winckworth's and subgenera listed by Wenz into these
(1934, p. 12) arguments. The Entocon- three families, so the total number only
chidae and Enteroxenidae have been re- is given.
moved from this superfamily to an order 26. The name Melongenidae is pre-
Parasita within the Euthyneura after Ti- ferable to Galeodidae because Galeodes
kasingh and Pratt (1961). Roding 1798 is a junior homonym (Clench
21. Amplocamus Dall, 1895, formed and Turner, 1956).
Wenz's family Anaplocamidae in the Ca- 27. The orders of the Euthyneura are
lyptraeacea. This genus is included in mostly from Zilch (1959-60), listed in
the Pleuroceridae (Cerithiacea) after the sequence of numbers assigned to them
Morrison (1954). by Boettger (1955, p. 263). The ordinal
22. The Xenophoridae have been trans- name Anaspidea is used instead of Ap-
ferred from the Strombacea to the Calyp- lysiacea, and the Notaspidea are ranked
traeacea following Morton (1958a). as a separate order, following Odhner
22A. The Eratoidae are classed in the (1939).
Lamellariacea instead of Cypraeacea fol- 28. The term Entomotaeniata was pro-
lowing Fretter and Graham (1962, p. 626- posed by Cossmann (1896, p. 5) as a sub-
629). order to include the Nerineidae and re-
23. The name Amphiperatidae, used by latives (the mesogastropod Nerineacea of
Wenz, isbased upon a genus which is no- later authors). Subsequently Cossmann
menclatorially invalid under Opinio^ 261 (1921, p. 209-210) concluded that the Py-
Commission on Zoo-
of the International ramidellidae were descended from the ex-
logical Nomenclature. Ovulidae is the tinct Nerineacea, although he did not for-
next most commonly used name. mally include the Pyramidellidae in the
24. Oocorythidae, ranked as a separate Entomotaeniata. Cossmann's summary is
family by Wenz, have been included in so incisive that we quote it:
the Tonnidae following Turner (1948, p.
181).
25. The arrangement of the
tropoda is that of Wenz as modified by
Risbec (1955). The composition of the
Neogas- -
"The origin of the Pyramidellidae
seems evident to me: the Cretaceous
System includes some Ne like shells
{Itieria and especially Itriivia) of which
the columellar plication and heterostro-
suborders Stenoglossa and Toxoglossa is phic protoconch have the greatest analogy
that of Risbec, so that the Mitridae of with those of Pyramidella. Furthermore,
Wenz have been divided. The Mitrinae it seems incontestable that the posterior
and Cylindrinae made up the restricted notch of the lip in the Entomotaeniata,
family Mitridae, which has been trans- exaggerated to the point where it gener-
ferred from the Volutacea in the Steno- ates a subsutural band (Essais V, livr.
glossa to the Toxoglossa as a separate II), could have become reduced in the
superfamily. TheVexillinae have been dis- first pyramidellids to a very weak sinus
sociated and assigned to two different or even to a simply protractive outline
stenoglossan superfamilies: Piisia and of the posterior part of the outer lip, ex-
two included subgenera to the Muricidae; actly as in the opisthobranchs which also
and the fifteen other Vexillinae to the have the protoconch heterostrophic. Con-
Nassariidae. Risbec's name Terebracea sequently the Pyramidellidae are descend-
is rejected in favor of the older and ed from the latter by way of the Ento-
more common term Conacea. motaeniata. But, while the opisthobranchs
.
have persisted to the present with their Knight and others (1960). The Pyrami-
primitive and very ancient characters, dellidae are transferred from the Meso-
their indirect heirs the Pyramidellidae gastropoda to the Euthyneura following
are today very distinct in their anatomi- Fretter and Graham (1949).
cal features, and they resemble each 29. The scope of the Cephalaspidea is
other only by their protoconch and by that of Odhner (1939) and Thiele (1929-
the protractive outline of their lip." 1935) except that the Akeratidae are in-
[Translated from the original French with cluded in the Anaspidea after Boettger
changes to conform with the conventional (1955). The Cephalaspidea of Zilch (1959-
orientation of shells] 1960) are the Cephalaspidea, Acochlidioi-
This inferred phylogeny was doubted dea, and Philinoglossoidea of this classi-
byWenz (1938-1944, p. 64), who thought fication.
the Nerineacea and Pyramidellidae more Odhner (1939) grouped the families of
probably had an earlier, common ances- Cephalaspidea into suborders on the basis
tor. Wenz went on to add: of two characters: degree of development
''One might query, whether the Neri- of shell, and presence or absence of
neacea should not be recognized as a parapodia. Neither Boettger nor Zilch
strongly aberrant side branch of the opis- used formal categories between family
thobranchs, and the Pyramidellacea also and order, but we are establishing super-
brought into closer systematic relation- families based on Boettger' s inferred
ship with them than is usual. Only in- lineages. Boettger's hypothetical lines of
creasing anatomical investigation of the descent were derived largely from degree
pyramidellid genera and of the families of development of six characters: (1) de-
thought to be related can provide a degree of detorsión of the visceral connec-
cision." [Translated from the original tives, (2) shortening of the visceral con-
German]. nectives, (3) change in mutual relationship
An affinity between Pyramidellidae and of individual ganglia and their possible
Nerineacea has been inferred by many fusion, (4) position of the pharyngeal nerve
paleontologists, for example d'Orbigny ring before or behind the pharynx, (5)
(1842-1843, p. 73) and Stoliczka (1867- loss of shell, and (6) loss of operculum.
1868, p. 172). Cossmann's appreciation 30. The Acteonellinae of Zilch (1959-
of the significance of the heterostrophic 60) are ranked as a family after Pche-
protoconch reflects keen insight. Now lintsev (in Pchelintsev and Korobkov,
that the Pyramidellidae have been recog- 1960). In this group we include Cylin-
nized as Euthyneura the inclusion of other dritella White, 1887, and Peruviella 01s-
many-whorled high- spired shells in the son, 1944, after Zilch, but v.e exclude
subclass seems less anomalous than pre- Troostella Wade, 1926, after Pchelintsev.
viously. We approve the grouping of 31. Following Evans (1950) and Morton
Pyramidellacea and Nerineacea in one (1958b) we remove Volvatella Pease, 1860,
order Reviving Cossmann' s Entomotaeni-
. s sa Ev2ins, 1950, from theDiaph-
ata also obviates proposing a new ordinal anidae in the Cephalaspidea and group them
name for the Pyramidellacea as was sug- as a family Arthessidae in theSacoglossa.
gested by Morton (1958, p. 177). 32. Marcus and Marcus (1956) have
Within the Nerineacea the classification established the family Cylindrobullidae
has been modified slightly from that of for Cylindrobulla Fischer, 1857. The
Wenz. The Nerinellidae of Pchelintsev genus shows a mixture of primitive and
(in Pchelintsev and Korobkov, 1960) have specialized characters, but seems most
been accepted without modification. The probably a primitive Cephalaspidean with
family includes 7 genera from Wenz' s affinities to the sacoglossan Arthessidae.
Nerineidae. The Cylindrobullidae do not fit readily into
The Pyramidellacea are composed of any other group of Cephalaspidea and
Pyramidellidae and Streptacididae after we have segregated them as a superfamily
18 TAYLOR AND SOHL
Cylindrobullacea. tention to the uncertain affinities of the
33. Ordinal rank for Acochlidiidae and group, Dall' s Payettinae are ranked as a
relatives is after Odhner (1938, 1939, family and grouped in the same super-
1952) and Marcus (1953). Rank of the family as the Latiidae.
families and numbers of their genera are 38. The scope of the Lymnaeacea of
after Odhner (1952) and Marcus (1953). Zilch (1959-60) has been reduced by sep-
Odhner's name Acochlidiacea is changed arating off two superfamilies, the Ancy-
to Acochlidioidea because we prefer to re- lacea and Acroloxacea, mainly on the ba- -
strict the ending -acea to superfamilies. sis of the classification of Baker (1956)
34. Ordinal rank for the Philinoglos- and the work of Bondesen (1950). The
idae is after Odhner (1952) and Marcus numbers of genera are practically all from
1953). The number of genera is after Zilch. The distinction between Ancylacea
Marcus (1953). The name Philinoglos- and Lymnaeacea was first made by Baker
sacea is changed to Philinoglossoidea be- (1956) who distinguished the superfamilies
cause we prefer to restrict the ending "Ancyloidea" and "Lymnoidea". Baker's
-acea to superfamilies. names have been modified to conform to
35. The superfamilies of Basomma- the customary superfamily endings in mol-
tophora are those of Zilch (1959-60) with lusks. The unique features of the Aero-
an additional one (unnamed) for the La- loxidae have been recognized previously,
tiidae and Chilinidae, which Morton (1955) although the basic differences were not
suggested should be dissociated from the expressed by ranking them formally as a
Lymnaeacea (see also note 37). superfamily.
36. Pseiidorhytidopilus Cox, 1960
The present groupings reflect more
(Knight and others, 1960, p. 237) has been
clearly the relationships of the various
included in the SLphonariidae with simi-
limpet-like freshwater snails to the forms
lar forms placed there by Zilch (1959-
with coiled shells. The patelliform groups
60).
that were originally united in the com-
36A. The family Stenacmidae, included
posite family "Ancylidae" have gradu-
in the Amphibolacea by Zilch (1959-60) is
ally been recognized to fall under four
based upon an Epitoniid (Mesogastropoda)
separate families: the restricted Ancy-
according to Robertson and Oyama(1958).
lidae, Acroloxidae, Lancidae and Latiidae
37. The Payettiidae were established
(note 37) in different superfamilies.
by Dall (1924) as a subfamily Payettinae
for Payettia Dall, 1924. The subfamily The patelliform Acroloxidae (equiva-
was listed under the Planorbidae, but per- lent to the Ancylinae of Walker, 1923)
haps inadvertently. Henderson (1935, p. form a distinct family according to Bon-
267) recorded that Dall "wrote that he desen (1950), who emphasized that they
had intended to place them in the Ancy- have little in common with other Ancyli-
lidae, but that probably Payettia and Latia dae in the broad sense. Not only are
really belong together in a separate they less similar to the Planorbidae in
group.". The ranking of Payettia as a their internal organization than are the
synonym of Amphigyra Pilsbry, 1906, by restricted Ancylidae (see below), but
Zilch (1959-60) was probably based on they are not closely related to other
the works by Wenz (1923, p. 1704) and Basommatophora. "As regards the other
Hannibal (1912). Yen (1944) considered European patelliform freshwater snail,
the genus as one of the Ancylidae. The the dextrorse Acroloxus laciistris (L.),
Payettiidae may be most closely related a study of its egg capsules leaves the
to the Latiidae (listed here under an un- direct impression that we are here con-
named superfamily), or possible to Wal- cerned with a species with an isolated
ker's (1923) Ancylastruminae (included position in the system. The highly de-
in the Ancylidae of the Ancylacea). Part- viating structure of its capsule with the
ly for convenience, and partly to call at- slightly irregular eggs, apparently ar-
ranged without any regular order, and 39. Subdivision of the Stylommato-
the entire absence of any sign of curv- phora into four suborders, and of the
ing or spiral torsion makes the capsule Sigmurethra into three infraorders, is
of this species the most remarkable after Baker (1955, 1962). The family
among the capsules of the freshwater groupings by Zilch (1959-60) have been
pulmonates." (Bondesen, 1950, p. 111- maintained except where distributed in
112). Differences in spermatogenesis different superfamilies by Baker.
also "would tend to further separate 40. The Achatinellacea may possibly
Acroloxus from other Basommatophora" be an unnatural group. Baker (1956)
(Burch, 1961, p. 16). Hubendick (1962) suggested that the Partulidae may belong
has supported the separate family status either near the Ceriidae (in theClausi-
of the Acroloxidae and inferred from the liacea of the Mesurethra), or near the
structure of the genitalia that they are Pupillacea in the Orthurethra. The Acha-
most closely related to the Latiidae. The tinellidae he included in the Pupillacea.
distinctive features pointed out by these Cooke and Kondo (1960) have concluded
authors are here formally recognized by that the Achatinellidae and Tornatellinidae
grouping the Acroloxidae in a separate are most reasonably grouped as one fam-
superfamily. ily Achatinellidae.
Separate family rank for the patelli- 41. Recognition of a superfamily Cio-
form Lancidae, instead of subfamily sta- nellacea for the Cionellidae and Amas-
tus within the Lymnaeidae, is adopted tridae ows Baker (1956). Cochlicopa
following Baker (1925b). Only three valid Ferussac, 1821, is not a senior syno-
genera or subgenera are recognized. nym of Cionella Jeffreys, 1829, or even
Zalophancylus Hannibal, 1912, was based a member of the same family (Kennard,
upon the external mold of a fossil fish 1942, Pilsbry, 1948, p. 1047). The name
vertebra (Hanna, 1925). Cionellidae, used by Pilsbry, is there-
The number of genera of Lymnaeidae fore substituted for Zilch's Cochlicopidae.
is that of Zilch's (1959-60) Lymnaeinae. 42. Baker (1961) ranked the Ceriidae
The Ancylidae as listed here are most and Clausiliidae as separate families
of Walker's (1923) family, after the re- within the same superfamily of the Me-
moval of the following groups: the An- surethra. The Filholiidae are placed in
cylinae Walker (1923) = Acroloxinae
( the Clausiliacea with a query after Zilch
Thiele (1931)), the Protancylinae, Neo- (1959-60). The Megaspiridae are included
planorbinae and Lancinae. These re- after Baker (1961). The Acavidae of Zilch
stricted Ancylidae correspond to the An- have been divided by Baker (1955, 1962)
cylidae, Ferrissiidae, and Rhodacmeidae into Dorcasiidae and Strophocheilidae in
of Zilch (1959-60). the Mesurethra, and Acavidae in the Holo-
The scope of the Planorbidae is basi- podopes in the Sigmurethra. The high-
cally that of Zilch, with addition of his spired, many-whorled shells of Ceriidae,
Neoplanorbidae. Thus from the Ancylidae Clausiliidae, and Megaspiridae have been
ofWalker (1923) both the Neoplanorbinae retained in the superfamily Clausiliacea
and Protancylinae have been transferred which has much the same scope as that
to the Planorbidae. The scope of the of Zilch. Corillacea and Strophocheilacea
Neoplanorbinae is that of Walker, so that have been added as coordinate categories
two genera (instead of only one as in with the Clausiliacea.
Thiele, 1929-35, v. 1, p. 480) have been The name Cerionidae has been emended
added to the Planorbidae. Protancylus to Ceriidae following Baker (1957).
Sarasin, 1898, forming Walker's Protan- 43. Aillya Odhner, 1927, has been re-
cylinae, was transferred to the Planor- moved from the sigmurethran Amphibuli-
bidae by Pilsbry and Bequaert (1927, p. midae and segregated as the heterure-
132). thran family Aillyidae after Baker (1955).
The Physidae have been accepted as in 44. The number of genera of Athora-
Zilch (1959-60). cophoridae is from Solem (1959, p. 44).
20 TAYLOR AND SOHL
45. The Systrophiidae have been in- ly because it cannot be placed readily
cluded with a query in the Zonitacea, fol- anywhere else. Torrechrysias Moreno,
lowing Baker (1956), instead of in a su- 1936, was treated as a synonym of Chry-
perfamily with the Rhytididae and Haplo- sias Pilsbry, 1929, by Zilch (1959-60)
trematidae as classified by Zilch (1959- but is considered valid following Wurtz
60). Polygyratia Gray, 1847, is included (1955).
in the Systrophiidae after Baker (1925a) 48. The subdivision of the Sacoglossa
rather than in the Camaenidae as Zilch and the ranks of these subdivisions are
(1959-60) grouped it. from Kawaguti and Baba (1959). Their
Baker (1956) transferred the three gen- name Tamanovalvidae is rejected because,
era of Austroselenitinae from the Haplo- according to Keen and Smith (1961) and
trematidae to the Streptaxidae with doubt. Baba (1961), Tamanovalva Kawaguti and
46. The number of genera of Philomy- Baba, 1959, is invalid and because the
cidae isfrom Solem (1959, p. 77). older name Juliidae is available. Julia
47. Baker (1956,1962) has divided the Gould, 1862, is known to be a sacoglossan
Oleacinidae of previous classifications (Morrison, 1961) and hence we accept
into two families. The restricted Olea- Thiele's (1929-35) family Juliidae for this
cinidae are included in theholopod super- group. The number of genera in the fam-
family Oleacinacea. The Spiraxidae, with ily is from Keen and Smith (1961). We
subfamilies Spiraxinae, Streptostylinae here also replace Tamanovalvacea Kawa-
and Euglandininae, are included in the in- guti and Baba, 1959, byJuliacea. Berthe-
fraorder Holopodopes and superfamily liniidae Baba, 1961, is a much younger
Achatinacea. Counts of genera are based name than Juliidae. In this classification
on Zilch (1959-60) as follows: Oleacinidae only the one family Juliidae is recognized
are Zilch's " Varicelleae" and Oleacina within its superfamily, and hence the name
R'dding, 1798, in the strict sense. Spirax- Bertheliniacea Baba, 1961, is invalid.
idae are Zilch's Spiraxinae, " Euglan- 49. The number of genera of Gymno-
dineae", and " Streptostyleae", except for somata is from Thiele (1929-35).
Oleacina. 50. The division of the Notaspidea into
The Ammonitellidae were classified Umbraculacea and Pleurobranchacea is
as a subfamily of the Camaenidae by that ofOdhner (1939).
Zilch (1959-60). They were considered 51. Thiele's (1929-35) classification
a distinct family by Wurtz (1955), and of theNudibranchia is much out of date,
thought by Baker (1956) to belong either but there is no comprehensive revision
in the Polygyracea or Endodontacea. of the group available. The outline gi-
Baker
(1956) ranked the Thysanophor- ven here is an attempt to fit Thiele's
idae as a family of either Polygyracea groups into the framework established by
or Endodontacea. The group was included Odhner (1934, 1936, 1939, 1941).
in the Polygyridae with doubt by Zilch Odhner (1939) and following him Boett-
(1959-60). ger (1955) have divided the nudibranchs
The Sagdidae have been transferred into four suborders. To these four, we
from the Polygyracea to Oleacinacea after add a fifth one for the aberrant genus
Baker (1956). Gouostouiopsis Pilsbry, Rhodope, following Thiele (1929-35), Hoff-
1889, was thought to belong to the Sagdi- man (1932-39, p. 193), and Marcus and
dae rather than Camaenidae by Wurtz Marcus (1952). The endings of the names
(1955). have been changed to reserve -acea for
The Oreohelicidae are ranked as a fam- superfamily rank only.
ily, instead of a subfamily of Camaen- Thompson (1961) has recognized two
idae, after Wurtz Solaropsis
(1955). types of larval shells in the Nudibranchia
Beck, 1837, is not one of the restricted and pointed out their significance for
Camaenidae according to Wurtz (1955), classification. Most of the known spe-
but is nevertheless counted in this fami- cies have type 1 (spiral) which occurs
also in the orders Notaspidea and Saco- and Marcus (1957). These are the Dori-
glossa. In the suborders Dendronotoidea didae, Dendrodoridinae and Phyllidiidae
and Eolidoidea both type 1 and type 2 of Thiele.
(egg-shaped) are present. In the classi- 53. Odhner' s classification (1936) of
fication outlined herein, the two types the Dendronotoidea has been accepted
never occur within the same family, and with only minor changes. Tritoniidae
within the Eolidoidea they do not occur has been substituted for Duvauceliidae,
within even the same infraorder. and Tritonia Cuvier, 1803, has been con-
52. Odhner (1934) made a twofold pri- sidered distinct from Duvaucelia Risso,
mary subdivision of the doridoids, into 1826, following Odhner (1939). The works
Gnathodoridacea Eudoridacea, the
and of Odhner (1936) and Thiele (1929-35)
latter being divided again into Crypto- have been combined in counting the gen-
branchia and Phanerobranchia. Marcus era of Tritoniidae.
(1957, 1961) has added Bergh's old group 54. The threefold division of the Ar-
Porostomata as a third primary subdi- minoidea is that of Odhner (1939). In
vision. To preserve the family group- the Euarminoidea, Odhner' s three fami-
ings of present classification without in- lies Heterodorididae, Arminidae, and
troducing more than one category between Doridoididae are equivalent to Thiele' s
suborder and superfamily, we introduce Arminidae and Doridoididae. In the Pa-
a fourfold division of the suborder into chygnatha, the families are those of both
the infraorders Gnathodoridoidea, Crypto- Odhner and Thiele except that the name
branchia, Phanerobranchia, and Poro- Antiopellidae is used following Odhner in
stomata. place of Thiele' s Zephyrinidae. The com-
The Gnathodoridoidea include two fami- position of the Leptognatha is after Odh-
lies, monotypic as in Thiele' s classifi- ner (1939). Goniaeolididae is changed
cation. in conformity with the original spelling
The Cryptobranchia include the doridids of GonieoUs Sars, 1859. This family
and close relatives. Dorididae are split and also Heroidae are used as in Thiele' s
into many families by Pruvot-Fol (1954), classification. The Charcotiidae are ac-
but following Odhner (1934) only Chromo- cepted in the sense of Odhner (1934), so
dorididae, Dorididae, andHalgerdidae are that Charcotia Vayssiere, 1906, and Pseu-
recognized. These three include all of dotritonia Thiele, 1912, are removed from
Thiele' s (1929-35) Dorididae except for Thiele' s Notaeolidiidae. Telarma Odhner,
the Dendrodoridinae. The Hexabranchi- 1934, is the third genus of this family.
dae are monotypic, as in Thiele. 55. Division of the Eolidoidea into
The Phanerobranchia are accepted in three groups is from Odhner (1934, 1939).
the sense ofOdhner (1941), who divided The Pleuroprocta include two families.
them into Suctoria and Nonsuctoria. If The Notaeolidiidae as restricted by Odh-
these groups are ranked as superfamilies ner (1934) include only one genus. The
they should have names based on a typi- second pleuroproct family of Odhner
cal genus, but for present purposes no (1939), Coryphellidae, is included in
new formal names are necessary and Thiele' s Flabellinidae.
Odhner' s terms are listed in parentheses. The Acleioprocta of Odhner (1939) are
The four families of Nonsuctoria are made up of three families, Eubranchidae,
those of Odhner (1941); they form part Cuthonidae, and Calmidae. Three addi-
of Thiele' s Polyceridae. The Onchidori- tional families, Fionidae, Flabellinidae,
didae and Goniodorididae of the Suctoria and Pseudovermidae are recognized by
make up the rest of Thiele' s Polyceridae. Marcus and Marcus (1955), Marcus (1961),
The Corambidae and Vayssiereidae have and Pruvot-Fol (1954). These six fami-
been accepted in the sense of Thiele. lies include the Tergipedidae, Fionidae,
Two families have been segregated in Calmidae, Pseudovermidae, and most of
the Porostomata by Pruvot-Fol (1954) the Flabellinidae of Thiele' s classification.
22 TAYLOR AND SOHL
The cleioproct Eolidoidea are classed BABA, Kikutarô, 1961, The shells and radulae
in Berthelinia, a bivalvedsacoglossan genus.
by Odhner (1934, 1939) in four families,
Venus, 21: 389-401, pi. 21.
which are equivalent to Thiele' s Aeo- BAKER, H. ., 1925a, Agnathomorphous Aula-
lidiidae. The four families recognized copoda. Nautilus, 38: 86-89.
here are those which Marcus (1955, 1957, 1925b. Anatomy of Lanx, a limpet-
,
1958) has modified from Odhner's classi- like lymnaeid mollusk, Proc. Calif. Acad.
Sei., ser. 4, 14: 143-169,
fication. Ranking and numbers of genera
, 1955, Heterurethrous and aula-
of Facelinidae and Favorinidae are after copod. Nautilus, 68: 109-112.
Marcus Odhner's Aeolidiidae and
(1958). , 1956, Family names in Pulmonata.
Spurillidae are combined after Marcus Nautilus, 69: 128-139; 70: 34.
(1955, 1957) into the Aeolidiidae whose 1957, Families of Pulmonata. Nau-
,
tilus, 70(3): 141-142.

number of generals counted from Thiele' s 1961, Puerto Rican pupillids and
,
Aeolidiinae. Glaucinae of Thiele are clausilioids. Nautilus, 75: 33-36.

ranked as a family after Odhner. , 1962, Puerto Rican Holopodopes.
Thirty-one genera of the Coryphellidae, Nautilus, 75: 116-121.
Eubranchidae, and Flabellinidae have not 1962, Puerto Rican oleacinoids.
,
Nautilus, 75: 142-145.

been allocated to a specific family, but BOETTGER, C. R., 1955, Die Systematik der
have been included in the count of genera euthyneuren Schnecken. Zool. Anzeiger,
within the Eolidoidea. Suppl., 18: 253-280.
The monotypic Myrrhinidae of Thiele' s BONDE SEN, Poul, 1950, A comparative mor-
phological-biological analysis of the egg
classification have been listed as Eoli-
capsules of freshwater pulmonate gastro-
doidea incertae sedis. pods, Natura Jutlandica, 3: 1-208, pi. 1-9.
56. Soleolifera was used by Thiele BURCH, J. ., 1962, Some cytological aspects
for the Rathouisiidae and Veronicellidae of Acroloxus lacustris (linnaeus). Amer.
only, but we follow Zilch in applying this Malacol. Union Ann, Rept. 1961, p. 15-16.
,
name to the order. For Soleolifera in CLARKE, A. H., Jr., 1961, Abyssal mollusks
from the south Atlantic Ocean. Bull. Mus.
Thiele' s sense we substitute Veronicel- Compar. Zool., Harvard College, 125: 345-
lacea. The numbers of genera of Onchi- 387, pi. 1-4.
diidae are from Solem (1959, p. 37) and CLENCH, W. J. and R. D. TURNER, 1951, The
of Rathouisiidae and Veronicellidae from genus Epitonium in the western Atlantic,

Thiele (1929-35). Part 1. Johnsonia, 2: 249-288.
:
1956, The family Melongenidae in

,
the western Atlantic. Johnsonia, 3: 161-188.

COOKE, M., Jr., and Yoshio KONDO, 1960,
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ZUS AMME NF ASSUNG

UMRISS EINER KLASSIFIKATION DER GASTROPODEN
Die Systematik grösserer Abschnitte der asse Gastropoda ist in verschiedenen

neueren Veröffentlichungen zusammenfassend behandelt worden. Lebende und fossile
Archäogastropoden und andere paläozoische Gastropoden werden von Knight u. a.
(1960) erfasst, Zilch (1959-60) behandelt umfassend die lebenden sowie fossilen
schalentragenden euthyneuren Schnecken, während Wenz (1938-44) als jüngste um-
fassende Quelle für lebende und fossile post-paläozoische Meso- und Neogastropoden
anzusehen ist.
Unser Umriss einer Einteilung der Gastropoden wurde zwar hauptsächlich aus
den obigen Quellen aufgestellt, wurde jedoch, insbesondere für die schalenlosen
Gruppen, auch aus anderen Arbeiten ergänzt, wie in den Anmerkungen angegeben.
Dieser Umriss* reicht nicht unter Familienrang herab. Nur wenige von anderen als
unseren Hauptquellen vorgeschlagenen Familien oder Superfamilien wurden darin
aufgenommen. Ebenfalls herangezogene neuere und allgemein zusammenfassende
Arbeiten sind die von Korobkov (1955) Ptschelintsev und Korobkov (1960) (angeführt
unter Pchelintsev) und Termier und Termier (1952). In einigen Fällen, besonders
was die Neogastropoden anbelangt, bestehen ziemlich allgemein Meinungsverschie-
denheiten über die von Wenz aufgestellten Familien. Wir haben mangels einer jün-
geren monographischen Bearbeitung, trotzdem seine Einteilungbeibehalten, haben
jedoch die alternativen gebräuchlichen Benennungen in Klammern hinzugefügt.
Erklärungen darüber wie die verschiedenartigen Enteilungen in Einklang gebracht
wurden sind in den Anmerkungen zu finden.
Die Zahl der Gattungen und Untergattungen in der Gastropoden (7 ,324) stammt
mit nur wenigen Abänderungen aus obigen Arbeiten und ist wann immer möglich
für jede der Familien angegeben (linke Spalte der "Classification"). Später be-
schriebene Gattungen wurden nur selten aufgenommen (wie angemerkt). Unsere
Einteilung ist daher zu verschiedenen Graden veraltet und enthält viele der Schwä-
chen der angeführten zusammenfassenden Werke.
Die Gastropoden sind hier in die 2 Unterklassen Streptoneura und Euthyneura
eingeteilt, wobei, nach Boettger (1955) und Zilch (1959-60), die altbekannten Opistho-
branchia und Pulmonata durch Spengels Euthyneura ersetzt werden, während die
Benennung Streptoneura, wegen der parallellen Ableitung des Wortes, den Namen
Prosobranchia vorzugsweise ersetzt. Obwohl die Streptoneura (4,218 Gattungen und
Untergattungen) die grössere der beiden Gruppen darstellen, sind sie in bloss 3
Gruppen eingeteilt, während die Euthyneura (3,106 Gattungen und Untergattungen)
grössere Verschiedenheiten des Aufbaues aufweisen und in 14 Ordnungen zerfallen.
Die Grössenverhältnisse dieser Ordnungen, nach der Zahl ihrer Gattungen und
Untergattungen berechnet, sind in Abbildungen 1 und 2 dargestellt, wobei die erstere
unserer eigenen, sowohl lebende als auch fossile Formen enthaltenden assifikation
entspricht, und die letztere derjenigen von Thiele (1929-35), welche nur lebende
Formen umfasst. Der einzige markante Unterschied zwischen den beiden Darstel-
lungen besteht darin, dass in der letzteren die Stylommatophora, hauptsächlich auf
Kosten der Archeogastropoda, umfangreicher sind.
*Die Sternchen bezeichnen durchwegs fossile Gruppen.

26 TAYLOR AND SOHL
Im allgemeinen scheinen innerhalb der Gastropoden, mit Ausnahme der Ordnung

Archäogastropoda, die Einschnitte in Familien und Superfamilien ungefähr gleich-
wertigen morphologischen Unterschieden zu entsprechen. Die genannte Ordnung
zeigt eine feinere Einteilung, indem viele Familien und Unterfamilien verhältnis-
mässig wenige Gattungen enthalten.
RÉSUMÉ
e'bAUCHE D'UNE CLASSIFICATION DES GASTÉROPODES
La Classification des principaux groupes de la classe des gastéropodes se

trouve résumée dans plusieures publications plus ou moins récentes. Ainsi Knight
et autres (1960) couvrent les archéogastéropodes vivants et fossiles ainsi que les
autres groupes paléozoiques. Récemment, Zilch (1959-60) traite des mollusques
euthyneures pourvus de coquilles, vivants et fossiles, tandis que l'oeuvre de Wenz
(1938-44) représente la plus récente source comprehensive pour les méso- et néo-
gastéropodes post-paléozoiques vivants et fossiles.
Notre classification gastéropode a été esquissée principalement d'après ces
sources, mais elle a aussi été supplémentée par d'autres travaux, surtout en ce
qui concerne les groupes atestes, comme indiqué dans les notes. Cette classiñ-
cation* ne descend qu'au niveau familial. Familles et sous-familles proposées par
d'autres auteurs que ceux déjà cités ne sont inclus que dans une faible mesure. Les
principaux travaux additionnels récents et généraux pris en considération sont ceux
de Korobkov (1955), de Ptchélintsev et Korobkov (1960) (voir Pchélintsev) et de
Termier et Termier (1952). Dans certains cas, spécialement en ce qui concerne
les néogastéropodes, il y a désagrément assez uniforme au sujet de la classification
familiale de Wenz. Nous avons pourtant retenu sa classiñcation faute de traité
monographique plus récent, ajoutant toutefois, entre parenthèses, les désignations
alternatives courantes. De quelle manière les classification divergentes ont été
réconciliées est expliqué dans les notes.
Le nombre de genres et sous-genres gastéropodes (7,324) est derivé princi-
palement des sources indiquées ci-haut est et donné pour chaque famille (colonne
gauche, classification) dans la mesure du possible. Les genres décrits après la
publication de ces travaux ne sont que rarement inclus. Notre classification est
donc périmée à dégre variable et participe des faiblesses de ces ouvrages généraux.
Les gastéropodes sont ici divisés en 2 sous-classes, les streptoneures et les
euthyneures, ces derniers étant formés par la fusion des opisthobranches et des
pulmones, selon Boettger (1955) et Zilch (1959-60), tandis qu'en vertu de sa déri-
vation similaire, le terme streptoneure remplace celui de prosobranche. Quoique
les streptoneures (4,218 genres et sous-genres) forment le plus grand de ces 2
groupes, ils ne sont divisés qu'en 3 ordres, pendant que les euthyneures (3,106
genres et sous-genres), de plus grande diversification structurelle, sont divisés en
14 ordres. Les dimensions relatives de ces ordres, selon le nombre de leurs
genres et sous-genres, sont illustrées par les figures 1 et 2, dont la première se
base sur la présente classification comprenant groupes vivants et fossiles et la
seconde sur celle de Thiele (1929-35) ne comprenant que formes vivantes. La seule
différence frappante entre ces deux illustrations est, dans la seconde, l'accrois-
sement des stylommatophores au dépens des archéogastéropodes.
Les coupures familales et sur- familiales dans les gastéropodes semblent cor-
respondre à peu près au même degré de differentiation morphologique, à l'exception
de l'ordre des archéogastéropodes qui a été plus finement divisé et dont les familles
et surfamilles n'ont que relativement peu de genres.
*Les groupes pourvxu d'un astérisque sont uniquement fossiles.

RESEÑA
UNA RESEÑA DE LA CLASIFICACIÓN DE LOS GASTRÓPODOS
Varias publicaciones recientes han resumido la clasificación de gran parte de
la clase Gastropoda. Knight y otros han estudiado los arqueogastrópodos fósiles y
vivientes, así como otros gastrópodos paleozoicos. Zilch (1959-1960) ha estudiado
los fósiles y vivientes con concha de la subclase Euthyneura. La fuente de información
más reciente acerca de mesogastrópodos y neogastrópodos fósiles y vivientes es
Wenz (1938-1944).
Hemos llevado a cabo una reseña de la clasificación de los gastrópodos basada
principalmente en esta información, suplementándola con información especial para
los grupos con o sin conchilla como indicamos en nuestras notas. Esta clasiñcación*
se extiende únicamente a la familia. Nos hemos mostrado conservadores en lo que
se refiere al reconocimiento de familias y superfamilias propuestas en otras fuentes
de información distintas a las que han servido para estos trabajos básicos. También
hemos considerado otros trabajos generales llevados a cabo recientemente por Korob-
kov (1955), Pchelintsev y Korobkov (1960)y Termiery Termier (1952). La forma en
que estas clasiñcaciones divergentes han sido reconciliadas se explica en nuestras
notas.
En algunos casos, especialmente entre los neogastrópodos, existe una discre-
pancia uniforme en lo que se refiere a la clasificación por familias de Wenz. Debido
a que no existe ningún tratamiento monográfico reciente para estos grupos, nos
hemos limitado a retener la clasificación de Wenz, dando entre paréntesis una lista
de alternativas comunes.
Damos una lista (columna izquierda en la clasificación) del número de géneros y
subgéneros de los gastrópodos (7 ,324) siempre que nos es posible. Estas figuras han
sido tomadas de las fuentes de información mencionadas anteriormente con las
modificaciones indicadas. Los géneros descritos después de la publicación de estos
estudios se incluyen muy raremente, por lo tanto nuestra clasificación está en cierto
modo atrasada incluyendo muchos de los puntos débiles de los trabajos generales ya
mencionados.
En este trabajo, los gastrópodos se dividen en 2 sub-clases, Streptoneura y
Euthyneura, siendo la ultima mencionada formada por los opistobranquios y pul-
monados, dejando el término Steptoneura para los prosobranquios, debido a la deri-
vación similar del nombre. A pesar de que los Streptoneura constituyen el grupo
más grande (4,218 géneros y subgéneros) se dividen únicamente en 3 órdenes, mien-
tras que los Euthyneura (3,106 ge'neros y subgéneros), que son más diversos en su
estructura, se dividen en 14 órdenes. La medida relativa de estos órdenes, basadas
en el mismo número de géneros y subgéneros, se muestra en las figuras 1 y 2.
Nuestra clasificación, incluyendo fósiles y vivientes, constituyen la base de la figura
1. La clasificación de Thiele (1929-1935) incluyendo únicamente las formas vi-
vientes, constituyen la base de la figura 2. La única diferencia entre los dos cuadros
es el hecho de que en la figura 2, los Stylommatophorahan aumentado principalmente
a expensas de los arqueogastrópodos.
En general creemos que las incisiones familiares y superfamiliares a través
de los gastrópodos, corresponden en el mismo grado a la diferencia morfológica,
con la excepción de que los arqueogastrópodos se han dividido más finamente que
otras órdenes, resultando así que muchas de las familias y superfamilias tienen
relativamente muy pocos géneros.
*Taxa marcada con un asterisco se refiere iónicamente a los fósiles.

28
', TAYLOR AND SOHL
,, , : . .
,. . ..
-
,
"Handbuch der Paläozool cgie" (Wenz und Zilch)
.
"Tr-estise on Inverteorate Faleontolcgy" (Knlght and others)
,
. . ,- ,,
:) ).) ,
ura (4218
.
(7324
Streptoneura Euthiyneura. str-eptone-
. 14
huthyneura (3106
INDEX OF SCIENTIFIC NAMES
Abyssochrysidae, 10 Aporrhaidae, 10 Casimiria, 15

Acavidae, 11,19 Aranucidae, 12 Cassididae, 10
Achatinacea, 11,20 Archaeogastropoda, 7,8,13,15 Catantostomatidae, 9
Achatinidae, 11 Architectonic acea, 10,14 Cavolinidae, 11
Achatinellacea, 11,19 Architectonicidae, 10,14,15 Cephalaspidea, 7,11,17
Achatinellidae, 11,19 Arionidae, 12 Ceriidae, 11,19
Aciculidae, 1,14 Ariophantacea, 12 Cerionidae, 19
Acleioprocta, 13,21 Ariophantidae, 12 Ceritellidae, 11
Aclididae, 10 Arminidae, 12,21 Cerithiacea, 10,13,14,15,16
Acmaeidae, 9 Arminoidea, 12,21 Cerithiidae, 10,14,15
Acmeidae, 14 Arthessa, 17 Cerithiopsidae, 10
Acochlidiacea, 18 Arthessidae, 12,17 Charcotia, 21
Acochlidiidae, 11,18 Assimineidae, 9,13 Charcotiidae, 12,21
Acochlidioidea, 8,11,17,18 Asterophilidae, 10 Chilinidae, 11,18
Acroloxacea, 11,18 Ataphridae, 9 Chlamydephoridae, 11
Acroloxidae, 11,18,19 Athoracophoracea, 11 Chondrinidae, 11
Acroloxinae, 19 Athoracophoridae, 11,19 Chondropomidae, 9
Acroloxus, 18,19 Atlantacea, 10 Choristidae, 9
Acroreidae, 11 Atlantidae, 100 Chrom odorididae, 12,21
Acteonacea, 11 Atyidae, 11 Chrysias, 20
Acteonellidae, 11 Aulacopoda, 12 Cingulopsidae, 9,14
Acteonellinae, 17 Austroselenitinae, 20 Cirtgulopsis , 14
Acteonidae, 11 Barleeia, 14 done lia, 19
Adeorbidae, 9,14 Barleeinae, 14 Cionellacea, 11,19
Aeolidiidae, 13,21,22 Basommatophora, 11,18,19 Cionellidae, 11,19
Aeolidiinae, 22 Bathydorididae, 12 Cirridae, 9
Agathirses, 15 Bellerophontacea, 8 Cithna, 14
Aglajidae, 11 Bellerophontidae, 8 Clausiliacea, 11,19
Aillya, 19 Bertheliniacea, 20 Clausiliidae, 11,19
Aillyidae, 11,19 Bertheliniidae, 20 Cleioprocta, 13
Akeratidae, 12,17 Bithyniidae, 9,13 Clionidae, 12
Amaltheidae, 10 Bornellidae, 12 Cliopsidae, 12
Amastridae, 11,19 Brachytremidae, 10 Clisospiracea, 9
Amberleyacea, 9 Bradybaenidae, 12 Clisospiridae, 9
Amberleyidae, 9 Buccinacea, 10 Cocculinacea, 9
Ammonitellidae, 12,20 Buccinidae, 10,16 Cocculinidae, 9
Amphibolacea, 11,15,18 Buccinulidae, 10,16 Cochlicopa, 19
Amphibolidae, 11 Bulimidae, 9,13 Cochlicopidae, 11,19
Amphibulimidae, 12,19 Bulimulacea, 12 Cochostomatidae, 9
Amphigyra, 18 Bulimulidae, 12 Codonochilidae, 9
Amphiperatidae, 10,16 Bullacea, 11 Coelostylinidae, 10
AmpuUariidae, 9,13 BuUidae, 11 Colombellinidae, 10
Anadromidae, 12 Bursidae, 10 Columbellidae, 10,15
Anaplocamidae, 15,16 Burtinella, 15 Conacea, 10,16
Anaplocamus, 16 Caecidae, 10 Conidae, 10
Anaspidea, 8,12,16,17 Calceolata, 14 Coralliophilidae, 10
Ancylacea, 11,18 Caliphyllidae, 12 Corambidae, 12,21
Ancylastrmninae, 18 Callomphala, 14 Coreospiridae, 8
Ancylidae, 11,18,19 Calmidae, 13,21 Corillacea, 11,19
Ancylinae, 19 Calyptraeacea, 10,15,16 Corillidae, 11
Anguillospira, 15 Calyptraeidae, 10 Coryphellidae, 12,21,22
Anomphalacea, 9 Camaenidae, 12,20 Craspedopominae, 13
Anomphalidae, 9 Cambodgia, 14 Craspedostomatacea, 9
Anopsiidae, 12 Cancellariidae, 10 Craspedostomatidae, 9
Anthracopupa, 13 Caporbis, 14 Crossostomatidae, 9
Antiopellidae, 12,21 Capulidae, 10 Cryptobia, 15
Aplysiacea, 12,16 Carbonispira, 13 Cryptobranchia, 12,21
Aplysiidae, 12 Carinariidae, 10 Ctenosculidae, 10
30 TAYLOR AND SOHL
Cuthonidae, 13,21 Euomphalidae, 8,13 Hydatinidae, 11
Cyclophoracea, 9,13,15 Euomphalopteridae, 9 Hydrobiidae, 9,13
Cyclophoridae, 9 ETiStomidae, 10 Hydrocenidae, 9
Cyclostrema, 13 Euthyneura, 7,8,10,14,15,16,17 Hydrococcidae, 9
Cyclostrematidae, 13,14 Facelinidae, 13,22 Idioraphe, 14
Cylindrinae, 16 Fasciolariidae, 10 Iravadiidae, 9
Cylindritella, 17 Favorinidae, 13,22 Itieria, 16
Cylindrobulla, 17 Ferrissiidae, 19 Itieriidae, 11
CylindrobuUacea, 11,18 Ferrussaciidae, 11 Itruvia, 16
Cylindrobullidae, 11,17 Ferussininae, 13 Janthinidae, 10
Cymatiidae, 10 Ficidae, 10 Julia, 20
Cymbuliidae, 11 Filholiidae, 11,19 Juliacea, 12,20
Cypraeacea, 10,16 Fimbriidae, 12 Juliidae, 12,20
Cypraeidae, 10 Fionidae, 13,21 Kittlidiscidae, 9
Cyrtolitidae, 8 Fissurellacea, 9 Lacunidae, 9
Dawsonellidae, 9 Fissurellidae, 9 Lagini ops idae, 12
Deianiridae, 9 Flabellinidae, 13,21,22 Lamellariacea, 10,16
Dendrodorididae, 12,21 Fossaridae, 10,14 Lamellariidae, 10
Dendrodoridinae, 21 Fusinidae, 10 Lancidae, 11,18,19
Dendronotidae, 12 Galeodes, 16 Lancinae, 19
Dendronotoidea, 12,21 Galeodidae, 10,16 Latia, 18
Desmopteridae, 11 Gastropteridae, 11 Latüdae, 11,18,19
Diaphanacea, 11 Girtyspira, 14 Laubellidae, 9
Diaphanidae, 11,17 Glaucidae, 13 Lavige riidae, 15
Diastomidae, 10 Glaucinae, 22 Laxispira, 15
Didianema, 14 Glauconia, 14,15 Lemintina, 15
Dihelice, 15 Glauconiidae, 10,14,15 Lepetellidae, 9
Dironidae, 12 Gnathodoridacea, 21 Lepetidae, 9
Dorcasiidae, 11,19 Gnathodoridoidea, 12,21 Leptognatha, 21
Dorididae, 12,21 Goniaeolididae, 21 Leucodiscus, 14
Doridoidea, 12,21 Gonieolididae, 12 Leucorhynchia, 14
Doridoididae, 12,21 Gonieolis, 21 Lilax, 15
Doridoxidae, 12 Goniodorididae, 12,21 Limacidae, 12
Dotonidae, 12 Gonostomopsis, 12 Limapontiidae, 12
Duvaucelia, 21 Gosseletinidae, 8 Liotiinae, 13
Duvauceliidae, 21 Grangerellidae, 9,13 Littorinacea, 9
Ekadantinae, 13 Gymnentome, 14,15 Littorinidae, 9
Elachorbis 14
, Gymnosomata, 8,12,20 Lomanotidae, 12
Elasmonematidae, 9 Gymnodorididae, 12 Lophospiridae, 8
EUobiacea, 11 Hainesiinae, 13 Loxonematacea, 10
Ellobiidae, 11,13 Halgerdidae, 12,21 Loxonematidae, 10
Elysiacea, 12 Haliotidae, 9 Luciellidae, 8
Elysüdae, 12 Hancockiidae, 12 Lymnaeacea, 11,18
Endodontacea, 12,20 Haplotrematidae, 11,20 Lymnaeidae, 11,19
Endodontidae, 12 Harpidae, 10 Lymnaeinae, 19
Enidae, 11 Hedylopsidae, 11 Macluritacea, 8
Enteroxenidae, 11,16 Helcionellacea, 8 Macluritidae, 8
Entoconchidae, 11,16 Helcionellidae, 8 Macro »iphalina, 14
Entomotaeniata, 8,10,16,17 Helicacea, 12 Madrellidae, 12
Eolidoidea, 12,21,22 Helicidae, 12 Magilidae, 10
Eotomariidae, 8 Helicarionidae, 12 Maizaniidae, 9
Epitoniacea, 10,15 Helicinidae, 9,13 Marginellidae, 10
Epitoniidae, 10,15 Helicotomidae, 8 Mathildidae, 10,14,15
Epitonimn, 15 Helminthoglyptidae, 12 Maturipupa, 13
Eratoidae, 10,16 Heroidae, 12,21 Meekospira, 14
Euarminoidea, 21 Heterodorididae, 12,21 Meekospiridae, 10,14
Eubranchidae, 13,21,22 Heterurethra, 11 Megalo»iphalus, 14
Euconulidae, 12 Hexabranchidae, 12,21 Megaspiridae, 11,18,19
Eudoridacea, 21 Hipponicacea, 10 Megatyloma, 14
Euglandininae, 20 Hipponicidae, 10 Melanellacea, 10,16
Eulimacea, 10,16 Holopeidae, 9 Melanellidae, 10,16
Eulimidae, 10,16 Holopoda, 12 Melaniidae, 10
Euomphalacea, 8 Holopodopes, 11,19,20 Melanopsidae, 10,15
Melongenidae, 10,16 Orthalicidae, 12 Porostomata, 12,21

Mesogastropoda, 9,15,17,18 Orthurethra, 11,19 Portlockiellidae, 9
Mesurethra, 11,19 Otinidae, 11 Potamididae, 10
Metoptomatidae, 9 Otoconchidae, 12 Poteriidae, 9
Microdomatacea, 9 Ovulidae, 10,16 Procerithiidae, 10
Microdom atidae, 9 Oxynoacea, 12 Procymbuliidae, 11
Microhedylidae, 11 Oxynoidae, 12 Progrange re lia, 13
Micromelaniidae, 9 Pachygnatha, 12,21 Prosobranchia, 8
Milacidae, 12 Paedophoropodidae, 10 Protancylinae, 19
Mitracea, 10 Palaeotrochacea, 9 Protancylus, 19
Mitridae, 10,16 Palaeotrochidae, 9 Provermicularia, 15
Mitrinae, 16 Palaeozygopleuridae, 10 Psendobrochidium, 15
Modulidae, 10 Paosia, 14 Psetidoglauconia, 14
Murchisoniacea, 9 Parasita, 7,11,16 Pseudomelaniacea, 10,14,15
Murchisoniidae, 9 Paraturbinidae, 9 Pseudomelaniidae, 10,14
Muricacea, 10 Parmacellidae, 12 Pseiidomesalia, 14,15
Muricidae, 10,16 Partulidae, 11,19 Pseudophoracea, 9
Myrrhinidae, 13,22 Patellacea, 9 Pseudophoridae, 9
Naricidae, 10 Patellidae, 9 Pseudorhytidopilus 18 ,
Nassariidae, 10,16 Payettia, 18 Pseudorotella, 14

Nassidae, 10 Payettiidae, 11,18 Pseutosacculidae, 10
Naticacea, 10 Payettiinae, 18 Pseiidotritonia, 21
Naticidae, 10 Peraclidacea, 11 Pseudovermidae, 13,21
Neogastropoda, 7,10,16 Peraclididae, 11 Pseudozygopleuridae, 10
Neoplanorbidae, 19 Peruviella, 17 Pterotracheidae, 10
Neoplanorbinae, 19 Phanerobranchia, 12,21 Pulmonata, 8
Neptuneidae, 10,16 Phanerotrematidae, 8 Pupillacea, 11,19
Nerinea, 16 Phasianellidae, 9,13 Pupillidae, 11
Nerineacea, 11,16,17 Phenacolepadidae, 9 Pupinidae, 9
Nerineidae, 11,17 Philinacea, 11 Purpurinidae, 9
Nerinellidae, 11,17 Philinidae, 11 Pusia, 16,
Neritacea, 9,13 Philinoglossacea, 18 Pyramidella, 16
Neritidae, 9 Philinoglossidae, 11,18 Pyramidellacea, 11,17
Neritopsidae, 9 Philinoglossoidea, 7,11,17,18 Pyramidellidae, 11,14,15,16,17
Nododelphinulidae, 9 Philomycidae, 12,20 Pyramidulidae, 11
Nonsuctoria, 12,21 Phyllidiidae, 12,21 Pyrenidae, 10
Notaeolidiidae, 12,21 Phylliroidae, 12 Pyxipoma,. 15
Notaspidea, 8,12,16,20,21 Physidae, 11,19 Raphischismatidae, 9
Notobranchaeidae, 12 Phymatopleuridae, 9 Raphistomatidae, 8
Notodiaphanidae, 11 Pilidae, 9 Rathouisiidae, 13,22
Notodorididae, 12 Plagiothyridae, 9 Retusidae, 11
Nudibranchia, 12,20 Planaxidae, 10 Rhodacmeidae, 19
Odontostomidae, 12 Planitrochidae, 9 Rhodope, 20
Oleacina, 20 Planorbidae, 11,18,19 Rhodopidae, 12
Oleacinacea, 12,20 Platyacridae, 9 Rhodopoidea, 12
Oleacinidae, 12,20 Platyceratacea, 9 Rhytidacea, 11
Oleidae, 12 Platyceratidae, 9 Rhytididae, 11,20
Olividae, 10 Plethospiridae, 9 Ringiculidae, 11
Omalaxidae, 10,14 Pleurobranchacea, 12,20 Rissoacea, 9,13,14
Omalogyra, 14 Pleurobranchidae, 12 Rissoellidae, 9,14
Omalogyridae, 9,14 Pleuroceridae, 10,15,16 Rissoidae, 9,14
Omphalotrochidae, 8 Pleurodiscidae, 11 Rissoella, 14
Onchidiacea, 13 Pleuroprocta, 12,21 Rostellorbis, 14
Onchidiidae, 13,22 Pleurotomariacea, 8 Runcinidae, 11
Onchidorididae, 12,21 Pleurotomariidae, 9 Sacoglossa, 8,12,17,20,21
Onychochilidae, 8 Pneumodermatidae, 12 Sagdidae, 12,20
Oocorythidae, 16 Polyceridae, 12,21 Scalacea, 10,15
Opisthobranchia, 8 Polygyracea, 12,20 Scalidae, 10,15
Orbitestellidae, 9 Polygyratia, 20 Scaphandridae, 11
Orculidae, 11 Polygyridae, 12,20 Schizogoniidae, 9
Oreohelicldae, 12,20 Polytremariidae, 9 Scissilabra, 14
Oriostomatacea, 9 Pomatiasidae, 9 Scissurellidae, 9
Oriostomatidae, 9 Porcelliidae, 9 Scyllaeidae, 12
32 TAYLOR AND SOHL
Segmentella, 15 Subulitidae, 10 Trochidae, 9,15

Segnenzia, 15 Succineacea, 11 Trochomorphidae, 12
Seguenziidae, 10,15 Succineidae, 11 Trochonematacea, 9
Serpulorbis, 15 Suctoria, 12,21 Trochonematidae, 9
Sigmurethra, 11,19 Symmetrocapulidae, 9 Trochotomidae, 9
Sinuitidae, 8 Synceratidae, 9 Troostella, 17
Sinuopeidae, 8 Syrnolopsidae, 10,13 Trimcatellidae, 9
Siphonariacea, 11 Systrophiidae, 12,20 Tubinidae, 9
Siphonariidae, 11,18 valva, 20 Turbinidae, 9,13
Siphonium, 15 Tamanovalvacea, 20 Turridae, 10
Skeneidae, 9,13,14 Tamanovalvidae, 20 Turritellidae, 10,15
Skeneopsidae, 9,14 Teirwstoma, 14 Umbraculacea, 12,20
Solariacea, 14 Telarma, 21 Umbraculidae, 12
Solarüdae, 10,14 Temnotropidae, 9 Unnamed superfamily, 11
Solariorbis, 14 Tenagodus, 15 Urocoptidae, 12
Solaropsis, 20 Terebracea, 16 Urocyclidae, 12
Soleolifera, 8,13,22 Terebridae, 10 Valloniidae, 11
Spiratellacea, 11 Tergipedidae, 21 Valvatacea, 9
Spiratellidae, 11 Testacellacea, 12 Valvatidae, 9
Spiraxidae, 11,20 Testacellidae, 12 Vanikoridae, 10
Spraxinae, 20 Thaisidae, 10 Vasidae, 10
Spiroglyphus 15
, Thecosomata, 8,11 Vayssiereidae, 12,21
Spirostylidae, 10 Thiaridae, 10,13,14,15 Velainellidae, 9
Spurillidae, 22 Thliptodontidae, 12 Vermetidae, 10,14,15
Starkeytm, 14 Thyrophorellidae, 12 Vermicularia , 15
Stenacmidae, 16,18 Thysanophoridae, 12,20 Veronicellacea, 13,22
Stenoglossa, 10,15,16 Titiscaniidae, 9 Veronicellidae, 13,22
Stenothyridae, 9 Tonnacea, 10 Vertiginidae, 11
Stephopoma, 15 Tonnidae, 10,16 Vexillinae, 16
Stiliferidae, 10 Tornatellinidae, 19 Vitrinellidae, 9,13,14
Stiligeridae, 12 Tornidae, 9,14 Vitrinidae, 12
Stomatellidae, 9 Torre chry Sias, 20 Viviparacea, 9,13
Streptacididae, 11,17 Toxoglossa, 10,16 Viviparidae, 9,13
Streptaxacea, 11 Tra^ysmidae, 9 Volutacea, 10,16
Streptaxidae, 11,20 Trajanella, 14 Volutidae, 10
Streptoneura, 7,8 Trajanellidae, 14 Volvatella, 17
Streptostylinae, 20 Trichotropidae, 10 Weeksiidae, 8,13
Strombacea, 10,16 Trigonochlamydidae, 12 Xancidae, 10
Strombidae, 10 Trimusculidae, 11 Xenophoridae, 10,16
Strophocheilacea, 11,19 Triophidae, 12 Z alo pliancy lu s , 19
Strophocheilidae, 11,19 Triphoridae, 10,15 Zephyrinidae, 21
Struthiolariidae, 10 Tritonia, 21 Zonitacea, 12,20
Stylommatophora, 7,11,19 Tritoniidae, 12,21 Zonitidae, 12
Subulinidae, 11 Trochacea, 9 Zygitidae, 9
Subulitacea, 10,14 Trochaclisidae, 9 Zygopleuridae, 10
11, 1962, 1(1): 33-53
CRITICAL CATALOGUE OF THE NOMINAL GENERA AND SPECIES OF

NEOTROPICAL PLANORBIDAE
Harold W. Harry
Ida, Louisiana, U.S.A.
ABSTRACT
The genera of Neotropical Planorbidae have only a few species as far north as
the southern United States of North America. Of the 11 nominal genera which are
based on type species of the Neotropics, only 3, Taphius, Drepanotrema and
Acrorbis, are considered valid. An additional unnamed genus is recognized but
deliberately left unnamed here. Although the Nearctic genus Helisoma does ex-
tend as far south as Central America, it is pointed out that the citation of other
Nearctic and Palearctic genera from the Neotropics lacks confirmation by ana-
tomical data. Some Neotropical and African planorbid genera may be identical,
but as the former seem to have priority in all known cases, the question has not
been pursued.
An attempt is made to cite the original references and type localities of all
nominal planorbid species of the Neotropics. Of these, 207 are arranged in 21
"species groups." The members of each group are probably synonyms. Thirty
two species are Some of these seem to be distinct
lumped as incertae sedis.
from the 21 species groups, but lack of anatomical data prevents their being
placed in genera. Eleven nomina mida and 4 extralimital species are listed.
An extensive bibliography is given.
In the preparation of anatomical ac- cies. Fewadditions or changes have been

counts of the Puerto Rican Planorbidae I made manuscript since it was put
in the
found it difficult to determine what names, aside in the spring of 1958.
both generic and specific, ought to be ap- Over two hundred and fifty species of
plied to them. The present catalogue was planorbid snails were named from the
therefore compiled as a guide. An alpha- Neotropics between 1798 and 1957. Most
betical or chronological arrangement of of them were placed in the genus Planor-
the species and genera seemed of little bis, which served during the nineteenth
value, owing to the large number of spe- century as a general depository for nearly
cies described, and the many genera with every species of the family. Owing chief-
which they have been combined. The ar- ly to the works of Pilsbry (1934), F.
rangement as here presented was chosen Baker (1945) and Hubendick (1955), it is
partly on the basis of anatomical and eco- now apparent that a large number of ge-
logical studies, tobe presented elsewhere, nera may be recognized in the family.
and partly on the basis of studies of the These genera are based mainly on in-
collections and libraries of the U.S. Na- ternal anatomy rather than shell charac-
tional Museum and the Museum of Zoology ters. The genus Planorbis,3.s restricted
of the University of Michigan. I am in- by the workers cited, is now known to
debted to Dr. Harald Rehder and Dr. J.P.E. live only in the Palearctic faunal realm.
Morrison of the former institution, and Several other genera have been estab-
to Dr. Henry van der Schalle of the lat- lished for the species of the Neotropics.
ter, for their generosity in allowing me It is becoming increasingly evident that
free access to the important collections there are far fewer biological species of
of which they are custodians. I am also Planorbidae in the Neotropics than the
grateful to Dr. Lobato Paraense and Dr. number of nominal species recorded in
Walter Biese for providing anatomical the literature.
material of several South American spe- At the present stage of knowledge there
(33)
34 H. W. HARRY
can be no general agreement on the de- area, which usually have very limited
finition of the various genera, or their ranges. A few species are included in
limits (i.e., which species are to be in- the following catalogue which have their
cluded in each genus). There are also type localities in the United States, either
likely to be differences of opinion on because they have been cited from the
which of the nominal species are to be Neotropics, or seem (e.g. Planorbis eii-
recognized as biologically valid. Pro- cosmius Bartsch) to belong to Neotropi-
bably in both instances some arbitrary cal genera. Fossil genera and species
decisions will ultimately have to be made. have not been included.
The present catalogue tries merely to
summarize the nomenclature to date, and
thus to bring into sharper focus the many CHRONOLOGICAL LIST OF THE
problems involved, rather than to settle NOMINAL PLANORBID GENERA
them. OF THE NEOTROPICS
The interpretation of some early de-
scriptions of species depends on a pro- The Bulinidae and Planorbidae are
cess of elimination of the biological spe- sometimes considered as subfamilies of
cies known to occur in the area of the the Planorbidae. I prefer to consider
type locality. There are some type lo- them as families within the superfamily
calities which are open to doubt (e.g., Planorbacea,-"- for reasons to be given
Planorbis glabratus Say: see Pilsbry elsewhere. A catalogue of all nominal
1934). Of the many species described species of Plesiophysaf which is the only
from unknown localities (see Dunker 1848), known genus of Bulinidae in the New
only those referred to the Neotropics by World, has been given by Bequaert and
later authors have been included in the Clench (1939).
present list. It is now evident that some Neotropi-
cal planorbid species, at least in the

Taphiinae,âre probably congeneric and
GEOGRAPHICAL LIMITS perhaps even conspecific with African
species. In all cases presently known,
There is a rather well defined division the Neotropical genera have precedence
between the Neotropical andNearctic Pla- over the nominal African genera (e.g.,
norbidae. The line of separation is ap- Biomphalaria Preston 1910, Afroplanorbis
proximated by the southern boundary of Thiele 1931). Unfortunately, litüe atten-
the United States. Few species cross it. tion has been given to the African Pla-
Helisoma, which is predominantly a Ne- norbidae other than the Taphiinae, and so
arctic genus with several species, extends it is not practical at present to comment
into the Antilles and to Central America, on the probable relationships of the other
but its presence in South America is very groups on the two continents.
dubious. Drepanotrema and Taphhis are The species of Neotropical Planorbidae
predominantly Neotropical genera, each have sometimes been cited in the genera
with one or two species extending into Gyraidus, Promenetiis, Anisiis, Segmen-
Texas, Louisiana or Florida. In general, tina, Platwrbula, PImwrbis and perhaps
the species of Neotropical Planorbidae others. Such combinations with extra-
are widely distributed within their faunal limital genera have yet to be substanti-
realm. This is in striking contrast to ated by anatomical studies. In the fol-
the terrestrial gastropods of the same lowing list names proposed as subgenera
^This superfamily name is here used for the first time. ED.
^ Plesiophysa is not usually included in the bulinlds but placed in the separate planorbid sub-
family Plesiophysinae. ED.
^This subfamily name is here used for the first time. ED.
.
A CATALOGUE OF NEOTROPICAL PLANORBIDAE 35
or sections are included as genera, since several grounds. accepted, Armige rus
they would have that legal status accord- Clessin would have priority over Obstruc-
ing to the International Rules of Zoolog- tio Haas, but both names seem super-
ical Nomenclature. Similarly, subspe- fluous, since the type species show few
cific names and varieties are treated as or no characters of importance to sepa-
species. rate them from Taphius.
HELISOMA Swainson 1840. Type-spe- TROPICORBIS Brown and Pilsbry 1914.
cies, by original designation, Planorbis Type-species by original designation, PZa-
bicarinatus Sowerby (which is P, anceps norbis liebmanni Dunker (which is Pla-
Menke). A discussion of the taxonomy norbis havanensis Pfeiffer). This genus
of this genus has been given by F. C. is not well differentiated from Taphius,
Baker (1945). This is the only Nearctic and may be superfluous.
genus definitely known, on the basis of PLATYTAPHIUS Pilsbry 1924. Type-
anatomical data, to extend into the Neo- species by original designation, PZanoröes
tropics. heteropleurus Pilsbry and Vanatta. The
PLANORBINA^ Haldeman 1843. The anatomy of the genotype shows no major
vague description accompanying Halde- differences from Taphius, according to
man' s proposal is obviously open to a the account of Hubendick (1955, 1955a),
variety of interpretations, and he cited and Haas (1955) found intergrades in shell
no species in this group. Dall (1905) se- form between this nominal species and
lected as type-species Planorbis olivaceus Taphius andecolus (d'Orbigny). This ge-
"Spix" Wagner (which is Planorbis gla- neric concept may be superfluous.
bratiis Say). Pilsbry (1934) insisted that FOSSULORBIS Pilsbry 1934. Type-
P. olivaceus did not comply with the ori- species by original designation, PZa«or&¿s
ginal description of the genus. No species cultratus d'Orbigny (which is Planorbis
could be attributed to the genus with cer- kermatoides d'Orbigny). This generic
tainty before Dall's arbitrary selection concept may be superfluous.
of a genotype in 1905. Meanwhile the AUSTRALORBIS Pilsbry 1934. Type-
genus Taphius had been made available. species by original designation, Planorbis
TAPHIUS^H. and A. Adams 1855 .Type- guadaloupensis Sowerby (which is Pla-
species by monoty^y, Planorbis andecolus norbis glabratus Say). Proposed to re-
d'Orbigny. Unless some earlier name is place Planorbina "Haldeman" Dall 1905,
discovered, this genus will stand for a the anatomy shows no characters of ge-
large number of Neotropical and African neric importance to separate it from
Planorbid snails. Taphius, and this genus may be super-
DREPANOTREMA Fischer and Crosse fluous .
1880. Type-species by monotypy, Pla- ACRORBIS Odhnerl937. Type-species

norbis ysabelensis Crosse and Fischer by original designation, Acrorbis Petri-
(which is Planorbis anatinum d'Orbigny) cola Odhner. A very distinctive genus
ARMIGERUS 4 dessin 1884. The type- with only one known species.
species, by designation of Morrison (1947), OBSTRUCTIO Haas 1939. Genotype by
is Planorbis albicans Pfeiffer. H. . original designation, Planorbis janeirensis
Baker (1947) objected to Morrison's re- Clessin (which is probably Planorbis al-
surrection of this overlooked name on bicans Pfeiffer). Probably superfluous.
'*Sincemuch consideration has lately been given to the generic status of the various planorbid
species acting as vectors of Schistosoma mansoni, attention is drawn to a recent review, dis-
cussing relationships and priorities, by Barbosa et al. (1961, Ann. Mag. Nat. Hist., Ser. 13,
4:371-375) and to a request to the International Commission on Zoological Nomenclature by
Wright (1962, Bull. Zool. Nomencl., pt. 1, 19:39-41) for suppressing the above terms. Note
further that Walter (1962, Malacologia, 1(1): 115-137) considers Taphius to be generally distinct
from the group in question. ED.
-
36 H. W. HARRY
LATEORBIS F. .
Baker 1945. Type- 3. APPLANATUS von Martens 1899- Planorbis
speciesby original designation, Planorbis tenuis var. applanatus von Martens 1899, Biol.
palUdus C. B. Adams. Probably super- Centr. Amer., Moll., p. 384, PI. 21, Fig. 3.
(Central Mexico, Plateau of Mexico). Not P.
fluous .
applanatus Thomae 1854; see P. pertenuis

F.C. Baker 1940.
THE NOMINAL SPECIES OF 4. AURICULATUS Clessin 1884. Planorbis au-
NEOTROPICAL PLANORBIDAE riculatus Clessin 1884, Syst. Conch. -Cab.
Mart, and Chem. Ed. 2, Planorbis, p. 163,
In the following list, the citation of the PI. 24, Fig. 10. (Jamaica, Bluefields).
original description of each nominal spe- 5. BELIZENSIS Crosse and Fischer 1879- Plan-
cies is given. It includes references to orbis belizensis Crosse
and Fischer 1879,
any figures accompanying that description. Jour, de Conchyl. 27:342. Not figured. (Belize
The type locality is added in parentheses. (British Honduras)). 1880, Misc. Sei. .,
Moll., 2:68, PI. 32, Fig. 6, 6a, 6b.
Subsequent references are rarely cited,
except when they are emendations made 6. BOUCARDl Fischer and Crosse 1880. Planor-
by the original author. Thus, d'Orbigny bis tenuis var.boucardi Fischer and Crosse
(1837) redescribed and figured species

1880, Misc. Sei. ., Moll. 2:61, PI. 32, Fig.
3a-b. (Near Mexico City).
which he had described earlier (1835).
It notable that his later descriptions
is
7. CARIBAEUS d'Orbigny 1841. Planorbis caribae-
us d'Orbigny 1841, in Sagra's Hist. .Cuba,
differ somewhat from his original ones,
.
Moll., 1:193, PI. 13, Figs. 17-19. ("Environs

particularly in the number of whorls gi- de la Havane. aussi à la Vera Cruz, au
. .
ven. Dunker redescribed and figured spe- Mexique").

cies in the Conchylien-Cabinet which he CHAPALENSIS Pilsbry 1920. Planorbis tenuis
8.
had only described earlier (1848). Many Chapalensis Pilsbry 1920, Proc. Acad. Nat.
of the descriptions are so vague as to Sei. Phila., p. 192-193, Fig. 1, p. 193. ("Laguna
render identification almost impossible, de Chápala, State of Jalisco" (Mexico)).
and these will consequently be interfireted 9. CHIAPASENSIS Fischer and Crosse 1880.
differently by different authors. The spe- Planorbis ancylostomus var. chiapasensis
cies of Morelet (1849, 1851) are parti- Fischer and Crosse 1880, Mise. Sei. .,
Moll., 2:63-64, PI. 34, Fig. 5 ("E'tat de
cularly difficult to identify. The group-
Chiapas" (Mexico)).
ing of trivial names into genera and sub-
families requires some arbitrary deci- 10. COLON "Clessin" von Martens 1899. Planor-
sions. These will be necessary until stu- bis colon "Clessin" von Martens 1899, Biol.
dies have been made on the anatomy of Centr. Amer., Moll., p. 399- Error for P. coton
Clessin 1884, q.v.
all nominal species, preferably from their
type localities. Within each group, the 11. CONTRERAS! Pilsbry 1920. Planorbis con-
nominal species are arranged alphabeti- trerasi Pilsbry 1920, Proc. Acad. Nat. Sei.
cally. Phila., p. 193, text Fig. 2. ("Laguna de
Chápala, State of Jalisco"(Mexico)).
12. COSTARICENSIS Preston 1907. Planorbis cos-

HELISOMATINAE taricensis Preston 1907, Ann. and Mag. Nat.
Hist., 7 Ser., 20:496-497, text Fig. 16. ("Cata-
Genus Helisoma lina, Province of Guavacaste, Costa Rica").
Group of Helisoma foveale Menke 13. COTON "Morelet" Clessin 1884. Planorbis
coton "Morelet" Clessin 1884, Syst. Conch.
1. AFFINIS C.B. Adams 1849. Planorbis affinis
Cab. Mart, and Chem. Ed. 2, Planorbis, p. 209,
C.B. Adams 1849, Contrib. Conch. No. 3, p.
PI. 33, Fig. 3. (Locality unknown). Clessin
44. Not figured. (Jamaica).
spelled it coton on page 209, in the index and
2. ANCYLOSTOMUS Crosse and Fischer 1879- in the explanation of the figures. He did not
Planorbis ancylostomus Crosse and Fischer correct it in the errata, yet von Martens (1899)
1879, Jour, de Conchyl. 27:341. Not figured. changed the spelling (to P. colon, q.v.) and
(Vera Cruz, Mexico). 1880, Misc. Sei. ., attributed it to Central America, suggesting it
Moll., 2:63, PI. 32, Fig. 5a, 5b. is P. carihaeus d'Orbigny.
- -
14. EQUATORIUS Cousin 1887. Planorbis equa- 25. MINOR von Martens 1899- Planorbis carihaeus
toriiisCousin 1887, Bull. See. Zool. France, var. minor von Martens 1899, Biol. Centr.
12:263-264, PI. 4, Fig. 8. ("Equateur") Amer., Moll., p. 388. Not figured. ("E. Mexico:
spelled "aequatortus" on the plate. Vera Cruz in brackish marshes. N. Guatemala:
Coban").
15. EUDISCUS Pilsbry 1934. Helisoma duryi
eudiscus Pilsbry 1934, Proc. Acad. Nat. Sei. 26. MYSAURUS Mabille 1895- Planorbis mysaurus
Phila., 86:42-43, PI- 9, Figs. 4-9- ("Miami Mabille 1895, Bull. Soc. Philomatique Paris,
River above Miami" (Florida)). Has been cited 8 Ser., 7(2):63-64. Not figured. (LowerCali-
by several authors as occurring in the Antilles. fornia (Mexico)). Germain, 1921, Cat. Planor-
bidae, p. 54, decided this is a synonym of
16. EXAGGERATUS von Martens 1899- Planorbis
P. tumidus Pfr., after examining the type.
tenuis var. exaggeratus von Martens 1899,
Biol. Centr. Amer., Moll., p. 385- Not figured. 27. NICARAGUANUS Morelet 1851- Planorbis
("Central Mexico, Lake Patzcuaro"). nicaraguanus Morelet 1851, Test. Noviss. Ins.
Cub. et Amer. Centr. Pt. 2, p. 14. Not figured
17. FOVEALIS Menke 1830. Planorbis fovealis
("H. lacum nicaraguanensem").
Menke 1830, Synop. Method. Mollusc p. 37,
,
cites "Lister Conch, tab. 140 fig. 47," where 28. PAUCISPIRATUS Clessin 1885- Planorbis
the word "lam" appears adjacent to the figure. paucispiratus Clessin 1885, Syst. Conch. -Cab.
This has been interpreted by Pilsbry (1934:45) Mart, and Chem. Ed. 2, Planorbis, p. 223-224,
to be Jamaica. PL 33, Fig. 8- (Locality unknown). Von Mar-
tens, Biol. Centr. Amer., p. 400, noted its re-
18. FRAGILIS Dunker 1850. Planorbis fragilis
semblance to P. ivyldi Tristram.
Dunker 1850, Syst. Conch. -Cab. Mart, and
Chem., Ed. 2, Planorbis, p. 46-47, PL 10, 29- PERTENUIS F.C. Baker 1940. Planorbis
Fig. 41-43- ("In der Nähe von Mexico mit tenuis vat. pertenuis F.C. Baker 1940, Nautilus
Planorbis lenuis Phil."). 54:97. New name for P. tenuis var. applanatus
von Martens 1899-
19. GUATEMALENSIS Clessin 1884. Planorbis
guatemalensis Clessin 1884, Syst. Conch. 30. PERUVIANUS Broderip 1832. Planorbis peru-
Cab. Mart, and Chem., Ed. 2, Planorbis, p. vianas Broderip 1832, Proc. Zool. Soc. London
209-210, PL 32, Fig. 7. ("Centralamerika, p. 125, Not figured. ("Hab. in Peruvia (Mala-
Guatemala"). briga, province of Truxillo)").
20. HUMILIS C.B. Adams 1851- Planorbis humilis 31. SALVINI "Tristram" Clessin 1884. Planorbis
C.B. Adams 1851. Contrib. Conch. No. 8, p. sö/f»«/ "Tristram" Clessin 1884, Sysc Conch.
131-132. Not figured. (Jamaica). Cab. Mart, and Chem. Ed. 2, Planorbis, p.
207-208, PL 31, Fig. 8. ("Centralamerika,
21. INTERMEDIUS "Philippi" Dunker 1850. Plan-
Guatemala").
orbis "Philippi" Dunker 1850,
intermedins
Conch. -Cab. Mart, and Chem. Ed. 2, Planorbis, 32. SINUOSUS Bonnet 1864. Planorbis sinuosus
p. 39, as a synonym of P. tumidus Pfeiffer.
Bonnet 1864, Rev. et Mag. de ZooL, p. 280,
(No locality, evidently Mexico). PL 16, Figs. PL 22, Fig. 3, 3a. (Rio Grande River, New
18, 19 (appeared 1844). Clessin, 1884, Ibid., Mexico). Pilsbry (1934:44) thought this is a
p. 196, PL 11, Fig. 1,2 (appeared 1882). (Vera
synonym of P. caribaeus d'Orbigny.
Cruz, Mexico). Not P. intermedias T. de Char- 33. SOLIDUS "Wiegmann" von Martens 1899-
pentier 1837, N.D. Allg. Schweiz. Gesellsch. Planorbis solidus "Wiegmann" von Martens
1, P- 21. 1899, Biol. Centr. Amer., Moll., p. 384i von
22. JUVENILIS von Martens 1899. Planorbis tenuis Martens cites only "Mus. Berol." as the source
var. juvenilis von Martens 1899» Biol. Centr. of Wiegmann's name. He proposed the new name
Amer., MolL, p. 384, PL 21, Fig. 4. (Central P. juvenilis to replace it, as P. solidus was
Mexico: ditches near the city of Mexico). preoccupied by P. solidus C. Thomae 1845,
Proposed as a new name for P. solidus Weig- Jahrb. Ver. Nat. Nassau 2:153, and P. solidus
man, q.v. Dunker 1850 Syst. Conch. -Cab. p. 60.
STREBELIANA Fischer and Crosse 1880-

23. LENTUS Say 1834. Planorbis lentus Say 1834, 34.
Amer. Conch. 6:6, PL 4, Fig. 1. (Mexico, Ojo Planorbis ancylostomus var. B. Strebeliana
de Agua, and Canal at New Orleans, La.) Fischer and Crosse 1880, Misc. Sei. .,
Moll., 2:63-64. Not figured. (Laguna des Cocos
24. MEXICANUS "Ziegler"Dunker 1850. Planorbis
and Rio Tenoya, in the state of Vera Cruz
mexicanus "Ziegler" Dunker 1850, Syst.
(Mexico)).
Conch. -Cab. Mart, and Chem., Ed. 2, Planorbis,
TENUIS "Philippi" Dunker 1850. Planorbis
p. 45, as a synonym of P. tenuis "Philippi" 35.
Dunker. No locality. tenuis "Philippi" Dunker 1850, Syst. Conch.-
-
38 H. W. HARRY
Cab. Mart, and Chem., Ed. 2, Planorbis, p. DREPANOTREMATINAE

45, PI. 9, Figs. 14-16, 17-19 (appeared 1850);
PI. 16. Fig. 23-25 (appeared 1844, without
Genus Drepanotrema
name). ("Häufig in Gräben der Umgegend von
Mexico mit IJmnaeus subulatus Dkr."). Group of Drepanotrema anatinum
36. TUMENS Carpenter 1856. Planorhis tumens d'Orbigny
Carpenter 1856, Cat. Mazatlan Shells in Brit.
Mus. p. 181. Not figured ("Mazatlan, not com- 42. ANATINUS d'Orbigny 1835. Planorbis anatinus
mon"). Carpenter realized this was an extreme- d'Orbigny 1835, Mag. de Zool. V(62):28. Not
ly variable species, and noted that this species, figured. (Rio Parana, Argentina, from the
P. affinis C.B. Adams and P. lentus Say "may stomach of a duck). 1837, Voyage Amer.
hereafter be proved identical." Mérid., p. 351, PI. 45, Figs. 17-20.
37. TUMIDUS Pfeiffer 1839- Planorbis tumidus 43. ARACASENSIS "Gundlach" Pfeiffer 1879-
Pfeiffer 1839, îegm. Arch. f. Naturgesch. p. Planorbis aracasensis "Gundlach" Pfeiffer
354, Not figured. (Cuba). The original descrip- 1879, Malakozool. Blatt. 4:179- Not figured.
tion is merely ^'Planorbis tumidus Pfr. Spec- (Aracas Lake, Tinidad). The original descrip-
imina incompleta, Pi. fragili affina." The tion consists only of"15. Planorbis Aracasen-
species is scarecly recognizable froni this sis Gund. Eine sehr niedliche und winzige Art
description, if the name is not nude. aus dem See Aracas bei Trinidad." Several
authors have considered this term a nomen
38. TUMIDUS "Pfeiffer" Dunker 1850. Planorbis
nudum. No species could be definitely recog-
tumidus "Pfeiffer" Dunker 1850, Syst. Conch.
nized from it.
Cab. Mart, and Chem., Ed. 2, Planorbis, p. 39,
PI. 7, Figs. 10-12. (San Juan and Havana,
44. ARACASENSIS "Gundlach" Clessin 1884.
Cuba; Vera Cruz and Vampa, Mexico). P. in- Planorbis aracasensis "Gundlach" Clessin
termedius Philippi and P. caribaeus d'Orbigny 1884, Syst. Conch. -Cab. Mart, and Chem., Ed.
are listed as synonyms. The original citation 2, Planorbis, p. 143, PL 15, Fig. 7. (Esperanza,
of Pfeiffer's is given, but since that is too Pinos de Rio, Cuba).

inadequate for recognition, the name should 45. ESPERANZENSIS Tryon 1866. Planorhis es-
date from Dunker's description, as Pilsbry peranzensis Tryon 1866, Amer. Jour. Conch.
(1934) noted. 2:10, PI. 2, Figs. 11-13 (Esperanza, Cuba).
39- UHDEI von Martens 1899- Planorbis tenuis v. 46. HALDEMANI C.B. Adams 1849- Planorbis
uhdei von Martens 1899» Biol. Centr. Amer., haldemani C.B. Adams 1849, Contrib. Conch.
Moll., p. 385, PI. 21, Fig. 2. (Central America). No. 3, P- 43- Not figured. (Jamaica).
40. WYLDI Tristram 1861. Planorbis y W» Tristram 47. INVOLUTUS "Dunker" Clessin 1884. Planor-
1861, Proc. Zool. Soc. London p. 232. Not bis involutus "Dunker" Clessin 1884, Syst.
figured. ("Lake of Dueñas" (Guatemala)). Conch. -Cab. Mart, and Chem., Ed. 2, Planorbis,
p. 143- As a synonym of P. aracasensis
"Gundlach" Clessin.
Group of Helisoma eyerdami Clench 48. ISABEL ".Morelet" Sowerby 1877. Planorbis
Isabel "Morelet" Sowerby 1877 in Reeve's
and Aguayo 1932
Conch. Icon. V. 20, Planorbis. PI. 12, Fig.
101. (Locality unknown). Goodrich and van der
41. EYERDAMI Clench and Aguayo 1932. //e/isowa Schalie cite this species from Guatemala
eyerdami Clench and Aguayo 1932, Proc. New
(1937, Moll. Peten..., p. 33), and Aguayo
England Zool. Club 13:38. Not figured. ("Lake
(1933, Nautilus 47:64) considered it a synonym
Miragoane, two miles southeast of Miragoane, of D. anatinum d'Orbigny.
Haiti.") Clench and Aguayo 1937, Mem. Soc.
Cubana Hist. Nat. 11:68, PI.
49. NIGELLUS Lutz 1918. Planorbis (Spiralina)
7, Fig. 7.
Two nigellus Lutz 1918, Mem. Inst. Osw. Cruz
paratypes of this species in the Univer-
sity of Michigan Museum of Zoology show a 10:55- Not figured. (Pool near Manguinhos
(Brazil)).
prominent, thickened keel on both shoulders,
and a slight flare of the lip in these positions. 50. YSABELENSIS Crosse and Fischer 1879.
I could not determine whether the embryonic Planorbis ysabelensis Crosse and Fischer
whorl was angled. This species seems to be 1879, Jour. Conchyl. 27:342-343- Not figured.
distinct from the other Neotropical populations (Balancan, province of Tabasco, Mexico, and
of Helisoma, and it may even be a Taphius, Yzabel, Guatemala). 1880, Miss. Sei. .,
Anatomical studies are needed to settle the Moll., 2:75-76, PL 33, Fig. 2, 2c.
question. ^This Subfamily name is here used for the first time. ED.
-
Group of Drepanotrema limayanum V(62) p. 27. Not figured. (Province of Lima,

Republic of Peru). 1837, Voy. Amér. Mérid. p.
Lesson
350-351, PI. 45, Figs 5-8.
51. AHENUM H.B. Baker 1930. Dre¡>anotrema
a/'ewMwH.B. Baker 1930,Occ. Pap. Mus. Zool.,
64. PURUS von Martens 1868. Planorbis purus von
Univ. Mich. 210:49-50, Figs. 2-4 (Bejuma,
Martens 1868, Malakozool. Blatt. 15, p. 190,
Venezuela).
Not figured. (Rodersberg, Jacuhy Region,
52. CATILLUS Anton 1839- Planorbis catillus Brazil).
Anton 1839, Verz. d. Conch.... p. 51- Not
65. REDFIELDI C.B. Adams 1849. Planorbis red-
figured. (Lima (Peru)).
fieldi C.B. Adams 1849, Contrib. Conch. No.
53. DECLIVIS "Gould" Clessin 1884. Planorbis 3, p. 43. Not figured. (Jamaica).
"Gould" Clessin 1884, Syst. Conch.
declivis
66. SCHUBARTI Haas 1938. Hippeutis schubarti
Cab. Mart, and Chem., Ed. 2, Planorbis, p.
Haas 1938, Arch. f. Molluskenk., 70:49-50,
142-143, PI. 17, Fig. 3. (Cuba). Not P. declivis
Fig. 7 (p. 47). (Acude Triumpho, 1000 m alt..
Genth 1848, nor P. declivis Tate 1869, nor P.
State of Pernambuco, Brazil).
declivis Sowerby 1877.
67. SUCCINEUS Sowerby 1877. Planorbis succineus
54. FAUSTI Ferguson and Gerhardt 1956. Drepan-
Sowerby 1877, Reeve's Conch. Icon. Vol.
in
otrema fausti Ferguson and Gerhardt 1956,
20, Planorbis, PI.3, Figs. 19a, b. (No locality).
Bol. Ofic. Sanit. Panamericana 41:340-341,
It was noted in the index of that volume that
Fig. 10, 11. (Sabana Seca, Puerto Rico).
this entry represents P. redfieldi C.B. Adams,
55. HOFFMANI F.C. Baker 1940. Drepanotrema and that P. succineus was an error.
hoffmani F.C. Baker 1940, Nautilus '54:96-97,
68. SUMICHRASTI Crosse and Fischer 1879.
PI. 8. (Isabela, Puerto Rico).
Planorbis sumichrasti Crosse and Fischer
56. JAMAICENSIS "Dunker" Clessin 1884. Plan- 1879, Jour, de Conchyl. 27:342. Not figured.
orbis jamaicensis "Dunker" Clessin 1884, (Cacoprieto, Isthmus of Tehuantepec, Mexico).
Syst. Conch. -Cab. Mart, and Chem., Ed. 2, 1880, Miss. Sei. ., Moll., 2:69, PL 33,
Planorbis, p. 126. As a synonym of P. surin- Figs. 6-6d.
awews/s "Dunker" Clessin. Not P. jamaicensis
69. SURINAMENSIS "Dunker" Clessin 1884. Plan-
Bolten 1798.
orbis surinamensis "Dunker" Clessin 1884,
LANERIANUS d'Orbigny 1841. Planorbis lan-
57.
«5
Moll.,
d'Orbigny 1841, in Sagra, Hist. .. Cuba,
1:195, PI. 14, Figs 1-4. (Near Havana,
Syst. Conch. -Cab. Mart, and Chem., Ed. 2,
Planorbis, p. 126, PL 17, Fig. 11. (Jamaica,
and the region of Paramaribo, Surinam).
Cuba).
70. TAENIATUS Morelet 1849. P/a«orèis taeniatus
58. LENZI van Benthem Jutting 1943- Anisus Morelet 1849, Test. Noviss. Ins. Cub. et Amer.
(Gyraulus) lenzi van Benthem Jutting 1943, Centr. Pt. 1, p. 17. Not figured. (Isle of Pines
Arch. Hydrobiol. 39:480, text Fig.
f. and d. (Cuba)).
("Rio S3o Francisco, Sao Pedro Dias-Bucht,
Pernambuco " (Brazil)). Group of Drepanotrema kermatoides
59. LIMAYANA Lesson 1830. Planorbis limayana d'Orbigny
Lesson 1830, Voyage autour du monde . . .
ANITENSIS Cooper 1893. Planorbis (Anisus)
71.
V. 2, Pt. 1, p. 330. Not figured. ("Entre Callao anitensis 1893, Proc. Calif. Acad.
Cooper
et Lima au Pérou" (Peru)).
Sei. Ser. 2, V. 3, p. 341-342, PL 14, Fig. 8.
60. LUCroUS 1839. Planorbis lucidas
Pfeiffer (Santa Anita, Lower California and 10 miles
Pfeiffer 1839, Wiegm. Arch. f. Naturgesch. p. from San Jose del Cabo, Lower Calif. (Mexico)).
354. Not figured. (Cuba).
72. BARBADENSIS "Dunker" Clessin 1884. Plan-
61. MELLEUS Lutz 1918. Planorbis melleus Lutz orbis barbadensis "Dunker" Clessin 1884,
1918. Planorbis melleus Lutz 1918, Mem. Inst. Syst. Conch. -Cab. Mart, and Chem., Ed. 2,
Oswaldo Cruz 10:54, PL 16, Fig. 5a,b,c,d. Planorbis, p. 118-119, PL 11, Fig. 4. (Bar-
(Manguinhos and Meyer in Rio de Janeiro; bados).
kilometer 22 of the Leopoldina Railway;
73. BONARIENSIS Strobel 1874. Planorbis kerma-
Aracaju; Parahyba (Brazil)).
toides bonariensis Strobel 1874, Mat. Malac.
62. MENISCUS Guppy 1871. Planorbis meniscus Argentina, p. 33-34, PL 2, Fig. 1. (Palermo
Guppy, 1871, Amer. Jour. Conch. 6:310. Not and Belgrano, vicinity of Buenos Aires,
figured. (Chatham River, Erin, Trinidad). Argentina).
63. PAROPSEIDES d'Orbigny 1835- Planorbis 74. CULTRATUS d'Orbigny 1841. Planorbis cul-
paropseides d'Orbigny 1835, Mag. de Zool. tratus d'Orbigny 1841, in Sagra, Hist. Cuba, . .
,
40 H. W. HARRY
Moll., Vol. 1, p. 196, PI. 14, Figs. 5-8. (Cuba? figured. (Guadeloupe, Antilles). 1876, Jour, de
probably Martinique). Conchyl. 24:388, PL 11, Fig. 3-
75. DEPRESSISSIMUS Moricand 1839- Planorbis 87. CHITTYI Aguayo 1935- Drepanotrema chittyi
defrressissimus Moricand 1839, Mém. Soc. Aguayo 1935, Mem. Soc. Cubana Hist. Nat.
Phys. Hist. Nat. Genève, 8:143-144, PI. 3, 9:121. New name for Planorbis angulatus
Fig. 10, 11. (Bahia, Brazil). Chitty 1853, q.v.
76. DUENASIANUS Tristram 1861. Planorbis 88. CIMEX Moricand 1839- Planorbis cimex Mori-
duenasianus Tristram 1861, Proc. Zool. Soc. cand 1839- Mém. Soc. Phys. Hist. Nat. Genève
London p. 232. Not figured. (Lake of Dueñas 8:143, PI. 3, Figs. 8-9- (Bahia, Brazil).
(Guatemala)).
89. LABROSUM Pilsbry 1934. Drepanotrema cul-
77. HARRYI Ferguson and Gerhardt 1956. Drepan-
tratum labrosum Pilsbry 1934, Proc. Acad.
otrema harryi Ferguson and Gerhardt 1956,
Nat. Sei. Phila. 86:61, PL 11, Figs. 9-11.
Bol. Ofic. Sanit. Panamericana 41:337-340,
(Brownsville, Texas).
Figs. 4-6 (p. 338) and Fig. 13 (p. 339). (St.
Croix, Virgin Islands). 90. MACNABL\NUS C.B. Adams 1849- Planorbis
macnabianus C.B. Adams 1849, Contrib. Conch.
78. KERMATOIDES d'Orbigny 1835- Planorbis No. 3, P- 43- Not figured. (Jamaica).
kermatoides d'Orbigny 1835, Mag. de Zool.
V(62):27. Not figured. (Province of Lima, 91. PISTIAE H.B. Baker 1930. Drepanotrema
Peru). 1837, Voy. Amér. Mér., Moll., p. 350, cimex pistiae H.B. Baker 1930, Occ. Pap.
PI. 45, Figs. 1-4. Mus. Zool., Univ. Mich. 210:50-51, PL 30.
Fig. 1, (Tucacas, Venezuela).
79. NORONHENSIS Smith 1890. Planorbis noron-
hensis Smith 1890, Jour. Linn. Soc. Lond. 92. POEYANUS Clessin 1884. Planorbis poeyanus
20:502-503, PI- 30, Figs. U-llb. (The lake on Clessin 1884, Syst. Conch. -Cab. Mart, and
the southwest corner of Fernando Noronha Chem., Ed. 2, Planorbis, p. 205-206, PL 31,
(Brazil)). Fig. 2. ("Die Antillen; St. Domingo (Coll.
Dunker) Havannah (Coll. Morelet)").
80. PANUCO Pilsbry 1934. Drepanotrema cultratum
panuco Pilsbry 1934, Proc. Acad. Nat. Sei. 93. UNGULATUS "Chitty" Sowerby 1877. Planor-
Phila. 86:60-61, PL 11, Figs. 4-5a. (Pasture bis ungulatus "Chitty" 1877, in Reeve's
west of San D i e g u i t o, San Luis Potosi, Conch. Icon. Vol. 20, Planorbis. PL 8, Fig.
Mexico). 62. (Jamaica). Error for P. angulatus Chitty.
81. PENINSULARIS Cooper 1893- Planorbis Group of Drepanotrema heloicus

(AnisusF) peninsularis Cooper 1893, Proc. d'Orbigny
Calif. Acad. Sei., Ser. 2, 3:342, PL 14, Fig. 9.
("With P. anitensis, in the same laguna"). 94. CASTANEONITENS Pilsbry and Vanatta 1896.
Planorbis castaneonitens Pilsbry and Vanatta
82. PULCHELLUS "Philippi" Clessin 1884. 1896, Proc. Acad. Nat. Sei. Phila. p. 561-562,
Planorbis "Philippi"
pulchellus Clessin PL 27, Figs. 10-12. (Ponds and small stream
1884, Syst. Conch. -Cab. Mart, and Chem.,Ed. 2, near Maldonado, Uruguay).
Planorbis, p. 137, PL 11, Fig. 6; PL 16, Figs.
12-14. (Bolivia). 95. HELOICUS d'Orbigny 1835. Planorbis heloicus
d'Orbigny 1835, Mag. de ZooL, V(62):27. Not
83. TANCREDII Paravicini 1894. Planorbis tan-
figured. (Montevideo, Uruguay). 1837, Voy.
credii Paravicini 1894, Boll. Mus. Zool. Anat.
Amér. Mérid. p. 349, PL 45, Figs. 9-12.
Comp. Univ. Torino 9(181):8-9. Not figured.
This species is probably very circumscribed.
(Asuncion, Paraguay).
I have seen material in the U.S. National Mu-
84. TENUISSIMUS "Philippi" von Martens 1873. seum and University of Michigan Museum of
Planorbis lenuissimus "Philippi" von Martens Zoology which is apparently paratypic of
1873, Binn. Moll. Venez., p. 197. As a syno- Pilsbry and Vanatta's species. The general
nym of P. cultratus d'Orbigny. form and color of the shell suggest a species
close to D. limayanum, yet the absence of any
Group of Drepanotrema cimex Moricand spiral sculpture on the shell, and the fact
that d'Orbigny (1837) says the animal is uni-
85. ANGULATUS Chitty 1853- Planorbis angulatus
formly blackish in color, leaves much room for
Chitty 1853, Contrib. Conch. No. 1, p. 18-19.
doubt about its systematic position.
Not figured. (Near the head of the navigable
part of Black River, St. Elizabeth (Jamaica)). Genus undescribed: Group of "Planorbis"
Not. P. angulatus Wood, 1838. See D. chittyi. salleanus Dunker 1853
86. BAVAYI Crosse 1875- Planorbis bavayi 96. CIRCUMLINEATUS Shuttleworth 1854. Planor-
Crosse 1875, Jour, de Conchyl. 23:329. Not bis circumlineatus Shuttleworth 1854. Mitth.
Naturf. Gesellsch. Bern. p. 96-97- Not figured. Lauricocha, unde flumen Maranon sive Ama-
(Humacao, Puerto Rico). zonas nascitur").
97. NORDESTENSIS Lucena 1954. Tropicorbis 106. MONTANUS d'Orbigny 1835- Planorbis mon-
nordestensis Lucena 1954, Rev. Brasil. Malar. tanus d'Orbigny 1835, Mag. de Zool. V(62):26.
D. Trop. 6(3):329-331; 1955, Jour, de Conchyl., Not figured. ("Habitat in lacu Titicaca re-
95:20-22. These references I have not seen. publica Boliviana). 1837, Voy. Amér. Mérid.,
But see Paraense and Deslandes 1958:281. p. 345-346, Pl. 44, Figs. 5-8.
98. SALLEANUSDunlcer 1853- Planorbis salleanus 107. PENTLANDI "Valenciennes" Beck 1837.
Dunker 1853, Proc. Zool. See. Lond. 21:53-54. Planorbis pentlandi "Valenciennes" Beck
Not figured. (St. Domingo). 1837, Index, moll, praes. ... p. II9. Not de-
scribed or figured. As a synonym of P. ande-
99. SANTACRUZENSIS "Nevill" Germain 1923-
colus d'Orbigny.
Planorbis (Gyraulus) santacruzensis "Nevill"
Germain 1923, Cat. Planorbidae, Rec. Indian 108. TITICACENSIS Clessin 1884. Planorbis
Museum 21, p. 138-139, Fig. 18-21 (p. 139), titicacensis Clessin 1884, Syst. Conch. -Cab.
PI. 4, Figs. 10, 13 and 14. (St. Croix, Virgin Mart, and Chem., Ed. 2, Planorbis, p. 147, PL
Islands). 12, Fig. 23-25. (Lake Titicaca).
100. SIMMONSI Ferguson and Gerhardt 1956. Dre-
Group of Taphius pronus von
panotrema simmonsi Ferguson and Gerhardt
1956, Bol. Ofic. Sanit. Panamericana 41:336-
Martens 1873
337, Figs. 1-3 (p. 338) and Fig. 12 (p. 339). 109. COSTATUS Biese 195L Taphius costatus
(Sabana Seca and Arroyo, Puerto Rico). Biese 1951, Bol. Mus. Nac. Hist. Nat. (Chile)
25:116-117, Fig. 5, p. 130; Pl. 6, Figs. 1-3
Genus Acrorbis ("Cuchicha, 3,800 m. de altura" (Chile)).

Topotypic material identified by Biese in the
101. PETRICOLA Odhner 1937. Acrorbis petricola University of Michigan Museum of Zoology is
Odhner 1937, Ark. Zoologi Bd. 29B, No. 14,
apparently of this group.
p. 1-8. Ten text figures. (Santa Catharina,
Brazil). 110. PRONUS von Martens 1873. Planorbis pronus
von Martens 1873, Binnenmoll. Venez, p. 198-
TAPHUNAE 199, PI. 2, Fig. 5. ("Valenciasee" (Vene-
zuela)).
Genus Taphius 111. SOLroULUS Clessin 1885. Planorbis solidulus
Clessin 1885, Syst. Conch.-Cab. Mart, and
Group of Taphius andecolus d'Orbigny
ehem., Ed. 2, Planorbis. p. 224, PL 33, Fig.
102. ANDECOLUS d'Orbigny 1835- Planorbis ande- 10 ("?Coll. Morelet"). Of unknown locality,
colus d'Orbigny 1835, Mag. de Zool. V(62):26. this species is included here because von
Not figured. ("Habit, lacu Titicaca (república Martens, 1899, 400 (Biol. Centr. Amer.) notes
p.
Boliviana)"). 1837, Voy. Amér. Mérid., p. and P. subpronus,
its similarity to his P. />ro«tts
346, Pl. 44, Figs. 1-4. and von Martens claimed he examined Clessin's
103. CONCENTRATUS Pilsbry 1924. Planorbis material.
(Taphius) andecolus concentratus Pilsbry 1924, 112. SUBPRONUS von Martens 1899- Planorbis
Proc. Acad. Nat. Sei. Phila., p. 50, PI. 4, (Taphius) subpronus von Martens 1899, Biol.
Figs. 2, 2a, 2b. (Lake Titicaca, Peru). Centr. Amer. p. 396, PL 21, Fig. 15- ("S.E.
104. HETEROPLEURUS Pilsbry and Vanatta 1896. Mexico: Amatitan, State of Tabasco").
Planorbis heteropleurus Pilsbry and Vanatta 113. TRIGYRUS Philippi 1869- Planorbis trigyrus
1896, Proc. Acad. Nat. Sei. Phila. p. 562, PL Philippi 1869, Malakazool. Blatt. 16:39. Not
26, Figs. 1,2 and 3- (Lake Titicaca). figured. ("Specimen unicum cum Pl, helophilo
This species was designated the genotype d'Orb. in litore Peruviae ad Pimentel lectum
of Platytaphius by Pilsbry 1924, Proc. Acad. vidi").
Nat. Sei. Phila., p. 51. Hubendick (1955,
1955a) examined the anatomy of this species, caloderma Pilsbry 1923
Group of Taphius
and failed to note any character of significance
from Taphius s.s. Haas (1955) considers P.
114. CALODERMA Pilsbry 1923- Planorbis calo-
heteropleurus to be merely a variant of Taphius
derma Pilsbry 1923, Nautilus p. 143-144. Not
andecolus d'Orbigny.
figured. ("Esmerelda, Guatemala"). F.C.
105. LAURICOCHAE Philippi 1869- Planorbis Baker 1945, Moll. Fam. Planorbidae, Pl. 115,
lauricochae Philippi 1869, Malakozool. Blatt. Figs. 2-4, figured cotypes. Anatomical studies
16:38. Not figured. ("Habitat in lacu andino may show this species to be a Helisoma.
42 H. W. HARRY
Group of Taphius eucosmius Bartsch 1908 Spix to a species which was not that of Spix,
and which had received no name of its own.
115. EUCOSMIUS Bartsch I9O8. Planorbis eucosmius Not P. confusus Rochebrunne 1881.
Bartsch 1908, Proc. U.S. Nat. Mus. 33:699- 126. CUMINGIANUS Dunker 1848. Planorbis cuming-
PI. 57, Figs. I-3. ("Greenfield Pond, Wilming- ianus Dunker 1848, Proc. Zool. Soc. London
ton, North Carolina" (U.S.A.)).
p. 41. Not figured. (Locality unknown), 1850,
116. VAUGHANI Bartsch 1908. P/awor¿ís eucosmius Syst. Conch. -Cab. Mart, and Chem., Ed. 2,
vaughani Bartsch 1908, Proc. U.S. Nat. Mus. Planorbis, p. 49, PI. 8, Fig. 1-3- (No locality).
33:699-700, Pi. 57, Figs. 4-6 ("Burkes Place, Placed in the synonymy of Pi. olivaceus Spix
Louisiana" (U.S.A.)). by Germain, 1921, Cat. Planorbidae, p. 45-
I am indebted to Dr. J. P.E. Morrison for 127. DENTIFER Moricand 1853- Planorbis dentifer
calling my attention to the striking similarity
Moricand Jour, de Conchyl. 4:37. Not
1853,
of these shells to Taphius. The band
spiral
figured. (Lake Baril, near Bahia, Brazil). See
of dark brown color is distinctive, and not found
Paraense, 1957, Proc. Mai. Soc. London, 32: 175-
in any other planorbid, to my knowledge.
128. FERRUGINEUS "Spix" Wagner 1827. Planor-
Group of Taphius glabratus Say 1818 bis ferrugineus "Spix" Wagner 1827, Test.
Fluv. Bras p. 26, PL 18, Fig. 1. As a
117. ALBESCENS "Spix" Wagner 1827. Planorbis synonym of P. olivaceus "Spix" Wagner.
albescens "Spix" Wagner 1827, Test. Fluv.
Bras. p. 27, as a synonym of P- lugubris 129. GUADALOUPENSIS Sowerby 1822. Planorbis
Wagner, q.v. guadaloupensis Sowerby 1822, Genera of Re-
cent and fossil Shells, No. 4. No description.
118. ANTIGUENSIS "Guilding" Sowerby 1877.
(Guadeloupe).
Planorbis "Guilding" Sowerby
antiguensis
1877, in Reeve's Conch. Icon. Vol. 20, Planor- 130. GLABRATUS Say 1818. Planorbis glabratus
bis, PI. 12, Figs. lOOa-b. (Antigua, West Say 1818, Jour. Acad. Nat. Sei. Phila. p. 280.
Indies). Not figured. ("South Carolina"). Pilsbry (1934)
argued that the locality was a mistake, and
119. ANTILLARUM Beck 1837. Planorbis antillarum
that Say's material probably came from Guade-
Beck 1837, Index, moll, praes. ., p. 120,
. .
loupe.
Refers to Chemnitz, Conch. -Cab., Ed. 1, Vol.
9, 12th Abt., Fig. 1118. (Antilles). 131. IMMUNIS Lutz 1923. Planorbis immunis Lutz
1923, Nautilus 37:36. New name for P. confusus
120. BECKI Dunker 1850. Planorbis becki Dunker
Lutz 1918, q.v.
1850, Syst. Conch. Cab. -Mart, and Chem., Ed.
2, Planorbis. p. 48, PL 8, Figs. 4-6 (Brazil) 132. LUGUBRIS Wagner 1827. Planorbis lugubris
Wagner 1827, Test. Fluv. Bras p. 27, PL 18,
121. BLAUNERI "Shuttleworth" Germain 1921.
Figs. 3-6. (Ilheos and Almada, Bahia, Brazil).
Planorbis (Planorbina) blauneri "Shuttleworth"
Germain 1921, Cat. Planorbidae, p. 47, text 133- LUTESCENS Lamarck 1822. Planorbis lutes-
Fig. 17, and PL 4, Figs. 2 and 7. (Vieques, cens Lamarck 1822, Anim. Sans Vert. 6(2):153-
east of Puerto Rico). Not figured. Locality unknown. Mermod (1952)
figures this species, compares it with P.
122. BOLIVIANUS "Philippi" Dunker 1850. Planor-
guadeloupensis Sowerby, and suggests it comes
bis bolivianus "Philippi" Dunker 1850, Syst.
from the Antilles or northern South America.
p. 60 and 408, PL 10, Figs. 35-37. (Bolivia). 134. NIGRICANS "Spix" Wagner 1827. Planorbis
nigricans "Spix" Wagner 1827, Test, Fluv.
123. CHRISTOPHORENSIS Pilsbry 1934. Australor-
Bras. ... p. 27. As a synonym of P. lugubris
bis glabratuschristophorensis Pilsbry 1934,
Wagner.
Proc. Acad. Nat. Sei. Phila. 86:58, PL 11,
Fig. 12. (Saint Christopher (St. Kitts), British 135. OLIVACEUS "Spix" Wagner 1827. Planorbis
West Indies). olivaceus "Spix" Wagner 1827, Test. Fluv.
Bras p. 26, PI. 18 Fig. 1-2. (Ilheos and
124. CONCAVOSPIRA Anton 1839- Planorbis con-
Almada, Bahia, Brazil).
cavospira Anton 1839, Verz. d. Conch. ., p. 50- .
51. Not figured. (South America). 136. REFULGENS Dunker 1853- Planorbis refulgens
Dunker 1853, Proc. Zool. Soc. London p. 54.
125. CONFUSUS Lutz 1918. Planorbis confusus
Not figured. (St. Domingo). Clessin, 1883, Syst.
Lutz 1918, Mem. Inst. Oswaldo Cruz 10:49-50,
PL 15, Figs. 2a-d. (Rio de Janeiro, Brazil).
p. 106, PL 18, Fig. 10, PI. 17, Fig. 5.
New name for P. ferrugineus d'Orbigny 1837,
Voy. Amer. Me'rid. Lutz thought d'Orbigny had 137. STRIATULUS "Richard" Beck 1837. Planor-
erroneously attached the name P. ferrugineus bis striatulus "Richard" Beck 1837, Index
moll, praes. ., P- 120. Not figured. As a syn-

. .
Group Taphius peregrinus
of
onym of P.guadaloupensis Sowerby with the
d'Orbigny 1835
reference "Guer. Jc vii 1 ?" given. I can
locate nothing in Guerin which would satisfy
148. ARGENTINENSIS Beck 1837. Planorbis pere-
grinus argentinensis Beck 1837, Index moll,
that citation.
praes. . .
., p. 120, as a subspecies of P. pere-
138. VIRIDIS "Spix" Wagner 1827. Planorbis viridis grinus d'Orbigny. Not figured or described.
"Spix" Wagner 1827, Test. Fluv. Bras p. 27. (Pampas (Argentina)).
As a synonym of P. lugubris Wagner.
149. CANONICUS Cousin 1887. P/ö«ori»s canonicus
139. XERAMPELINUS Drouet 1859, Planorbis Cousin 1887, Bull. Soc. Zool. France 12:264,
xerampelinus Drouet 1859, Essai Moll. Terr. PL 4, Fig. 11. (Lake Saint Paolo, near Quito,
Fluv. Guyane Française p. 76, PI. 2, Figs. Ecuador).
27-29- (French Guiana).
150. CENTIMETRALB Lutz 1918. Planorbis centi-
metralis Lutz 1918, Mem. Inst. Oswaldo Cruz
Group of Taphius tenagophilus
10:52-53, PL 17, Figs. 8a-d. (Several locali-
d'Orbigny 1835
ties in Eastern Brazil, and Paraguay).
140. BAHIENSIS Dunker 1850. Planorbis bahiensis 151. CHILENSIS Anton 1839. Planorbis chilensis
Dunker 1850, Syst. Conch. -Cab. Mart, and Anton 1839, Verz. d. Conch , p. 51, Not
Chem., Ed. 2, Planorbis, p. 51, PL 8, Figs. figured. (Chile).
13-18. (Bahia, Brazil).
152. LEVISTRIATUS Preston 1912. Planorbis levi-
striatus Preston 1912, Proc. Mai. Soc. London
141. BIANGULATUS Sowerby 1877. Planorbis bi-
("The Miguelete River,
10:107. Figured in text.
angulatus Sowerby 1877, in Reeve's Conch.
Montevideo"(Uruguay)). Paratypic material ex-
Icon. Vol. 20, Planorbis, PI. 4, Fig. 25- (Bra-
amined at the University of Michigan Museum
zil). Walker, 1918, Synopsis Freshwater Moll.
of Zoology is evidently this species. (UMMZ
N. Amer., p. 95, considered this species a
84088).
synonym of P. antros us Conrad, but other
authors, as Lutz, 1918, considered it Brazilian.
153- LIMAYANUS Beck 1837. Planorbis peregrinus
142. CHEMNITZIANA Beck 1837. Planorbis ten- limayanus Beck 1837, Index moll, praes. ., p. . .
agophilus hemnitziana Beck 1837, Index moll. 120. Not figured. (Lima, Peru). As a subspe-
praes p.
, 120, cites "C IX 1119-20?" cies of P. peregrinus d'Orbigny, without de-
evidently referring to the first edition of the scription. Not P. limayana Lesson 1830.
Syst. Conch. -Cab. of Chemnitz. (Bolivia).
154. MONTANUS Biese 1951. Tropicorbis montanus
143. CLEVEI "Jousseaume" Cousin 1887. Planor-^ Biese 1951, Bol. Mus. Nac. Hist. Nat. (Chile),
bis clevei "Jousseaume" Cousin 1887, Bull. 25:125-126. Fig. 3, p. 130; PL 6, Figs. 13-15.
Soc. Zool. France 12:263, PL 4, Fig. 9- (Rio Hurtado, Samo Alto, Prov. de Coquimbo
(Ecuador). (Chile)). Paratypic material in the University
144. MEGAS Pilsbry 1951. Australorbis bahiensis of Michigan Museum of Zoology (UMMZ 183447)
megas Pilsbry 1951, Nautilus 65=4, PL 9, is evidently this species.
Figs. 4,4a and 5 (of Vol. 64: not named there). 155- PATAGONICUS Beck 1837. Planorbis pere-
(Petropolis, State of Rio de Janeiro, Brazil). grinus patagonicus Beck
Index moll, 1837,
praes...., p. 120. Not figured. (Patagonia).
145. ORBIGNYANA Beck 1837. Planorbis tena-
gophilus orbignyana Beck 1837, Index moll, As a subspecies of P. peregrinus, without
description.
praes...., p. 120. (Argentina). Asa subspecies
of P. tenagophilus, with the citation "d'Orb. 156. PEDRINUS Miller 1879. Planorbis (Taphius)
V xliv 9-12," which is the typical species. pedrinus Miller 1879, Malakozool. Blatt, n.f.,
26:148, PL 7, Figs. 3aA,C. ("Chillo, Rio S.
146. PAYSANDUENSIS Marshall 1930. Planorbis Pedro" (Ecuador)).
paysanduensis Marshall 1930, Proc. U.S. Nat.
Mus. 77(2):4, PL 1, Figs. 1,4 and 6.(Paysandu, 157. PEREGRINUS d'Orbigny 1835. Planorbis pere-
grinus d'Orbigny 1835, Mag. de Zool. V(62):
Uruguay).
26-27. Not figured. ("Habit. Patagonia, Monte-
147. TENAGOPHILUS d'Orbigny 1835. Planorbis video (república Uruguayensi orientali); Pam-
tenagophilus d'Orbigny 1835, Mag. de Zool., pas; provincia Corrientes (república Argentina);
p. 347. Not figured. (Province of Corrientes,

provincia Rio-Grande (república Boliviana) et
Argentina; Province of Santa-Cruz and Chi- provincia Guayaquilensi (república Columbi-
quitos, Bolivia). 1837, Voy. Amér. Mérid., p. ana)"). 1837, Voy. Amér. Merid. p. 348, PL
237, PL 44, Figs. 9-12. 44, Figs. 13-16.
44 H. W. HARRY
158. PUCARAENSIS Preston 1909- Planorbis

araensis Preston 1909, Ann. Mag. Nat. Hist.
- Tristram 1861,
232. Not figured.
Proc.
("Lake
Zool. Soc.
of
London p.
Dueñas" (Guate-
Ser. 8, 3:512, PI. 10, Fig. 15, ("Pucará, Peru, mala)).
at an altitude of 12,500 feet"). Paratypic ma-
170. DUNKERIANUS Clessin 1884. Planorbis dun-
terial in the University of Michigan Museum of
kerianus Clessin 1884, Syst. Conch. -Cab.
Zoology (UMMZ 89735) is evidently this
Mart, and Chem., Ed. 2, Planorbis, p. 122,
species.
PI. 17, Fig. 14. ("Insel Cuba, San Juan").
159- SCHMIERERIANUS Biese 1951. Tropicorbis
schmiererianus Biese 1951, Bol. Mus. Nac.
171. EDENTATUS C.B. Adams 1851. Planorbis
dentiferus edentatus C.B. Adums 1851, Contrib.
Hist. Nat. (Chile) 25:122-124, Fig. 2, p. 130;
Conch. No. 8, p. 132. Not figured. (Jamaica,
PI. 6, Figs. 10-12. ("Salamanca, Rio Choapa"
Hatfield in Westmoreland).
(Chile). Several other localities are given).
Several lots in the University of Michigan Mu- 172. EDENTULA Fischer and Crosse 1880. Planor-
seum of Zoology, identified by Biese from the
localities he cited, are evidently this species. 1880, Miss. Sei. .
bula dentiens var. edentula Fischer and Crosse
Moll. 2:80-81, PL 34,
Figs. 6-6c. ("Colonie Anglaise de Belize").
160. STRAMINEUS Dunker 1848. Planorbis stramin-
eus Dunker 1848, Proc. Zool. Soc. London p. 173. EDENTULUS Clessin 1884. Planorbis edentu-
42. Not figured. (South America). 1850, Syst. lus Clessin 1884, Syst. Conch. -Cab. Mart, and
Conch. -Cab. Mart, and ehem., Ed. 2, Planorbis, Chem., Ed. 2, Planorbis, p. 220-221, PL 33,
p. 42, PI. 5, Figs. 7-9. Fig. 2. ("Central Amerika").
174. GEOSCOPUSPilsbryand Brown 1914. Segmen-

Group of Taphius albicans Pfeiffer tina obstructa geoscopus Pilsbry and Brown
1914, Proc. Acad. Nat. Sei. Phila. p. 431,
161. ALBICANS Pfeiffer 1839. Planorbis albicans
PI. 14, Figs. 4-5. (Antigua).
Pfeiffer 1839, Wiegm. Arch. f. Naturgesch. p.
354. Not figured. (Cuba). 175. INCERTUS Lutz 1918. Planorbis (Taphius)
incertus Lutz 1918, Mem. Inst. Oswaldo Cruz,
162. ARAKANENSIS "Gould" Sowerby 1877. Planor-
10, p. 54, PL 17, Fig. 9a,b,c, and lOd. (Para-
bis arakanensis "Gould" Sowerby 1877 in
hyba and Pernambuco, Brazil).
Reeve's Conch. Icon. Planorbis, PI. J2, No.
100. (Trinidad). 176. INCERTULUS "Lutz" Paraenseand Deslandes
1956. Planorbis incertulus "Lutz" Paraense
163. BERENDTII Tryon 1866. Planorbis (Planor- and Deslandes 1956, Rev. Brasil. Biol. 16
bula) berendtii Tryon 1866, Amer. Jour. Conch.
(l):96(From amuseum label of Lutz's, evident-
2:10, PI. 2, Figs. 14-16. ("Vera Cruz, Mexico; ly for P. incertus Lutz).
Orizaba, Mexico").
177. INSULARUM Pilsbry 1942. Tropicorbis havan-
164. CANNARUM Morelet 1849. Planorbis cannarum
ensis insularum Pilsbry 1942, Nautilus 56:8,
Morelet 1849, Test. Noviss. Ins. Cub. et Amer.
PL 1, Fig. 15. (Grand Cayman Island).
Centr. Pt. 1, p. 16. Not figured. ("H. Belise,
in littore Hondurasano"). 178. JANEIRENSIS Clessin 1884. Planorbis jan-
eirensis Clessin 1884, Syst. Conch. -Cab. Mart,
165. DECLIVIS Tate 1870. Planorbis declivis Tate and Chem., Ed. Planorbis, p. 123, PL
2, 18,
1870. Amer. Jour. Conch. 5:159. Not figured.
Fig. 8. ("Rio Janeiro in Brasilien").
(Acoyapa, Nicaragua). F.C. Baker 1940,
Nautilus 54(3):97, renamed this P. tatei, as 179. NIGRILABRIS Lutz 1918. Planorbis (Taphius)
Tate's name was preoccupied by P. declivis nigrilabris 1918, Mem. Inst. Oswaldo
Lutz
Genth, 1848. Cruz 10:53, PI. 16, Fig. 6a,c,d. (Rio de Jan-
eiro; Bahia and Natal, Brazil).
166. DENTATUS Gould 1844. Planorbis dentatus
Gould 1844, Boston Jour. Nat. Hist. 5:496, 180. PAPARYENSIS F. Baker 1914. Segmentina
PI. 24, Fig. 14. ("small lagoon at San Jorge"
paparyensis F. Baker 1914, Proc. Acad. Nat.
(Cuba)). Sei. Phila., p. 662-663, PI. 26, Figs. 9-11.
("Near the mouth of the main affluent of
167. DENTIENS Morelet 1849. Planorbis dentiens
Papary Lake"(Brazil)).
Morelet 1849, Test. Noviss. Ins. Cub. et Amer.
Centr. Pt. l,p. 18. Not figured. ("H. paludosa 181. PLANULATUS Clessin 1884. Planorbis planu-
circa Belize, in littore Hondurasano"). latus Clessin 1884, Syst. Conch. -Cab. Mart,
and Chem., Ed. 2, Planorbis, p. 163, PL 24,
168. DENTIFERUS C.B. Adams 1845. Planorbis
Fig. 8. (St. Thomas).
dentiferus C.B. Adams 1845, Proc. Boston
Soc. Nat. Hist. 2:17. Not figured. (Jamaica).
182. SHIMEKIF.C. Baker 1945. Trop»cor¿>is shimeki
169- DONBILLI Tristram 1861. Sementina donbilli F.C. Baker 1945, Moll. Fam. Planorbidae,
-
p. 218-219, PI. 134, Figs. 12-14, 28. ("Ome- Chem., Ed. 2, Planorbis. p. 222-223, PI. 33,
tope, Nicaragua"). Páratypes in the U.S. Na- Fig. 7. ("Havanah" (Cuba)).
tional Museum (USNM 534290) show this
194. ORBICULUS Morelet 1849. Planorbis orbiculus
species is P. albicans Pfeiffer. Morelet 1849, Test. Noviss. Ins. Cub. et Amer.
183. SCHRAMM! Crosse 1864, Planorbis schrammt Centr. Pt. 1, p. 17. Not figured. ("H. insulam
Crosse 1864, Jour, de Conchyl. 12:153, PI- 7, Carmen et pagum Palizada yucataneorum").
Fig. 2 (Guadeloupe). "Dunker" Clessin 1883. Planorbis
195. RIISEI
184. STAGNICOLA Morelet 1851- Planorbis stag- riisei "Dunker" Clessin 1883, Syst. Conch.
nicola Morelet 1851, Test. Noviss. Inc. Cub. Cab. Mart, and Chem. Ed. 2, Planorbis, p. 110,
et Amer. Centr. Pt. 2, p. 14-15, Not figured. PI. 17, Fig. 7. ("Jamaica, Puerto Rico").
("H. paludosa littoris cubensis, ad portem 196. STRICTUS Clessin Planorbis strictus
1885.
Bahia-Honda"). Clessin 1885, Syst. Conch. -Cab. Mart, and
185. TATEI F.C. Baker 1940. Tropicorbis tatei Chem., Ed. 2, Planorbis. p. 223, PI- 33, Fig.
F.C. Baker 1940, Nautilus 54:97. New name 4. ("'(Central Amerika?) Coll. Morelet").
for Planorbis declivis Tate 1869, q.v.
197. TEPICENSIS von Martens 1899. Planorbis
tepicensis von Martens
1899, Biol. Centr.
Group of Taphius havanensis
Amer. p. 393, PI. 21, Fig. 14. (N.W. Mexico:
Pfeiffer 1839 Tepic, State of Jalisco).
186. ANODONTA Pilsbry 1919. Planorbula obstructa 198. TERVERIANUS d'Orbigny 1841. Planorbis
anodonta Pilsbry 1919, Proc. Acad. Nat. Sei. terverianus d'Orbigny 1841, in Sagra, Hist.
Phila. p. 219. Not figured. (Reservoir four
,
Cuba, Moll., 1:194-195, PL 13, Figs. 20-23.
miles nocth of Guatemala City). ("environs de la Havane" (Cuba)).
187. DUNKERI F.C. Baker. Tropicorbis orbiculus 199. WEINLANDI Pfeiffer 1876. Planorbis ueinlandi
dunkeri F.C. Baker 1945, Moll. Fam. Planor- Pfeiffer 1876, Malakozool. Blatt. 23:172, 232,
bidae, p. 494 (Explanation of Plate 129). PL PL 2, Figs. 9-11, (Mountain streams near
129, Figs. 26-31, 32-36. (Dry pool near Tam- Jeremie, Haiti).
pico, Mexico, for and "Los
Figs. 26-31,
Canoas, Mexico" for Figs. 32-36). Proposed Group of Taphius helophilus
merely as a "new name," with no mention of d'Orbigny 1835
what it was to replace. Probably meant to re-
place P. haldemani Dunker, q.v.
200. ATACAMENSIS Biese 195 1. Tro/)ícor¿>»s ataca-
mensis Biese 1951, Bol. Mus. Nac. Hist. Nat.
188. HALDEMANIDunker 1850. P/íi«or¿>/s haldemani (Chile) 25:126-127, Fig. 4, page I3O; PL 6,
Dunker 1850, Syst. Conch. -Cab. Mart, and Figs. 16-18. ("Holotipo: Rio Copiapo. Copiapo
Chem., Ed. 2, Planorbis. p. 59, PL 10, Figs. (Canal Ojances) Prov. de Atacama, 370 m. de
38-40. (Mexico). Not P. haldemani C.B. Adams Atacama, 370 m. de altura" (Chile)).
1849.
201. HELOPHILUS d'Orbigny 1835. Planorbis helo-
189. HAVANENSIS Pfeiffer 1839. Planorbis havan- philus d'Orbigny 1835, Mag. de Zool. V(62):27.
ensis Pfeiffer 1839, Wiegm. Arch. F. Na- Not figured. ("Habit, provincia Limacensi
turgesch. p. 354. Not figured. (Havana, Cuba). (república Peruviana)"). 1837, Voy. Amér. Mérid.
p. 349, PL 45, Figs. 13-16.
190. LIEBMANNI Dunker 1850. Planorbis liebmanni
Dunker 1850, Syst. Conch. -Cab. Mart, and 202. INFLEXUS Paraense and Deslandes 1956.
Chem., Ed. 2, Planorbis, p. 59, PI- 10, Figs. Australorhis inflexus Paraense and Deslandes
32-34. ("Vera Cruz"(Mexico)).
1956, Rev. Brasil. Biol. 16:149-158, text Figs.
191. MICROMPHALUS "Dunker" Strebel 1873. 1-7. ("Type locality; Pouso Alegre, State of
Planorbis micromphalus "Dunker" Strebel
1873, Beitr. z. Kenntnis d. Fauna .
Land-u.
SÜSSW. -Conch, p. 47. Not figured. (No locality).
203.
Minas Gerais, Brazil").
URUGUAYENSIS
uruguayensis
Preston 1912. Planorbis
Preston 1912, Proc. Mai. Soc.
Said by Strebel to be a juvenile of P. haldemani London 10:107. Figured in text. ("Montevideo"
Dunker. (Uruguay)).
192. OBSTRUCTUS Morelet 1849. Planorbis ob-
structus Morelet 1849, Test. Noviss. Ins. Group of Taphius pallidus
Cub. et Amer. Centr. Pt. 1, p. 17. Not figured. C. B. Adams 1846
("H. insulam Carmen"). 204. DECIPIENS C.B. Adams 1849. Planorbis de-
193. OBVOLUTUS Clessin 1884. Planorbis obvolu- cipiens C.B. Adams 1849, Contrib. Conch.
tus Clessin 1884, Syst. Conch. -Cab. Mart, and No. 3, pp. 43-44. Not figured. (Jamaica).
-
46 H. W. HARRY
205. ISTHMICUS Pilsbry 1920. Planorbis isthmicus 1863, Proc. Acad. Nat. Sei. Phila., p. 146, PI.
Pilsbry 1920, Nautilus 33:78-79, Text figure. 1, Figs. 4-5- ("Panama").
(Panama City). 214. FILOCINCTUS Pilsbry and Ferriss 1906. P/önor-

206. MAYA Morelet 1869- Planorbis maya Morelet bis filocinctus Pilsbry and Ferriss 1906, Proc.
1849, Test. Noviss. Ins. Cub. et Amer. Centr. Acad. Nat. Sei. Phila., p. 165, PI- 9, Figs.
Pt. 1, p. 16. Not figured. ("H. cisternae 1-3- (San Pedro River, Benson, Arizona, in
civitatis Campeche" (Mexico)). drift debris). Possibly a synonym of one of the

Neotropical species.
207. PALLIDUS C.B. Adams 1846. Planorbis pal-
lidas C.B. Adams
1846, Proc. Boston Soc. Nat. 215. FUSCUS Dunker 1848. P/««orèis fuscus Dunker
Hist. 2:102. Not figured. (Jamaica). 1848, Proc. Zool. Soc. London p. 42. Not
figured. (Valparaiso (Chile)). 1850, Syst.
SPECIES INCERTAE SEDIS Conch. -Cab. Mart, and Chem., Ed. 2, Planorbis,
p. 52, PI. 8, Figs. 19-21.
The species listed here are placed in this cate-
gory for several reasons. In some instances I have 216. GRACILENTUS Gould 1855. Planorbis gracil-
not been able to recognize them from the descrip- entus Gould 1855, Proc. Boston Soc. Nat. Hist.
tions. In other cases the species seem quite dis- 5:129. Not figured. ("Great Colorado Desert
tinct, conchologically (e.g., Promenetus minutus lowlands" (U.S.A.)). Binney, 1865, Land and
Taylor), yet I am uncertain of the genus in which it fresh water shells of N. Amer., Pt. 2, p. 108,
should be placed. It is quite possible that additional considered this a synonym of P. liebmanni
genera will be discovered in the area, which are not Dunker, after examining authentic specimens
yet recognized. received from Gould. Other writers have con-
There are among these species several which are sidered the two species distinct.
smaller than any Neotropical species for which the 217. GUNDLACHI "Dunker" Clessin 1884. Planor-
anatomy is known, except AcrorAi's /)/«/
Odhner. bis gundlachi "Dunker" Clessin 1884, Syst.
This group includes P. hindsianus Dunker, P. Conch. -Cab. Mart, and Chem., Ed. 2, Planorbis,
panamensis Dunker, P. pfeifferi Strobe!, P. pfeifferi p. 146, PI. 17, Fig. 8. ("Die Insel Trinidad").
mendozanus Strobel, P. boetzkesi Miller and Pro-
218. HINDSIANUS Dunker 1848. Planorbis hindsianus
menetus minutus Taylor. Such material is very rare
Dunker 1848, Proc. Zool. Soc. London 16:41.
in the museum collections which I have studied.
Not figured. ("In insulam Puna in sinu ad
These species have probably been overlooked in the
Guayaquil" (Ecuador?)).
field.
Morelet 1851. Planorbis

219. HONDURASENSiS Clessin 1884. Planorbis
208. AERUGINOSUS hondurasensis Clessin 1884, Syst. Conch. -Cab.
aeruginosus Morelet 1851, Test. Noviss. Ins.
Mart, and Chem., Ed. 2, Planorbis, p. 164, PI.
Cub. etAmer. Centr. Pt. 2, p. 15- Not figured.
24, Fig. 2. ("Honduras bei Sta. Maria").
("H. paludosa circa lacum yzabalensem"
(Guatemala)). 220. JACOBEANUS "Valenciennes" Hupe 1854.
Planorbis jacobeanus "Valenciennes" Hupe
209- BOETZKESI Miller 1879. Planorbis (Gyraulus)
1854, Moluscos, Gay's Hist. Fisica y Politica
boetzkesi Miller 1879, Malakozool. Blatt, n.f.
de Chile, p. 124. Not figured. ("Estanques de
26:148, PI. 7, Figs. 4a-A-C. ("Chillo, Rio S.
Santiago" (Chile)).
Pedro cum praecedenti, crebrior" (Ecuador)).
221. KUHNERIANUS "Dunker" Clessin 1883.
210. BOUCARDIANUS Preston 1907. Planorbis
Planorbis kuhnerianus "Dunker"
Clessin
boucardianus Preston 1907, Ann. Mag. Nat.
1883, Syst. Conch. -Cab. Mart, and Chem., Ed.
Hist., Ser. 7, 20:497, text Fig. 17, p. 493
2, Planorbis. p. 108, PI. 11, Fig. 12. (Surinam),
(Mexico).
222. KUHNL\NA "Dunker" Clessin 1886. Planorbis
211. CIRCULARIS Clessin 1884. Planorbis circu- kuhniana "Dunker" Clessin 1886, Syst. Conch.
laris Clessin 1884, Syst. Conch. -Cab. Mart, Cab. Mart, and Chem., Ed. 2, Planorbis, p. 413
and Chem., Ed. 2, Planorbis. p. 221, PI. 33, and errata. A proposed substitute for P. kuhner-
Fig. 3- ("Central Amerika?"). ianus "Dunker" Clessin 1883-
212. DENTIFER Morch 1833- Planorbis dentifer
Morch 1833, Jour. Petit., No. 1, p. 37 (Bahia, 223. MENDOZANUS Strobel 1874. P/a«or¿)ís (Gyraul-
us) pfeifferi Strobel, var. mendozanus Strobel
Brazil). I have not been able to locate this
1874, Mat. Mai. Argent, p. 39-40, PI. 2, Fig.
reference. It is possibly a nude name, intro-
3, ("ban Carlos e Manantial de la Pirca nella
duced by Hupe (1854, p. 61), who gave only the
Sierra de Mendoza" (Argentina)).
information cited here. Not P. dentifer Moricand
1853. 224. MERIDAENSIS Preston 1907. Planorbis meri-
213. FIELDII Tryon 1863- Planorbis fieldii Tryon daensis Preston 1907, Ann. Mag. Nat. Hist.,
Ser. 7, 20:497, Fig. 18 (p. 493). (Merida, lineatus sonorensis J.G. Cooper 1893, Proc.
Yucatan). Calif. Acad. Sei., Ser. 2, 3:343, PI. 14. Figs.
lOa-d. (Near San Miguel, Sonora, Mexico). This
225. MEXICANUS "Dunker" Clessin 1886. Planor-
may be a synonym of P. salleanus Dunker.
bis mexicanus "Dunker" Clessin 1886, Syst.
Pilsbry(1948, Land Moll. N. Amer., 2:631-632)
Conch. -Cab. Mart. andChem., Ed. 2, Planorbis,
reproduced the original figure, noted that the
p. 408, PI. 12, Figs. 1-3. ("Mexico? (nach der
type was lost, and suggested it belonged to the
Benennung der Art.)").
Planorbidae.
226. MINUTUS Taylor 1954. Promenetus minutus
Taylor 1954 Revista Soc. Mal. "Carlos de la
238. THERMALUS Biese 1951- Taphius thermalus
Biese 1951, Bol. Mus. Nac. de Hist. Nat.
Torre," Havana, 9:37-38. Not figured. ("Allee
(Chile), 25:118-119, Fig. 6, p. 130; PI. 6,
Stream, opposite laboratory, Barro Colorado
p. 130; PI. 6, Figs. 4-6. ("Holotipo: Ojos de
Id., Gatun L., Panama Canal Zone").
Ascotan, Salar Ascotan" (Chile)).
227. MORELETIANUS Clessin 1884- Planorbis
239. UMBILICATUS Anton 1839. Planorbis umbili-
moreletianus Clessin 1884, Syst. Conch. -Cab.
catus Anton 1839, Verz. à. Conch...., p. 51.
Mart, and Chem., Ed. 2, Planorbis, p. 162, PI.
Not figured. (Chile). Not P. umbilicatus MUller
24, Fig. 1. (La Guayara, Venezuela).
1777. Biese, 1951, suggested Anton's name be
228. PANAMENSIS Dunker 1848. Planorbis panamen- suppressed, since he found nothing in Chile
sis Dunker 1848, Proc. Zool. Soc. London 16: to fit it.
41. Not figured. ("Hab. in rivulis Panama

(H. Cuming)").
NOMINA NUDA
229. PERFORATUS Anton 1839- Planorbis perfora-
tus Anton 1839, Verz. d. Conch...., p. 51. The matter of definition of this category is debat-
Not figured. ("Lima" (Peru)). able.Some students hold if anything
the view that
230. PERUVL\NUS "Mühl." Anton 1839- Planorbis at all is said about a species in naming it as new,
peruvianas "Mühl." Anton 1839, Verz. d. that name is not nude. Yet much may be said, and
Conch. . p. 50. Not figured. As a synonym of that species still not be recognizable. In the pres-
. .,
P. neglectus, with no author cited for the latter. ent list names are considered nude when nothing
(Malabrya in Peru). more was said about them than a mere listing of a
locality, or when only the name itself indicated the
231. PETENENSIS Morelet 1851- Planorbis petenen-
proposed species came from the Neotropics.
sis Morelet 1851, Test. Noviss. Ins. Cub. et
Amer. Centr. Pt. 2, p. 15- Not figured. ("H. 240. BRAZILIANUS Jay 1836. Planorbis brazilianus
lacum ytza Petenensium" (Guatemala)). Jay 1836, Catalogue Ed. 2, p. 77. No descrip-
tion, figure, citation or locality.
232. PFEIFFERI Strobel 1874. Planorbis pfeifferi
Strobel 1874, Mat. Mai. Argent, p. 39-40, PI- 241. CAPILLARIS Beck 1837. Planorbis capillaris
Beck 1837, Index moll, praes. p. 119- No
2, Fig. 2. ("Belgrano e Tigre presso Buenos
. . .,
Aires" (Argentina)). Not Planorbis pfeifferi citation, description or figure. (Mexico).
Krauss 1848, Sndafrik. Moll. p. 79. 242. COSTATUS "Férussac" Beck 1837. Planorbis
233. PHILIPPIANUS Dunker 1848. Planorbis philip- costatus "Fe'russac" Beck 1837 Index moll,
pianus Dunker 1848, Proc. Zool. Soc. London praes...., p. 120. No citation, description or
figure. (Brazil).
p. 43, Not figured. (Cochabamba, Bolivia).
1850, Syst. Conch. -Cab. Mart, and Chem, Ed. 243- CUMINGII Beck 1837. Planorbis cumingii
2, Planorbis, p. 47, PI. 5, Figs. 16-18. Beck 1837, Index moll, praes , p. 120. No
234. PLANUS CLESSIN 1884. Planorbis planus citation, description or figure. (Chile). Germain,
Clessin 1884, Syst. Conch. -Cab. Mart, and 1921, Cat. Planorbidae, p. 45, listed this as
Chem., Ed. 2, Planorbis, p. 222, PI. 33, Fig. synonym of P. chilensis Anton 1839-
6. ("?(Central-Amerika, Coll. Morelet)"). 244. LINNAEUS "Moricand" Hupe 1857. Planorbis
235- RAIMONDI Philippi 1869. Planorbis raimondi linnaeus "Moricand" Hupe 1857, Mollusques,
Philippi 1869, Malakazool. Blatt. 16:38-39. Not in Castelnau's Animaux. de l'Amérique du
. .
figured. ("In rivulis nemorum loco Peruviae Sud, p. 62. No citation, description or figure.
dicto 'Pampa del Sacramento' lectus"). (Brazil).
245. LUNDII Beck 1837. Planorbis lundiiBecV. 1837,

236. RETUSUS Morelet 1849. Planorbis retusus
Index moll. praes p. 120.,
No citation,
Morelet 1849. Test. Noviss. Ins. Cub. et Amer.
description or figure. (Brazil).
Centr. Pt. 1, p. 17. Not figured. ("H. insulam
Carmen"). 246. MINUTULUS Beck 1837. Planorbis minutulus
SONORENSIS Beck 1837, Index moll. praes p. 120. No
237. J.G. Cooper 1893- Helicodiscus ,
,
48 H. W. HARRY
citation, description or figure. (Antilles). amphiglyptus Pilsbry 1951, Nautilus 65:3, PI.
9, Figs. 6, 6a (of Vol. 64, where they were
247. MORICANDI Beck 1837. Planorbis moricandi
not named). ("Rio de Janeiro, Brazil" with
Beck 1837, Index moll. praes. ., p. 120. No . .
doubt). This is probably the juvenile of some

citation, description or figure. ( Bahia (Brazil)).
land snail.
248. REVENTLOWI Beck 1857. Planorbis reventloui
Beck 1837, Index moll. praes. . . ., p. 120. No
citation, description or figure. (Rio Janeiro BffiLIOGRAPHY
(Brazil)).
249. ROTA Beck 1837. Planorbis rota Beck 1937, ADAMS, C. ., 1845, Specierum novarum con-
Index moll. praes...., p. 120. No citation,
chyliorum in Jamaica repertorum, synopsis.
description or figure. (Peru).
Proc. Boston Soc. Nat. Hist. 2:1-28.
1846, (Minutes of the meeting of 18
,
250. SIMPLEX Beck 1837. Planorbis simplex Beck February 1846). Proc. Boston Soc. Nat.
1837, Index moll. praes. . . ., p. 120. No cita- Hist. 2:102-103.
tion, description or figure. (Mexico). 1849, Descriptions of supposed new
,
species of freshwater shells which inhabit

Jamaica. Contrib. to Conchol. No. 3, pp.
EXTRALIMITAL SPECIES 42-45.
1851, Descriptions of new varieties
,
Besides the names listed here, several African of shells, which inhabit Jamaica. Contrib.
to Conchol. No. 8, pp. 129-140.
species have been cited from the Neotropics. These
are omitted from the present catalogue. For a dis-
ADAMS, H. and A.ADAMS, 1853-1858 .The genera
of recent Mollusca. London. 3 Vols. (Plan-
cussion of the question of the conspecificity of the
orbidae in Vol. 2),
African and Neotropical planorbids, see Paraense
and Deslandes (1956a).
AGUAYO, . G., 1933, On the synonymy and dis-
tribution of Planorbis- anatinns 6.^ Orbigny.
251. CORNEUS Linné 1758. Helix cornea Linné Nautilus 47:64-68.
1758, Syst. Naturae Ed. 10, p. 770. This Eu- 1935, Esplcilegio de moluscos
,
ropean species, now placed in the genus Cubanos. Mem. Soc. Cubana Hist. Nat. 9:
Planorbarius, has been cited from Cuba 107-128
(Aguayo, 1935, Mem. Soc. Cub. Hist. Nat. , 1938, Los moluscos fluviátiles
9:120, as Planorbis) and Puerto Rico (various Cubanos. Mem. Soc. Cubana Hist, Nat. 12:
papers on parasitology by Hoffmann). These 203-242.
records were probably based on specimens of ANTON, H. E., 1839, Verzeichniss der Conchy-
Helisoma. There is no convincing account of lienwelche sich in der Sammlung von Herman
Planorbarius corneus L. from anywhere in the Eduard Anton befinden. Halle, i-xii, 1-110.
New World. BAKER, F. C, 1940, A new species of öre/)ano-
trema and some preoccupied planorbid
252. JAMAICENSIS "Bolten" Roding 1798. Planor-
names. Nautilus 54:96-97.
bis jamaicensis "Bolten" Roding 1798, Mus.
1945, The molluscan family Plan-
,
Boltenianum, p. 73- Refers to Gmelin, Chem-
orbidae. Univ. Illinois Press, Urbana, xxxvi,
nitz and Seba. (Jamaica). This species is the
1-530, Pis. 1-141,
first one placed in the genus Planorbis, to be BAKER, Fred, 1914, The land and freshwater
cited from the Neotropics. It is a member of mollusks of the Stanford Expedition to Brazil.
the Pleurodontidae. Proc. Acad. Nat. Sei. Phila. (1913) 618-672,
253.. PARVUS Say 1817. Planorbis parvus Say 1817, Pis. 21-27.
Nicholson's Encyclopedia Ed. 1, no pagination. BAKER, H. ., 1930, The mollusca collected by
PI. 1, Fig. 5- (Delaware (U.S.A.)). Cited from the University of Michigan— Williamson Elx-
Mexico byPilsbry (1891, Proc. Acad. Nat. Sei. pedition in Venezuela. Occ, Pap. Mus. Zool.
Phila., p. 322), but von Martens (1899, Biol.
Univ. Mich. No. 210:1-94,
Centr. Amer., Moll., p. 394) notes it is strange
1947, Clessin' s section of Ptoworöfs
,
armigeriis. Nautilus 61:71-72.

that noone else has found it in Mexico. Aguayo
BARTSCH, P., 1908, Notes on the freshwater
(1938, Mol. Fluv. Cubanos) listed it from Cuba,
with no specific location or comment. I have
moUusk Planorbis magnificus and descrip-
tions of two new forms of the same genus
seen a small lot from Chihuahua (Mexico) and
from the Southern States, Proc. U.S. Na-
two from Cuba, both in the U.S. National Mu-
tional Mus. 33:697-800.
seum, but the occurrence of this species in
BECK, H., 1837, Index molluscorum praesentis
the Neotropics needs confirmation.
aevi musei principis augustissimi Christiani
254. AMPHIGLYPTUS Pilsbry 1951. Australorhis Frederici. Hafniae, 1-24.
BENTHEM JUTTING, V. S. S. van, 1943, Ueber DROUET, H., 1859, Essai sur les mollusques
eine Sammlung nichtmariner Mollusken aus terrestres et fluviátiles de la Guyane Fran-
dem niederschlagsarmen Gebiete Nordost- çais, Mém, Soc, Acad, de l'Aube 23:7-116,
Brasiliens, Arch, Hydrobiol, Stuttgart. 39: Pis. 1-4.
458-489, .
DUNKER, G., 1848, Diagnoses specierumnovar-
BEQUAERT, J. and W. J. CLENCH, 1939, The um generis Planorbis collectionis Cumin-
genus Plesiophysa. J. Conchol, 21:175-178. gianae. Proc. Zool. Soc. London 16:40-43.
BIESE, W. A., 1951, Revision de los moluscos 1841-1850, Ptoworbes (in part). In
,
terestres y de agua dulce provistos de Systematisches Conchylien- Cabinet of Mar-

concha de Chile. Parte 4: Planorbidae. Bol. tini and Chemnitz, Ed. 2, Vol. 1, Pt. 17.
del Mus. Nac, de Hist. Nat. (Chile) 25:115- Pages 1-62 and Plates 1-10 and 16 were pub-
137. lished by Dunker. See Clessin.
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connues, déscrites par M. Bonnet. Rev. et Sexual apparatus of selected planorbid snails
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BRODERIP, W. J., 1832, (Minutes of the meet- somiasis. Bol. de la Ofic. San. Panameri-
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of land and freshwater mollusks of Antigua. du Mexique et du Guatemala, In Mission
Proc. Acad. Nat. Sei. Phila. 429-431. Scientifique au Mexique et dans l'Amérique
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Museum. Brit, Mus., London, viii, 1-552. GERMAIN, L., 1921-1924, Catalogue of the
(The pages containing the description of Planorbidae in the Indian Museum (Natural
Planorbis tuynens appeared in 1856). History), Calcutta. Rec, Indian Mus. 22:1-
CHITTY, E,, 1853, Descriptions of thirty sup- 210. Pis, 1-4,
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No, 5. Proc. New England Zool. Club 13: J. Nat. Hist. 4:485-498.
35-38. , 1855,
New species of land and fresh-
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MORICAND, S., 1839, Premier supplement au Acad. Nat. Sei. Phila. p. 561-565.
, 1919, Mollusca from Central Amer- Gesellsch. Bern. p. 33-56 and 89-103.
ica and Mexico. Proc. Acad. Nat. Sei. SMITH, E. 1890, Mollusca.
A., In Ridley,
Phila. 212-223, PI. 11. H. N.: Notes on the zoology of Fernando
, 1920, Mollusks from Lake Chápala, Noronha, J. linn. Soc. London, 20:473-570.
State of Jalisco and vicinity. Proc. Acad. SOWERBY, G. ., 1822-1834, The genera of
Nat. Sei. Phila., 192-194. recent and fossil shells. Stirling, London.
1920a, Some Auriculidae and Plan-
, A large number of plates and accompanying
orbidae from Panama. Nautilus 30:76-79. text without pagination. Part 4, Planorbis,
1923, (No title: description oiPlan-
,
dates from 1822. For dates, see Sherborn,
orbis caloderma, sp. n.) Nautilus 36:143- O. D, in Ann. Mag. Nat. Hist. 1894, Ser. 6,
144, PI. 115. 13:371.
, 1924, South American land and 1877, Monograph of the genus
.
fresh-water mollusks. Notes and descrip- Platwrbis. In Reeve's Conchologia Iconica

tions. Proc. Acad. Nat. Sei. Phila. 76:49-66. Vol. 20. 14 plates and accompanying pages
,1934, Review of the Planorbidae of (unnumbered) of description,
Florida, with notes on other members of the STREBEL, H,, 1873-1882, Beitrag zur Kenntniss
family. Proc. Acad. Nat. Sei. Phila. 86:29- der Fauna mexikanischer Land- und Suss-
66. wasser- Conchy lien. Abh. a. d. Gebiete d.
, 1942, Land mollusca of the Cayman Naturwiss. Naturwiss.
, Verein Hamburg,
Islands collected by the Oxford University Bd. 6, Abth. 1, 69 plates. Planorbidae,
Biological Expedition. Nautilus 56:1-9, PI. 1. pages 39-47, appeared in 1873.
, 1951, Notes on some Brazilian STROBEL, P., 1874, Materiali per una mala-
Planorbidae. Nautilus 65:306, and PL 9 costatica di terra e di acqua dolce dell'
VoL 64 (1950). Argentina Méridionale. Pisa. Ixxx, 1-142,
PILSBRY, H. A. and J. H. FERRISS, 1906, Mol- Pis. 1-2.
lusca of the Southwestern States II. Proc. SWAINSON, W., 1840, A treatise on malacology,
Acad. Nat. Sei. Phila. 429-431. London 1-419,
viii,
PRESTON, H. ., 1907, Descriptions of new TATE, R,, 1870, On the land and freshwater
species of land and freshwater shells from mollusca of Nicaragua. Amer. J. Conchol.
Central and South America. Ann. Mag. Nat, 5:151-162.
Hist. Ser. 7, 20:490-498. TAYLOR, D. W., 1954, A new Promete iz<s (Plan-
, 1909, New land, freshwater and orbidae) from Panama. Rev. Soc. Mala-
marine shells from South America. Ann. cologica "Carlos de la Torre" 9:37-38.
Mag. Nat. Hist. Ser. 8, 3:507-513, PL 10. TRISTRAM, H. ., 1861, Catalogue of a collec-
, 1912, Characters of three new tion of terrestrial and fluviatile mollusks,
epecies of freshwater shells from Uruguay. made by O. Salvin, Esq. in Guatemala. Proc.
Proc. MalacoL Soc. London 10:107. Zool. Soc. London, p. 229-233.
RÖDING, P. F., 1798, Museum Boltenianum. TRYON, G. W, Jr., 1863, Description of two new
Hamburg, viii, 1-199. species of fresh water mollusca from Pan-
SAY, T., 1816, Conchology. In Nicholson's ama. Proc. Acad. Nat. Sei. Phila. p. 146.
British Encyclopedia, First American Edi- ,1866, Descriptions of new exotic
tion Vol. 4. freshwater mollusca. Amer. J. Conchol. 2:
, 1818, Account of two new genera 8-11.
and several new species of fresh water and WAGNER, 1827, Testacea fluviatilia quae
J. A.,
land shells. J. Acad. Nat. Sei. Phila. p. 276- in itinereper Brasiliam annis 1817-1820
284. jussu et ... Munich, C, Wolf,
_,1830-1834, American conchology. WALKER, B,, 1918, A synopsis of the classifi-
New Harmony, Indiana. Series of plates and cation of the fresh-water mollusca of North
accompanying text without pagination. Plan- America, Mise, Publ, Mus, Zool,, Univ,
orbis lentus was published in 1834. Mich, No. 6:1-213.
SHUTTLEWORTH, R. J., 1854, Diagnosen Neuer WEIN LAND, D. F., 1876, Diagnosis molluscorum
Mollusken. No. 7. Beiträge zur näheren Haitiensium, Malakozool. Blatt. 23:170-174.
Kenntniss der Land- und Süsswasser- Mol- (contains Planorbis weinlandi, described by
lusken der Insel Portorico. Mitthl. Naturf. Pfleffer).
52 H. W. HARRY
Z US AMME N FASSUNG
EIN KRITISCHES VERZEICHNIS DER NOMINELLEN GATTUNGEN UND

ARTEN NEOTROPISCHER PLANORBIDAE
Die Verbreitung der neotropischen Planorbidengattungen reicht nach Norden
hin mit nur wenigen Arten bis in die südlichen Vereinigten Staaten von Nordamerika.
Von den 11 nominellen Gattungen deren typische Arten aus den Tropen der neuen
Welt stammen, werden hier nur 3 als gültig anerkannt: Taphius, Dre trema und
aber vorderhand absichtlich unbenannt. Obwohl eine nearktische ^,

Acrorbis. Eine weitere noch namenlose Grattung wurde zwar auch erkannt, bleibt
sich Zugebenermassen südwärts bis nach Zentralamerika hin erstreckt, muss

Helisoma,
jedoch betont werden, dass die Angaben über das Vorkommen anderer nearktischer
oder paläarktischer Arten in den Neuweltstropen jeglicher anatomischer Beweise
entbehren. Einige neotropische und afrikanische Gattungen sind zwar vermutlich
identisch; da jedoch die erstem in jedem Fall den letzteren gegenüber Priorität
zu haben scheinen, wird darauf nicht näher eingegangen.
Der vorliegende Katalog stellt einen Versuch dar, sowohl den locus typicus
wie auch die ursprünglichen diesbezüglichen Schriften sämtlicher jemals benannten
planorbiden Arten aus der Neotropen zusammenzufassen. Von diesen Arten wurden
207 in 21 "Arten- Gruppen" zusammengefasst, deren Mitglieder wohl synonym sein
dürften. Weitere 32 Arten wurden unter incertae sedis angeführt; diese scheinen
sich, zumindest teilweise, von den 21 Arten-Gruppen zu unterscheiden, können
aber, mangels anatomischer Angaben, keiner Gattung mit Bestimmtheit zugeteilt
werden. Schliesslich sind noch 11 nomina nuda sowie 4 extralimitale Arten ver-
zeichnet. Das Schrifttum ist weitgehend angegeben.
RESUME
UN CATALOGUE CRITIQUE DES GENRES PLANORBES

NÉOTROPICAUX NOMINAUX ET DE LEURS ESPÈCES
Vers le nord, les genres planorbes néotropicaux ne pénètrent que par quelques
espèces dans les régions méridionales des États Unis de l'Amérique du Nord. Des
11 genres nominaux dont les espèces-types proviennent des tropiques du nouveau
monde, seulement 3 sont considérés valides: Taphius, Drepanotrenia et Acrorbis.
Un genre additionel, point nommé encore et laissé ici délibérément innommé, a
été également reconnu. Quoique le genre néarctique Helisonia s'étend, sans aucun
doute, jusqu'en Amérique Centrale, il n'enest pas de même pour les autres genres
néarctiques ou paléarctiques dont la présence a été signalée dans les régions néo-
tropicales et il faut souligner que ces rapports sont dépourvus de toute corrobora-
tion anatomique. Remarquons aussi que certains genres néotropicaux et africains
sont probablement identiques; mais, comme dans tous les cas connus les premiers
semblent avoir precedence sur les derniers, la question n'a pas été approfondie.
Ce catalogue représente un essai d'assembler les références originales,
aussi bien que les localités-types, de toutes les espèces planorbides néotropicales
nommées. De ces espèces, 207 ont été divisées en 21 "groupes d'espèces" dont
les membres sont probablement synonymes; 32 autres espèces sont citées comme
incertae sedis ; elles semblent, partiellement du moins, se différentier des 21
groupes d'espèces, mais ne peuvent être assignées à aucun genre faute de détails
anatomiques. Finalement sont cités 11 no>ni)m nuda et 4 espèces extralimitales.
Une bibliographie étendue est donnée.
RESEÑA
UN CATALOGO CRITICO DE GENEROS NOMINALES Y DE ESPECIES DE
PLANORBIDAE NEOTROPICALES
Los géneros de Planorbidae Neotropicales tienen solamente unas pocas
especies septentrionales en la región Sur de los Estados Unidos de Norte América.
De los once géneros nominales que se basan en especies de tipos Neotropicales,
solamente 3 son considerados válidos: Taphiiis, D re ()a>io trema, y Acrorbis.
También se reconocen como válidos un género que se han dejado sin nombre
deliberadamente. El género Neartico Helisoma se extiende tan al Sur como Centro
América, mientras que la mención de otros géneros Nearticos y Palearticos que
llegan de la reglón Neotropical no está confirmada por datos anatómicos. A
pesar de que algunos géneros Neotropicales y Africanos son idénticos, el género
Neotropical parece tener prioridad en todos los casos conocidos, aunque no se
haya investigado extensamente la cuestión.
Hemos tratado de mencionar referencias originales y localidades típicas de
todas las especies nominales de los planorbidos de los Neotrópicos. Entre ellos,
207 se clasifican en 21 "grupos-especies". Los miembros de cada grupo son
sinónimos probablemente. Treintaidos especies están agrupadas enmcertae sedis.
Algunas de estas parecen diferenciarse de los 21 grupos de especies, pero la falta
de datos anatómicos no nos permite agruparlos en géneros. Once nomina mida y
cuatro especies extra son mencionadas lo mismo que se proporciona una amplia
bibliografía.
PLANORBIDAE
, .,
,
, -
; , .
, 11
,
Acrorbis.
-, 3: Taphius,
,-
Drepatwtrema
-
,
Helisoma
,,1.„,
, , , -
-
--
-
.
. . --,
-,
, . , 21 "; „"
„"
32
11
207
nomina muía
,
.
-
11, 1(1): 55-72
CYTOTAXONOMIC STUDIES OF FRESHWATER LIMPETS

(GASTROPODA: BASOMMATOPHORA)
I. THE EUROPEAN LAKE LIMPET, ACROLOXUS LACUSTRIS^
by J. B. Burch^
ABSTRACT
Acroloxus lacustris (Linnaeus) is a freshwater limpet common to Europe,
northern Asia and Caucasia. It has nearly always been assigned to the basom-
matophoran family Ancylidae, and hence is generally regarded as one of the most
specialized and phylogenetically advanced basommatophorans.
It is shown in this paper that in regard to certain details oí cytology, .
lacustris should not be considered closely related to other Ancylidae, but rather
placed in a family by itself, the Acroloxidae, a conclusion corroborated by other
authors on morphological grounds. Indeed, the various cytological differences
would tend to further separate Acroloxus from other Basommatophora. The dif-
ferences observed consist in the large size of the various cells of spermatogenesis,
the greater volume ratio of chromatin to cytoplasm, the relatively large size of the
chromosomes and the morphology of the mature sperm, whose heads are long and
thread-like, not bullet- or turnip- shaped like those found in other basommatophor
an snails. In addition, the chromosome number (n=18), although characteristic of
the Basommatophora in general, is different from that found in other freshwater
limpets (x or basic haploid number -15 in the Ancylinae-Ferrissiinae; n=17 in
the Laevapecinae).
The mitotic chromosomes of A. lacustris are metacentric as characteristic
of all Basommatophora; 6 pairs (including the 2 largest and the smallest) are
medianly constricted; the other 12 pairs are submedianly or subterminally con-
stricted. This is the first time the caryotype of any Euthyneuran snail has been
accurately determined and figured.
The phylogenetic position of the Acroloxidae may be close to the base of the
Basommatophora as suggested by Bondesen and Hubendick, but at the present state
of knowledge the evidence which would tend to support such a conclusion can not be
found in details of cytology. But, contrary to earlier views reached on purely ana-
tomical grounds, the position of the Ancylidae, as determined by their chromosome
numbers, should also be near the base of the Basommatophora, but not close to the
Acroloxidae because of the other cytological differences.
INTRODUCTION Most moUuscan systematists have

placed AcroZoxMS in the basommatophoran
Acroloxus lacustris (Linnaeus) is a family Ancylidae which originally contain-
freshwater limpet common to Europe, ed all freshwater limpets. However, Bon-
northern Asia and Caucasia. It is often deson (1950) and Burch (1961b) have con-
called the "lake limpet" because of its tended that the differences found between
preference for a lentic environment in Acroloxus and other ancylids in regard
contrast to the other common freshwater to egg- capsule morphology and spermato-
limpet of its region, Ancylus fluviatilis genesis are great enough to warrant its
Müller^, which inhabits rivers. separation as a distinct family. These
authors were not the first to appreciate
•'•This investigation was supported (in part) by

a research grant, 2E-41, from the National
Institute of Allergy and Infectious Diseases, "^Generic designations given here are in accord
U. S. Public Health Service. with Opinion 363 of the International Commis-
2 Museum and Department of Zoology, Univer- sion on Zoological Nomenclature (Hubendick,
sity of Michigan, Ann Arbor, Michigan, U.S.A. 1952; Hemming, 1955).
(55)
56
J. . BURCH
•^ '':
>»v
.7*«'
X
• "^ • » ^ v tie"
10
Al
• * •
mi
FIGS. 1-5, Chromosomes Acroloxus lacustris. Figs. 1-3. Spermatogonia! metaphase chromo-
oi
somes. The chromosomes are excessively contracted. A camera lucida drawing of these
in Fig, 1
chromosomes is shown in Fig. 16. The chromosomes in Figs. 2 and 3 have divided. Camera lucida
drawings of Figs. 2 and 3 are shown in Figs. 18 and 24. Fig. 4. Late prophase I (diakinesis)
chromosomes. Fig. 5. Metaphase I chromosomes.
FIG. 6. Spermatogonial late prophase chromosomes of Rhodacmea cahawbensis.
FIG. 7. Spermatogonial metaphase chromosomes of Laevapex fuscus. A camera lucida drawing
of these chromosomes is shown in Fig, 17.
. .
CYTOTAXONOMY OF ACROLOXUS 57
the differences exhibited by Acroloxns, taken along the River Thames at Sonning
as indicated by Walker's (1923) precedent (near Reading) on April 13, 1959, and
of placing it in a subfamily by itself from 7 specimens collected near East
among the Ancylidae. Most recently Bergholt, Suffolk, on June 26, 1959. The
Hubendick (1962) has also argued for tissues were killed, fixed and preserved
familial status ioT Acroloxns on morpho- in Newcomer's (1953) fluid and stainedby
the acetic-orcein squash technique (La
logical grounds.
Chromosome studies of freshwater lim- Cour, 1941) for chromosome studies, or
pets are rather sparse and to date are stained with haematoxylin and eosin for
restricted to only six publications (Le a general histological study of gameto-
Calvez and Certain, 1950; Burch 1959a, b, genesis. Tissues for the latter study
1960a, c; Burch, Basch and Bush, 1960). were washed in absolute alcohol, cleared
in chloroform, embedded in paraffin and
The present report presents cytological
information obtained in our laboratory sectioned at 15 miera. Shells of dupli-
cate specimens (from the Thames River
on the common European limpet, Acro-
loxus lacustris, and discusses the signi- at Sonning) have been deposited in the
ficance of these findings in respect to collection of the Museum of Zoology, Uni-

commonly held concepts of systematics versity of Michigan (UMMZ cat. no.
and phylogeny in freshwater limpet-like 207600).
mollusks Observations were made with aTiyoda-
Grateful acknowledgement is made to microscope using a 90X (n.a. 1.25) oil
Dr. Dorothea Franzen, Illinois Wesleyan immersion objective and 10-30X oculars.
University, Bloomington, Illinois, U. S. A., The chromosomes in Figs. 12-26 were
and to Mr. Jack Hayworth, University of drawn with the aid of a camera lucida
Reading, Reading, England, for supplying and reproduced at a table top magnifica-
me with the cytological material of Acro- tion of 4650X. Photographs (Figs. 1-11)
loxns lacustris. I am also indebted to were taken using a 20X ocular, oil im-
Dr. Henry van der Schalle, Museum of mersion objective, a Kodak Wratten 57A
Zoology, University of Michigan for faci- (green) filter, and Kodak High Contrast
lities and many kindnesses, to Mrs. Eliz- Copy and Ektachrome Type F films.
abeth Poulson, also from our Museum,
for technical assistance during part of
OBSERVATIONS
the work, and to Mrs Anne Gismann for
.
critically reading the manuscript. 1. Cytology of Acroloxns lacustris
a. Sperm atogonial Divisions

MATERIALS AND METHODS
Thirty-six chromosomes are easily
Specimens used in this study were ob- counted during metaphase of spermato-
tained by Dr. Dorothea Franzen and Mr. gonial divisions of Acroloxns lacnstris
Jack Hayworth from two localities in (Figs. 1-3, 16, 18, 24). Primary con-
England. The material examined constrictions can easily be observed in these
sisted of ovotestes from 3 specimens metaphase chromosomes, and in addition
FIG. 8-9. Chromosomes of Ancylus ßuviatilis. Fig. 8. Late Prophase I (diakinesis) chromo-
somes. A camera lucida drawing of the chromosomes of a cell similar to this one is shown in
Fig. 23. Fig. 9. Metaphase I chromosomes. A camera lucida drawing of these chromosomes is
shown in Fig. 26.
FIG. 10 Metaphase I chromosomes of Rhodacmea cahawbensis

FIG. 11 Metaphase I chromosomes of Laevapex fuscus . A camera lucida drawing of these
chromosomes is shown in Fig. 22
Figures 1-11 X1470.

58 J. . BURCH
^ 17
7 ^
<^
16 7
4f^
;
%9 %m
t
m ^ ^ ^
C3r
19
¿iS
&Q A
^ 20 ^ 21
m 5;
Ä ^ L Ci. ¿^?<.
CXD
D -
LfvxTí
23
12 ^ 13 14 15
to their constrictions they are often also 2.8, 2.7, 2.7, 2.7, 2.5, 2.5, 2.5, 2.5, 2.3,
very noticeable because they are non- 2.2, 1.9. Similar highly contracted chro-
staining or only lightly staining (Fig. 1; mosomes have been found in other spe-
the non- staining character associatedwith cies of basommatophoran snails and have
the primary constrictions is not shown been discussed by Burch (1960a).
in Fig. 16, and only
in the shaded chro-
b. Meiotic Divisions
mosomes Fig. 12).
in
During their period of maximum con- Eighteen bivalents can be readily ob-
traction, spermatogonial metaphase chro- served during late prophase (diakinesis)
mosomes appear to be divided into 6 and metaphase of the first meiotic di-
pairs of medianly or nearly medianly visions of spermatogenesis. The pairing
constricted homologues (shaded in Figs. behavior of the bivalents appeared to be
12-14) and 12 pairs of distinctly sub- normal, and during diakinesis the paired
medianly or subterminally constricted chromosomes were held together by one
homologues (solid in Figs. 12-14). The or more chiasmata. Details of the chro-
two largest pairs and the smallest pair mosome cycle of Acroloxiis laciistris
of chromosomes are medianly constricted. during meiosis appear to be similar in
One pair of chromosomes has secondary most respects to that of other basomma-
constrictions (the 10th pair shown in Fig. tophoran snails as described by Burch
12). (1960c).
There considerable variation in the
is
degree of contraction of spermatogonial
Mature Sperm
metaphase chromosomes. The extent of The mature sperm of Acroloxiis laciis-
this variability can readily be observed tris were first described by Retzius
in Figs. 1-3, 12-14, 16, 18, 24. Mea- (1906) and the present observations do
surements miera for the homologous
in not add additional information but merely
pairs of excessively contracted chromo- confirm his report. The mature sperm
somes shown in Fig. 1 are as follows as seen in the ovotestis occur in large
(arranged in decreasing order, see also bundles of many spermatozoa each. The
Fig. 12): 4.7, 4.1, 3.4, 3.1, 3.0, 3.0, 2.9, head segment or nucleus is extremely
FIGS. 12-14. Chromosomes of three spermatogonial cells oi Acroloxus laciistris. The chromo-
somes are paired and arranged according to decreasing lengths. Medianly constricted chromo-
somes are shaded. Fig. 12. Aligned chromosomes of Figs, 1 and 16. Fig. 13. Aligned chromosomes
of Figs. 2 and 18. Fig. 14. Aligned chromosomes of Figs. 3 and 24.
FIG. 15. Aligned mitotic metaphase chromosomes of Laevapex fiiscus from Figs. 7 and 17.
FIG. 16 Camera lucida drawing of mitotic metaphase chromosomes of Acroloxus lacustris

shown in Fig. 1 These chromosomes are shown paired and arranged according to decreasing
lengths in Fig. 12.
FIG. 17. Camera lucida drawing of mitotic metaphase chromosomes of Laevapex fus cus shown
in Fig. 7. These chromosomes are shown paired and arranged according to decreasing lengths in
Fig. 15.
FIG. 18. Camera lucida drawing of mitotic metaphase chromosomes of Acroloxus lacustris
shown in Fig. 2. These chromosomes are shown paired and arranged according to decreasing
lengths in Fig. 13.
FIG. 19. Camera lucida drawing of late mitotic prophase chromosomes oi Ancylus fluviatilis.
FIGS. 20-21. Camera lucida drawing of chromosomes of meiosis I of Ferrissia parallela. Fig.
20. Metaphase I. Fig. 21. Late Prophase I (diakinesis).
FIG. 22. Camera lucida drawing of Metaphase I chromosomes of Lae^ya/êAr/MscMS shown in Fig, 11.
FIG. 23. Camera lucida drawing of Prophase I (diakinesis) chromosomes of Ancylus fluviatilis.
A photograph of the chromosomes of a cell similar to this one is shown in Fig, 8.
Figures 12-23 X1940.
.
60 J. . BURCH
long and thin (thread-like), and, in acetic- the ancyline limpets, Rhodacmea cahaiv-
orcein squash preparations, can hardly bensis (Fig. 6; late spermatogonial pro-
be distinguished from the sperm tail seg- phase) and Ancyhis fluviatilis (Fig. 19;
ment. The contrast of this sperm mor- late spermatogonial prophase), and the
phology to that of other basommatophoran laevapecine^ limpet Laevapex fusciis
snails will be discussed below. (Figs. 7, 15, 17, 25; spermatogonial meta-
phase) .
2. Comparisons with Ancylids and comparing the chromosome number

In
other Basommatophora. of Acroloxus lacustris (n=18, 2n=36) with
After looking at many basommatophoran those known from other freshwater lim-
representing many groups, I pets (Table it is readily apparent that
I),
species,
have been struck by the general uniform- this although characteristic of
number,
ity of various details of spermatogenesis. the Basommatophora in general, has not
This includes the general appearance of yet been observed in other freshwater
the cells at the various stages, the vol- limpets. The numbers n=15, 30 and 60
ume ratio of the nuclear material to the (x=15) have been reported for the Ancy-
cytoplasm, the general size of the chro- linae and Ferissiinae (Burch, Basch and
mosomes, and the appearance of the Bush, 1960) and the number n-17 for the
sperm. In Acroloxus lacustris, I was Laevapecinae^ (Burch, 1960a, c) (see
surprised to find that these various de- Table I).
tails were strikingly different. When the caryotype oi Acroloxus lacus-

tris (Figs. 12-14) is compared to that
a. Spermatogonial Chromosomes of Laevapex fuscus (Fig. 15), the most
All of the cells of spermatogenesis obvious difference between the two (other
are much larger than those observed in than sizes of the chromosomes and their
other basommatophoran snails, and the number) is that the two largest pairs of
ratio of chromatin to cytoplasm is no- chromosomes of ,
lacustris are medianly
ticeably greater (although no actual quan- constricted, while the two largest pairs
measurements were made)
titative of L. fuscus are submedianly or subterm-
The mitotic metaphase chromosomes inally constricted. It would be very in-
observed during spermatogonial divisions teresting to compare the caryotypes of

are noticeably larger than any previously other species of freshwater limpets and
observed in Euthyneuran snails. Nor- to ascertain whether or not the above
mally contracted metaphase chromosomes differences will prove to be significant.

during these divisions measure from 7.0 b. Meiotic Chromosomes
miera for the largest chromosomes to
3.5 miera for the smallest. Measure- The meiotic chromosomes of AcroZoxHS
ments for spermatogonial chromosomes lacustris are considerably larger than
in similar preparations for other fresh- those seen by me or reported by others
water basommatophoran snails have been for any other basommatophoran snail.
given as 3.7 miera for the largest and These size differences, as compared to
0.7 miera for the smallest (Burch, 1960a, various ancylid limpets, are illustrated
c; Burch and Bush, 1960). Thus, in com- in Figs. 4, 5, 8-11, 20-23, 26. Late pro-
parative terms, the chromosomes of phase I chromosomes of A.
(diakinesis)
Acroloxus lacustris are approximately lacustris are shown in Fig. 4. As can
twice the size previously found for other readily be seen, these are much larger
species of the order. This size differ- than the chromosomes of the same stage
ence can be readily observed, as regards of Ancylus fluviatilis (Figs. 8, 23) and
the Ancylidae, by comparing the chromo-
somes of A. Zací^síns (Figs. 1-3, 12-14, In this paper the Laevapecinae are considered
16, spermatogonial metaphase)
18, 24; as a subfamily of the Ancylidae (see Footnote
8 and Table I).
with the corresponding chromosomes of
TABLE I: Chromosome Numbers In Freshwater Limpets^
Species
62 J. . BURCH
"
<''^
25^ ' 26
^
FIG. 24. Camera lucida drawing of mitotic metaphase chromosomes of Acroloxus lacustris
shown in Fig. 3. These chromosomes are shown paired and arranged according to decreasing
lengths in Fig. 14.
FIG. 25. Camera lucida drawing of mitotic prometaphase chromosomes of Laevapex fuscus.
FIG. 26. Camera lucida drawing of metaphase I chromosomes of Ancylus ßuviatilis shown in
Fig. 9.
Figures 24-26 X1940.
considered modified. Pri-

this type are Since the two modified sperm types
mitive sperm, although occuring in the found in the Basommatophora appear to
Placophora (^Polyplacophora), Soleno- be found also in the opisthobranchs, it
gastres (Âplacophora), Pelecypoda, Sca- might seem that the Basommatophora are
phopoda and Diotocardia, have not been really diphyletic, and that each group was
found in Euthyneura. derived independently from different opis-
The sperm of Acroloxus lacustris are thobranch ancestors. According to Fran-
highly modified. The nucleus, instead of zen (1956): *'If within a larger group a
being short and round or conical, is very subgroup is found which has an entirely
long, forming almost 1/4 the length of the different type and genesis of the sperm
spermatozoon. It is thread-like and hard- unaccompanied by any essential differ-
The acro-
ly thicker than the tail piece. ences in the way of fertilization, and if
some can hardly be distinguished from the same type of sperm is found within
the nucleus. another group the assumption is justified
The bullet- or turnip-shaped sperm that the sperm possesses aphylogenetico-
nuclei of the other Basommatophora would taxonomical significance." But after
at first sight appear much like that de- comparing the species he studied with
scribed above as primitive. However, current concepts of Euthyneuran phylogeny
Franzen considers them to also
(1955) (e.g., Boettger, 1955) I was unable to
be highly modified without " .any trace. .
correlate the recurrence of the two types
whatsoever of primitive conditions in any of sperm with any consistency. It would
of theforms." It is interesting to note seem therefore that the modified sperm

that Franzen also reports sperm ap- of both orders arose independently; or,
proaching this second modification from that the question of phylogeny and rela-
some opisthobranchs (Actaeon, Onchi- tionships in Euthyneuran snails has not
doris, Partidida, Tritonia). been finally settled.
DISCUSSION Thiele (1931) raised it to familial rank.

Later, Boettger (1955) reduced Latiinae
1 . Systematics
again to subfamilial rank, but within the
It seems desirable to present a brief Chilinidae.
historical account of systematics at the Gwatkin (1914) pointed out that in radu-
family level for freshwater limpets in lar characters and jaw the west Ameri-
general, since some present workers on can Lanx resembled the Lymnaeidae.
these snails are either unaware of their Pilsbry (1925) followed up this observation
classical groupings and of the changes and raised Walker's (1917) subfamily
subsequently made, or tend to ignore Lancinae to family status. He noted that
them. Moreover, since Acroloxus has Lanx had "no direct or near relation to
nearly always been placed in the rather Ancylidae. It belongs to a separate fam-
inclusive family Ancylidae, systematic ily, related to the Lymnaeidae somewhat
considerations of that family also, in as the Ancylidae are to thePlanorbidae."

part, involve Acroloxus. H. B. Baker (1925) showed conclusively
In 1923 Bryant Walker, the leading that anatomically Lanx was indeed a
authority on freshwater limpets, made lymnaeid, but retained the family name
the following observation concerning their Lancidae, because of its peculiar modi-
taxonomy: "The classification of the fication of the palliai complex. Thiele
Ancylidae is in a very unsatisfactory con- (1931) again subordinated Lancinae to
dition owing to the fact that . . . practi- subfamilial rank, but within the Lym-
known of the soft anatomy.
cally nothing is naeidae.
The simple form of the shell not only Pilsbry and Bequaert (1927) placed
renders the determination of the species Protancylus (sole genus ofWalker's (1923)
exceedingly difficult, but affords very subfamily Protancylinae) with the Planor-
slight indications of generic relations, bidae (Bulininae). Hubendick (1958) in-
and none as to the evolutionary history dependently concluded that this change
and affinities of the various groups." was necessary. Zilch (1959) also con-
Our state of knowledge concerning this sidered Protancylus as a piano rbid, but
perplexing group has advanced surpris- retained its subfamilial ranking.
ingly little since that time. The ana- Bondesen (1950) proposed splitting off
tomy of most freshwater limpets is still another limpet, Acroloxus from the An-
,
largely unknown. cylidae and placing it in its own family.

It has long been known that all fresh- In doing this he raised Thiele's (1931)
water limpet-like mollusks should not be subfamily Acroloxinae (Âncylinae Wal-
placed in the single family Ancylidae ker 1923) to the family Acroloxidae. The
Rafinesque 1815 (see also Phylogeny p. 64). reason for this elevation in the syste-
Some of the genera formerly includea matic position of Acroloxus was its strik-
have already been transferred to other ingly different egg- capsule morphology.
families on well founded anatomical There exist several other reasons for
grounds. As early as 1882 Hutton desig- separating the dextral Acroloxus irom the
nated a separate family Latiidae for the sinistral Ancylidae (as understood here)
New Zealand genus Latia because of the and, more recently Zilch (1959), Burch
difference in position of its eyes, its (1961b) and Hubendick (1962) also con-
peculiar radular teeth and lack of jaws. sidered the Acroloxidae as a distinct
However, Hutton' s paper was overlooked family.
by later authors. For example, Pelseneer The remaining taxa have generally been
(1901) and Walker (1923) both indicated left with the Ancylidae or, if raised to
that Latia should be put into a separate familial rank, retained as closely associ-
family, but neither actually made this ated families. The elevation of certain
change. Hannibal (1912) proposed the groups to family status has not been uni-
subfamily Latiinae, "n. sub-fam." and versally accepted, and the grounds for
64 J. . BURCH
these changes are usually not well found- ample. Walker (1923) stated: "The re-
ed? markable differences that have recently
Hannibal (1914) raised his 1912 sub- been discovered [by Gwatkin] in the rad-
family Laevapecinae to family rank. ulae of the various groups would certainly
Wenz (1938) raised Hannibal's (1912) sub- tend to strengthen the suggestions that
family Neoplanorbinae and Walker's (1917) have been made that the family [Ancyli-
subfamily Ferrissiinae to family status, dae] as now recognized is not of homo-
and also presumably Walker's (1923) An- geneous origin, but of diverse ancestry,
cylastruminae to Ancylastridae. Boettger and that the similarity in conchological
(1955) and Meyer (1955) also treated the characters should be considered rather
Ferrissiinae as a separate family. Zilch as an example of parallel development
(1959) recently raised Walker's (1917) than as indicating a common line of de-
subfamily Rhodacmeinae to familial rank. scent." Although Walker was perhaps
The descriptive cytological details and the first to state clearly such views on
comparisons shown in this paper strength- the polyphyletic origin of freshwater lim-
en Bondesen's contention that Acroloxiis pets, Hannibal (1914) was apparently ear-
should be separated at the family level lier thinking along similar lines (although
from other freshwater limpets. This his groupings are largely unnatural ones).
has already been pointed out in a brief Phylogenetic considerations of fresh-
abstract by Burch (1961b). Their higher water limpets go back many years, to
chromosome number (n=18) which differs Plate (1894), who considered Ancylus to
from that of theAncylinae andFerissiinae be a tectibranch. Pelseneer (1901) took
(x=15), and Laevapecinae (n=17) is also exception to this and placed Ancylus (A.
considered as an additional cytological fluviatilis + Acroloxiis lacustris) and
character speaking for separation. The Gundlachia near Planorbis with the Ba-
divergent chromosome number of the sommatophora. His reasons for con-
Laevapecinae would suggest possible ba- sidering Ancylus to be a basommatophor-
sic differences from the Ancylinae also, an were: 1) the shape and position of
which, if borne out by anatomical detail, its osphradium, 2) its palliai "gill", 3)
might lead to their removal from the its stomach with a pyloric caecum, 4) the
Ancylidae. As regards Acroloxiis, such position of the ventricle in relation to

a systematic separation as supported here the auricle in its heart, 5) the separation
seems justified also on more gross mor- of its male and female genital openings,
phological grounds, since considerable and short second pedal commissure.
6) its
differences in shell, radula and soft ana- He showed that Gundlachia was similar
tomy have been demonstrated between to Ancylus in its basic organization.
Acroloxiis and other freshwater limpets In relation to other Basommatophora,
(e.g., see Walker, 1923; Hubendick, 1960, Pelseneer considered Ancylus to be very
1962). specialized because of 1) its very concen-
trated nervous system, 2) its acquisition
2. Phylogeny
of a secondary gill, 3) its loss of the
For many years it has been known, or lung, 4) the displacement of its heart,
at least suspected, that the freshwater and 5) the early division of its herma-
limpets are not monophyletic. For ex- phroditic duct and the consequent great
length of oviduct. The position of the
its
"The Ancylidae are here considered to contain Ancylidae as now understood (and exclud-
the following genera: Ancylastrum (Ancy- ing /av/is), both in their relation to
lastruminae); Ancylus, ronde lia and Rhodac- the Planorbidae and in regard to the
mea (Ancylinae); Ferrissia, Gundlachia and other Basommatophora has not been chal-
Hebetancylus (Ferrissiinae); Anisancyliis Bur-
,
lenged by subsequent authors (e.g., see

nupia, Laevapex and Uncancylus (Laevapecinae).
This is a modification of Walker's (1923) clas- Hubendick, 1947; Meyer, 1955; Boettger,
sification. 1955).
Hubendick (1945) and others before him neuran snails needs to be mentioned since
consider Chilinidae and Amphibolidae to they have phylogenetic significance. If
be primitive Basommatophora. Since the one disregards erroneous and unreliable
more specialized Lymnaeidae appear to cytological reports (for discussion see
him (Hubendick, 1947) to be similar to Burch (1960c)) and considers chromosome
these two families in type of tentacles numbers in the various Euthyneuran
and in having a muscle on the ventral groups for which reliable information
side of the kidney, he considers the lym- regard to their presumed
is available, in
naeids to be the most primitive of the morphological advancement, one immedi-
" higher limnic Basommatophora" (-Bran- ately sees that there is a gradual in-
chiopulmonata) In addition, he considers
.
crease in chromosome numbers as one
some lymnaeids^ and Chilùm to have simi- goes up the evolutionary scale. Haploid
lar copulatory organs. numbers of the marine "opisthobranchi-
Since among the '' Branchiopulmonata" ate" snails are all less than 18 (Fig.
the Ancylidae and the Planorbidae are 27). The basic haploid number (x) for
most dissimilar to the Lymnaeidae, they the freshwater Basommatophora is 18
appear to Hubendick to be the most spe- (Burch 1960c). The haploid numbers of
cialized.
A))iphibola and ,
However, he points to certain
similarities that the Ancylidae have with
i.e., the presence
of an anal lobe, of lobate salivary glands,
and of a flagellum (he considers the fla-
nearly all members of the land-dwelling
Stylommatophora are greater than 18.
The increase in number is a gradual one
as one goes from one closely related
larger taxon to the next, indicating that
gellum of Ancyhis to be histologically change in chromosome number has re-
different from that of Amphibola: Chilina sulted by aneuploidy rather than poly-
does not have a flagellum). The simi- ploidy. This was first pointed out for
larities of the salivary glands he con- pulmonate snails by Husted and P. R.
siders unimportant. But the anal lobe Burch (1946). In those families where
he does not so easily dismiss. He (loc. polyploidy has been reported (Burch,
cit.) says: "The homologization of An- 1960b, d; Burch, Basch and Bush, I960)
cylus' lobe with one of those in Planor- the diploid number basic to the group is
bidae offers certain difficulties", and readily discerned. The great constancy
" as the shape of the lobe and the course of chromosome numbers within large tax-
of the rectum in the lobe agree well in onomic groups, especially in the lower
Ancylidae with those in Thalassophila Euthyneura, suggests that aneuploidy has
and Chilinidae, it might be possible for been a rather rare occurrence, and that,
the lobe to be homologous with the anal when it occurs, it usually involves a
lobe. .
." thereby indicating greater pri- whole major group, e.g., an order or
mitiveness. But Hubendick continues to family (sometimes only a genus in the
say that '* .if Ancylidae ought to be
. .
Stylommatophora). Polyploidy is rare,
derived from any recent types of higher and when it occurs it usually involves
limnic Basommatophora" then " the justi- species (perhaps genera in the Ancylidae),
fication for this last argument disap- and not larger groups. Also, chromo-
pears ."I
. .
some change by aneuploidy, when relat-
At this point, the nature of the vari- ing to higher taxonomic categories seems
ation of chromosome numbers in Euthy- to be by addition, rather than subtraction,
of chromosomes. (For a more detailed
9 He says that Lanx differs from other lymn- discussion of the above concepts see
aeids in that it lacks a praeputium, but H. B. Burch, 1961a; Burch and Heard, 1962).
Baker(1925)in his excellent detailed treatment With the above information at hand it
oi Lawc quite clearly describes the praeputium.
is then difficult to consider that the An-
I have also determined that Lanx has a well
developed praeputium which seems to differ cylidae as here understood (x=15; n=17)
but little from that of other Lymnaeidae. are highly specialized derivatives of the
66 J. . BURCH
60
36
34
32
30
^ 28
e
z
26
" ^^
24
e
¿ 22
20
X 1
16
14
12
10
pulmonata"
STYLOMMATOPHORA BASO MMATOP HORA

(n>18)
Branchiopulmonata
Phys idae
x=ir
F lanorbidae
Lymnae idae x=18
x=l
Aero loxidae Ancy lidae

n=l x=15,17
Archaeopu Imonat a
EL Lobiidae 'Ur-Basommatophora'
x=18 x=18 x=15,17
.
Amphibolidae •
Siphonariidae
n=18 n=16
"Opisthobranchiate"
anees tors
x=12,13,17
FIG. 28. Possible relationships of various taxa within the subclass Euthyneura based (in part) on
haploid chromosome numbers. For phy logenies based on various morphological considerations see
Hubendick (1947) Boettger (1955), Meyer (1955) and Morton (1955),
,
Planorbidae (x=18) as contended by Pel- the must be

origin of the Acroloxidae '*
seneer (1901), but it would seem that . . sought nearer to more primitive
.
they are perhaps much more primitive forms within the Basommatophora." Hu-
in nature. Brief attention has already bendick (1962) most recently has advo-
been called to this possibility by the au- cated that Acroloxus had an origin in
thor (Burch, Basch and Bush, 1960). common with the primitive and aberrant
Even tor Acroloxus despite agreement in
,
Latia of New Zealand. He bases his con-
the chromosome numbers (n=18), there clusions on what he considers similari-
seems little justification in leaving its ties in the radulae and reproductive tracts
connections with the Planorbidae, when of the two genera. However, Pelseneer
one considers the gross differences in (1901) who alone has studied the soft ana-
other details of cytology, discussed previ- tomy of Latia, pointed out certain im-
ously in this paper. portant dissimilarities between Latia and
Bondesen (1950), in considering the re- the ancylids (including Acroloxus lacus-
lationship of Acroloxus to other fresh- tris, with which he was familiar) that
water pulmonates as shown by compara- cannot be overlooked. These differences
tive egg capsule morphology, states that include the morphology of the nervous
. ,
68 J. ß. BURCH
system, the kidney, the lack of flagellum morphological data, which at present is
in Latia's male reproductive system (per- unavailable for many Euthyneuran groups.
haps a superficial character), and the Relationships of freshwater limpets to
presence of a well-developed pulmonary other Euthyneura as suggested by their
cavity in Latia's Concerning the radula chromosome numbers might be illustrated
of Acroloxus Hubendick (loc. cit.) states as in Fig. 28.
that it " . has certain important simi-
. .
larities with that of Latia ." But, after LITERATURE CITED

carefully examining the radula of Latia,
BAKER, H. ., 1925, Anatomy of Lanx, a limpet-
I find it very different irom Acroloxus
like lymnaeid mollusk. Proc. Calif. Acad,
and cannot understand Hubendick' s state- of Sei. (Fourth Series), 14(8): 143-169, pis.
ment, unless he considers the arched 11-14.
rows of radular teeth of Latia and the BOETTGER, C. R., 1944, Basommatophora, In:
semi-arched nature of those oi Acroloxus
to be important. The only other author
Grimpe und Wagler:
und Ostsee. Liet 35, Teil 242-478.
Die Systematik der euthy-
1955,
'
Tierwelt der Nord-
:
that I know of that has studied the radula neuren Schnecken. Verhandl. Deutsch. Zool.
of Latia is Hutton (1882), and his draw- Ges. Tubingen, 1954: 253-280.
ings also do not show the radular teeth BONDESEN, P., 1950, A comparative morpho-
logical-biological analysis of the egg cap-
of Latia to be similar to those reported
sules of freshwater pulmonate Gastropods.
by other authors for Acroloxus (e.g., Natura Jutlandica, 3: 1-208, pis. 1-9.
see Walker, 1925). BURCH, J. ., 1959a, Chromosomes of aquatic
The aberrant spermatozoon of Acro- pulmonate snails (Basommatophora). Amer,
loxus lacustris has been considered an Malacol. Union Ann. Reps. 1958, 25: 9-10.
extreme specialization when compared ,
1959b, Chromosomes of aquatic
pulmonate snails (Basommatophora), Dis-
with Lynnmea, Physa and Planorbarius
sert. Abstr., 20(4):1487-1488,
(see Retzius, 1904; also Boettger, 1944). 1960a, Chromosome morphology of
,
But if Bondesen's and Hubendick' s sug- aquatic pulmonate snails (Mollusca: Gastro-
gestions are correct in that Acroloxus poda). Trans. Amer. Microsc, Soc, ,79(4):
had its origin near to primitive forms 451-461,
1960b, Chromosomes of Gyraulus
within the Basommatophora, and if it ,
circumstriatus a freshwater snail. Nature,

,
might be assumed that such a starting 186(4723): 497-498.

point could be from a group more pri- 1960c, Chromosome studies of
,
mitive than those so far studied in re- aquatic pulmonate snails. Nucleus, 3(2):
177-208.
spect to spermiogenesis, then perhaps
1960d, Chromosome numbers of
the anomalous sperm morphology might
,
schistosome vector snails. Z. Tropenmed.

not by in the direction of extreme spe- Parasit., 11(4): 449-452.
cialization, but more closely like that of 1961a, The chromosomes of Plan-
,
less specialized living groups not yet orbarius corneus (Linnaeus), with a discus-
thoroughly investigated. Or perhaps the sion on the value of chromosome numbers in
snail systematics. Basteria, 25(4/5): 45-52.
aberrant sperm are related to more spe- 1961b, Some cytological aspects of
,
cialized species of a group less special- Acroloxus lacustris (Linnaeus), with a dis-
ized or more primitive than the Basom- cussion of systematics in fresh-water lim-
matophora (i.e., an " opisthobranch" pets. Amer. Malacol. Union Ann. Reps.
group) 1961, 28: 15-16.
BURCH, J. B.,BASCH, P. F. and BUSH, L. L.,
That a critical réévaluation of phylo- 1960, Chromosome numbers in ancylid
geny and phylogenetically significant char- snails. Rev, Portuguesa Zool. Biol. Ger.
acters in the so-called "higher limnic 2(3/4): 199-204.
Basommatophora" or " Branchiopulmon- BURCH, J. B. and BUSH, L, L, 1960, Chromo- ,
ata", indeed, in the entire Euthyneura, somes oi Physa gyrina ?>a.y (Mollusca: Pul-
monata). J. Conchy 1., 100: 49-54.
is badly needed is readily apparent. But BURCH, J, B. and HEARD, W. H. 1962, Chromo-
such a réévaluation of relationships must some numbers of two species of Vallonia.
await the proper accumulation of detailed (Mollusca: Stylommatophora: Orthurethra),
Acta Biol, Acad, Sei. Hung., 12(1): 305-312.
,
FRANZEN, A., 1955, Comparative morphologi- caryologiques sur quelques Pulmones ba-
cal investigations into spermiogenesis among sommatophores. . R
Acad. Sei., Paris,
Mollusca. Zool. Bidrag Uppsala, 30: 399- 231(16): 794-795.
456, pis. 1,2. MEYER, .
., 1955, Naturgeschichte der
, 1956,
On spermiogenesis, morphol- Strandschnecke Ovatella myosotis (Drapar-
ogy of the spermatozoon, and biology of naud).
fertilization among invertebrates. Ibid., 31: MORTON, J. E., 1955, The evolution of the El-
355-482, pis. 1-6. lobiidae with a discussion on the origin of
GWATKIN, H. M., 1914, Some moUuscan radu- the Pulmonata. Proc. Zool. Soc. Lond., 125
lae. Conchol., 14(5): 139-148.
J. (1): 127-168.
HANNIBAL, H., 1912, A synopsis of the recent NEWCOMER, E. H., 1953, A new cytological and
and tertiary freshwater Mollusca of the histological fixing fluid. Science, 118(3058):
Californian Province, based upon an onto- 161.
genetic classification. Proc. Malacol. Soc. PELSENEER, P., 1901, Études sur des Gastro-
Lond., 10: 112-211. podes pulmones. Mem. Acad. Roy. Sei.,
1914, Note on the classification of Lettr. Beaux- arts Belg., 54: 1-76, pis. 1-14.
,
the Ancylidae. Nautilus, 28(2): 23-24. PILSBRY, H. A., 1925, The family Lancidae
HEMMING, F. (Ed.), 1955, Opinion 363. Desig- distinguished from the Ancylidae. Nautilus,
nation, under the plenary powers, of a type 38(3): 73-75.
species in harmony with accustomed usage PILSBRY, H. A. and BEQUAERT, J., 1927, The
for the Müller
nominal genus "Ancylus" aquatic mollusks of the Belgian Congo, with
(. 1774 (Class Gastropoda). Opinions
F.), a geographical and ecological account of
and Declarations Rendered by the Interna- Congo malacology. Bull. Amer. Mus. Nat.
tional Commission on Zoological Nomencla- Hist., 53(2): 69-602.
ture, 11(12): 185-202. PLATE, L., 1894, Studien über opistopneumone
HUBENDICK, ., 1945, Phylogenie und Tier- Lungenschnecken. II. Die Oncidiiden. Ein
geographie der Siphonariidae. Zur Kenntnis Beitrag sur Stammesgeschichte der Pul-
der Phylogenie in der Ordnung Basomma- monaten. Zool. Jahrb., Anat, 7: 93-234.
,
tophora und des Ursprungs der Pulmona- RAFINESQUE, C. S., 1815, Analyse de la Na-
tengruppe. Zool. Bidrag Uppsala, 24: 1-216. ture, ou Tableau de l'Univers et des Corps
1947, Phylogenetic relations be-
,
Organisés. Palerme. p. 136-149.
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Ibid., 25: 141-164. der Evertebraten. Biol. Untersuch., N. F.,
1952, Proposed use of the plenary
, 11(1): 1-32, pis. 1-13.
powers to designate a type species for the 1906, Die Spermien der Gastro-
,
genus "Ancylus" Müller, 1774 (Class Gas- poden. Ibid., 13(1): 1-36, pis. 1-12.
tropoda) in harmony with established nomen- THIELE, J. 1931, Handbuch der systematischen
,
clatorial practice. Bull. Zool. Nomencl. Weichtierkunde. Jena 1929-35. 2: 377-778.

6(8): 227-230. TUZET, O. 1950, Le spermatozoïde dans la
,
(1958, A note on Protancylus P. and

,
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1960, The Ancylidae of Lake Ochrid
,
WALKER, ., 1917, Revision of the classifica-
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1962, Studies on Acroloxus (Moll.
, Nautilus, 31(1): 1-10, pis. 1-3.
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,
och Vitterhets-Samhälles Handlingar. Sjätte London: Printed for the Author. P. 82.
Följden. Ser. . 9(2): 1-68. 1925, New species of North Ameri-
,
HUSTED, L. and BURCH, P. R., 1946, The can Ancylidae and Lancidae. Univ. Mich.,
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14: 150-158, pis. 3 and 4. 960. Borntraeger, Berlin.
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LE CALVEZ, J. and CERTAIN, P. 1950, Données ,
70 J. . BURCH
ZUSAMMENFASSUNG
ZYTOTAXONOMISCHE STUDIEN ÜBER DIE NAPFSCHNECKEN
DES SÜSS WASSERS (GASTROPODA: BASOMMATOPHORA). I. DIE
EUROPÄISCHE BINNENSEENAPFSCHNECKE ACROLOXUS LACUSTRIS.
Acroloxus lacustris (Linnaeus) ist eine in Europa, Nordasien und Kaukasien

verbreitete Napfaschnecke die gewöhnlich der Basommatophorenfamilie Ancylidae
zugerechnet wird und daher auch zu den spezialisiertesten und phylogenetisch am
weitesten fortgeschrittenen Basommatophoren.
Es wird hier gezeigt, dass in Anbetracht gewisser Einzelheiten des Zellbaues,
A. lacustris nicht als eine den anderen Ancyliden verwandte Schnecke angesehen
werden sollte, sondern eher in eine Familie für sich (Acroloxidae)gestellt werden
sollte, wie es ja auch bereits von anderen Autoren aus morphologischen Gründen
befürwortet wurde. Ja darüber hinaus scheinen die beobachteten zytologischen
Unterschiede sogar für einen weiteren Abstand von den übrigen Basommatophoren
zu sprechen. Diese Unterschiede bestehen aus der beträchtlicheren Grösse der
verschiedenen spermatogenetischen Zellen aus deren im Verhältnis zum Zyto-
,
plasma grösserer Chromatinmasse, aus den verhältnismässig grossen Ausmassen

der Chromosome und aus der Gestalt der Spermatozoen, deren Köpfe lang und
fadenförmig und nicht geschoss- oder rübenförmig sind wie die der anderen Basom-
matophoren. Ausserdem ist die Chromosomenzahl (n=18) des Acroloxus obwohl ,
für die Basommatophoren kennzeichnend, von der der anderen Napfschnecken des
Süsswassers verschieden; die haploide Grundzahl ix) beträgt in den Ancylinae
und den Ferrissiinae 15, während die in den Laevapecinae n=17 beträgt.
Wie für sämtliche Basommatophoren charakteristisch, sind die mitotischen
Chromosome des Acroloxus lacustris metazentrisch. Sechs Paare, darunter die 2
grössten und das kleinste, sind median, die übrigen 12 submedian oder subzentral
abgeschnürt. Es wurde hier zum ersten Male die Kariotype einer euthyneuren
Schnecke genau festgestellt und abgebildet.
Es könnte wohl sein dass die Acroloxidae, wie es schon Bondeson und Huben-
dick vertraten, phylogenetisch in die Nähe der Wurzel des Basommatophoren-
stammes zu stellen wären, doch kann die Zytologie, zumindestens beim heutigen
Stand unseres Wissens, noch nicht das Beweismaterial für einen derartigen
Schluss liefern.
Auf Grund ihrer Chromosomenzahl wären auch die Ancylidae, im Gegensatz
zur üblichen, auf rein anatomischer Grundlage geschlossenen Folgerung, ebenfalls
in die Nähe der Wurzel der Basommatophoren zu versetzen, jedoch in Hinblick auf
die anderen zytologischen Unterschiede, nicht in die unmittelbare Nähe der
Acroloxidae.
RESUME
ETUDES CYTOTAXONOMIQUES SUR LES PATELUENS D'EAU DOUCE
(GASTROPODA: BASOMMATOPHORA) I. LA PATELLE
LACUSTRE EUROPÉENNE ACROLOXUS LACUSTRIS.
L'Acroloxus lacustris (Linné) est une patelle lacustre répandue en Europe, en
Asie septentrionale et en Caucasie, qui a généralement été rangée dans la famille
basommatophore des ancylides et, par là, parmi les basommatophores les plus
spécialisés et les plus avancés phylogénétlquement.
Nous montrerons dans cet exposé que, en vue de certains détails cytologiques,
A. lacustris ne saurait être placé dans le voisinage immédiat des ancylides, mais
plutôt dans une famille séparée, les Acroloxidae, conclusion qui d'ailleurs se trouve
aussi corroborée par d'autres auteurs pour raisons morphologiques. Plus que cela,
les différences cytologiques observées semblent même indiquer un certain écart
entre Acroloxus et les autres basommatophores. Ces différences se trouvent dans
la grandeur des cellules de la Spermatogenese, dans le volume supérieur de leur
chromatine par rapport au cytoplasme, dans les dimensions relativement grandes
des chromosomes et dans la morphologie des spermatozoïdes mûrs, dont les têtes
sont longues et ñliformes, ne montrant point la forme de balle ou de navet ren-

contrée chez les autres basommatophores. En outre, son nombre de chromosomes
(n=18), quoique caractéristique pour les basommatophores en général, diffère de
celui des autres patelles d'eau douce, dont le nombre haploïde fondamental de
chromosomes (x) est ISpour les Ancylinaeet Ferrissiinae, pendant que n, le nombre
haploïde, est 17 pour les Laevapecinae.
Les chromosomes mitotiques de A. lacustris sont métacentrlques de façon
caractéristique pour tous les basommatophores. Les constrictions sont médianes
dans 6 paires, comprenant les 2 plus grandes ainsi que laplus petite, et submédianes
ou subterminales dans les autres 12. C'est d'ailleurs la première fois que le cary-
otype d'un mollusque euthyneure ait été minutieusement observé et figuré.
Il se pourrait que la position phylogénétique des Acroloxidae serait à chercher
plutôt près de la base du tronc basommatophore comme l'ont suggéré Bondesen et
Hubendick, mais, à l'état actuel de nos connaissances, la cytologie ne peut pas
encore fournir les preuves d'une pareille conclusion.
Quant à la position des Ancylidae, elle se trouverait, contrairement aux con-
clusions antérieures basées sur une évaluation purement anatomique, également
proche de la base de l'arbre phylogénétique basommatophore relativement à leur
nombre de chromosomes, mais pas en proximité immédiate des Acroloxidae à
cause des autres différences cytologiques.
RESEÑA
ESTUDIOS CITOTAXONOMICOS SOBRE LAS LAPAS

DE AGUA DULCE (GASTROPODA: BASOMMATOPHORA) I. LA LAPA
DE LAGO EUROPEO, ACROLOXUS LACUSTRIS.
El Acroloxus lacustris (Linnaeus) es una lapita de agua dulce común en Europa,

norte de Asia y Caucasia. Casi siempre ha sido asignada a la familia basommato-
phora Ancylidae, considerada como una de las más especiales y filogenéticamente
avanzadas.
Se demuestra en este estudio que en base a ciertos detalles de citología A.
lacustris no debería considerarse muy relacionada a otros Ancylidae sino más
bien como constituyendo una familia por sí sola, Acroloxidae conclusión que ha
,
sido corroborada por otros autores, en base de conocimientos morfológicos.

Ciertamente las varias diferencias citológicas tienden a separar más los Acroloxus
de los otros Basommatophora. Las diferencias observadas las constituyen el
tamaño y mayor volumen de las varias células espermatogénlcas, el mayor
volumen de cromatina en relación al citoplasma, la relativa mayor medida de los
cromosomas y la morfología de la esperma madura cuyas cabezas son largas y en
forma de bala o de nabo como se encuentran en los otros caracoles basommato-
foros. En suma, el número de cromosomas (n=18), aunque característico de los
Basommatophora en general, es diferente del que se encuentra en otras lapas de
agua dulce (x o numero básico=J.5 en Ancylinae-Ferrissiinae;n=n en Laevapecinae).
Los cromosomas mitóticos del A. lacustris son metacéntricos como es
característico de todos los Basommatophora; 6 pares (incluyendo las 2 más
grandes y las más pequeñas) que están medianamente ligados. Los otros 12 pares
son submedianamente o subterminalmente ligados. Esta es la primera vez que el
La posición filogenetica de los Acroloxidae puede que este muy cerca de los
Basommatophora como sugerid Bondesen y Hubendick- Desafortunadamente al
presente estado de conocimiento la evidencia que ayudaría a sostener tal conclu-
sión no puede encontrarse en detalles de citología. Pero al contrario de opiniones
antiguas alcanzadas en base a puros conocimientos anatómicos, la posición de los
Ancylidae determinada por el número de cromosomas debería también estar muy
cerca de los Basommatophora, aunque no cerca a los Acroloxidae debido a las
otras diferencias citológicas.
72
.
. --
,
.
.
Acroloxus lascHstris
lacustris
, -
(L.)
, . .
, , ,,.,
-
Ancylidae,
- Acrol oxidas ;
-
-
, ,
;
). ; ,(
= 15 Ancyllnae -
(n=l8),
(
Ferrissiinae
.
;
lacustris
-
6
17 g
-
Laevapecinae)
2
.
,
12
,
, , .
,
,
, -
.
Euthyneure
Acroloxidae
,
-
•
,
. - Acroloxidae
Ancylidae ,
MALACOLOGIA, 1962, 1(1): 73-114
CONTRIBUTIONS TO THE MORPHOLOGY OF BULINUS TROPICUS

(GASTROPODA: BASOMMATOPHORA: PLANORBIDAE)
By
Ina Stiglingh, J. A. van Eeden and P. A. J. Ryke
Research,
Institute for Zoological
Potchefstroom University for Christian Higher Education,
Potchefstroom, Republic of South Africa.
ABSTRACT
A study was made on Bulmus (Bulinus) tropicus
(Krauss), an African fresh
water snail that intermediate hosts of the schistosomes of
is closely related to the
the hae»iatobiia)¡ group and which is also of veterinary importance as a carrier of
the trematodes Paramphistomiim and Cotylophoron. Special attention was paid to
three important taxonomic characters, the shell, radula and penial complex but the
other systems were also examined both morphologically and histologically.
The snail samples were selected both on the basis of their resemblance to
typical specimens of B. tropicus and their geographical distribution in the region of
the type locality. In spite of these precautions in the selection, the present study
of the shell once more emphasizes the notorious polymorphism of the species. The
important role of ecological factors in producing widely divergent shell types,
differing in both shape and texture, is discussed. Although two main adult types oc-
curred in our material, an elongated, narrowly umbilicate form associated with run-
ning water, and a squat, widely umbilicate form associated with pools (ñg, 3), there
was only one main type of juvenile, suggesting, as was observed by Schutte and van
Eeden (1959a) for BiompMlaria /)/(/, that the form of the juvenile is less affected
by the environment than is the case in the adults. This seems logical as the latter
have been exposed to the modifying influences for a longer period than the younger
snails. A conchometrical analysis was undertaken and the length, breath, aperture
height, aperture breadth and spire lengthof 171 snails from 9 localities were meas-
ured and various ratios calculated in order to discover the nature of the relation
between any two of these features. The angles of the spire and of the labrum de-
pression were also measured. Only comparison with corresponding values ob-
tained from other species will indicate which of these ratios differ sufficiently to be
of use taxonomically. However, though the ranges of these values are in some cases
ereat, the means for the 9 samples are usually fairly constant. An exception is the
ratio shell length to spire length in which the mean values vary between 6.7 and
15.2, The spire was, nevertheless, found to be relatively shorter in the smaller
°-
forms than in the large specimens. The apical spire angle is very variable: 58
134° .The mean value of the angle of labrum depression is 57° or 58° for all the
localities except one for which the value is 61° In respect of the shape of the aper-
.
ture the adult shells usually fell in two main groups. This feature seems to change
not only with the age of the snail, but probably also with varying ecological con-
ditions. Although the columella region is fairly uniform in each sample it varies
from one sample to the other.
The osphradium was found to be present outside the mantle cavity on the collar.
The gill (pseudobranch) is normal. The anal region differs slightly from previous
descriptions in that the anal lobe seems narrower and the rectal ridge more dis-
tinct. The length ratio Iddney/ureter was found to vary between mean values of 2.0
and 4.6 in the various samples.
Only two of the taxonomically important ridges are present on the roof of the
mantle cavity, the renal ridge and the median rectal ridge being absent. The inter-
mediate mantle ridge (between kidney and rectum) differs from the ñgure given by
Mandahl- Barth (1956) in being longer and in always extending posteriorly to meet
the lateral rectal ridge at the edge of the mantle cavity.
An account is given of the blood vessels draining the gill (pseudobranch). The
spherical bodies mentioned by Schutte and van Eeden (1959b) as occurring in the
pericardial cavity oi Biomphalaria pfeifferi were found to be present in a large num-
ber of specimens. They are thought to be the eggs of an unidentifled organism.
(73)
74 STIGLINGH, VAN EEDEN AND RYKE
In the digestive system the aw was found to
j differ from the descriptions offered
in the literature and to conform to the general pattern for Bulimis. A study of the
general proportions of the radula revealed that the mean ratio length/breath is fairly
constant (2.4 - 2.6) and this ratio may therefore prove to be useful in comparison
with related species. The sides of the teeth were found to be fluted and small den-
ticles were detected between the cusps of the centrals and at the base of the meso-
cone of the laterals. The mesocone of the laterals is almost never simply triangular
as is stated by Mandahl- Barth (1956) to be the condition in the'ß. tropicus group.
In certain instances it even approaches the arrowhead shape characteristic of the
B. truncatus group. In spite of the known limitations of a radula formula, a gen-
eralized formula for B. tropicus is offered: 1: 6+2 24 x 123.
:
The names applied to the various ganglia of the circumesophageal ring are
discussed and the nerves originating from them described. The salivary glands
pass through the nerve ring.
In the hermaphrodite gland there Is no division into male and female zones and
the oocytes were found to have double nucleoli as is the case in certain other
pulmonates. The carrefour region differs from that reported for the north American
planorbid Helisoma trivolvis and agrees with the condition found in the African
planorbid Biomphalaria pfeifferi. Attention is drawn to an irregular sac extending
from the uterus and partly surrounding the base of the oviduct. In the prostate gland
two main regions are distinguished, a peripheral opaque, yellowish region and a
more translucent whitish central region. The occurrence of a short internal ridge
in the proximal region of the penis is noted. The pilasters in the preputium extend
as far as the junction of the latter with the penis sheath. The penis sheath/preputium
ratio was found to vary considerably, but the means for the samples separately vary
only beWeen 1.2 and 1.6.
INTRODUCTION may resemble one species of Bulimis in

The interest African fresh-water
in the some respects and another species in
snail Bidiniis (B.) tropicus (Krauss) has, others, so that it is almost impossible
up till now, been centered mainly around to ascertain to which species a particu-
the veterinary and suspected medical im- lar specimen in reality belongs. Further
postance of this species in so far as it confusion arose from the fact that the
is known to act as the intermediate host criteria used in species discrimination
for the trematodes Paramphistomum and were not always well chosen. When the
Cotylophoron and is closely allied to shell alone was used to identify a spe-
species which carry schistosomes of the cies it was usually viewed as a dead,
haematobium group in Africa. There isolated object and neither the anatomy
have been few serious attempts at a of the snail producing the shell, nor the
thorough investigation of the morphology ecological factors influencing it, were
of African planorbids. Much has been taken into account Finally there was the
written on the problem of their classi- practice of sending snails for identifica-
fication and although the crux of the mat- tion to overseas authorities who often
ter is the intrinsic variability of the applied different names to the various
species studied, numerous other factors samples of the same species. In spite
have played a part in bringing about the of Cawston's warning that only confusion
general confusion on this issue which would arise as a result of giving too
now exists. Probably the most important many names, new names were freely
of these factors is that it was not ini- applied, as is testified by the long, but
tially realised by the taxonomists that, by no means exhaustive, list of synonyms
in the case of B. tropicus, they were given in Connolly (1939). Schweiz (1952a)
dealing with an extremely variable spe- suggests certain remedies for this un-
cies. Even when the variability was re- healthy state of affairs and the main
cognized, the nature and causes thereof ideas are that there should be a sound
were not, indeed are not, properly under- biological approach to these problems
stood. In certain instances a specimen and that ultimate proof of relationships
MORPHOLOGY OF BULINUS TROPICUS 75
should be obtained by carefully planned

breeding experiments which, unfortunate-
ly might also be subject to limitations.
Besides the taxonomic tangle of con-
fusion on the specific level, brief men-
tion should also be made of some obscure
relations on the subgeneric level. Wright
(1957) considers B. hemprichii depressus
Haas as a synonym of B. (Physopsis)
globosus (Morelet) while Mandahl- Earth
(1956) maintains that the former species
is a synonym of B. (Bulinus) tropicus,
whereas we are of the opinion that both
these authors might be at fault. In view
of these and other discrepancies much
careful morphological study will yet be
necessary in this group before the true
specific or strain relations can be ac-
curately assessed.
Since bulinids represent the most com-
monly occurring freshwater snails in
South Africa, a closer investigation of
the intraspecific variation of the shell of
B. tropicus and certain aspects of its
anatomy seemed to be indicated. The
results of investigation
this are dealt
with in the present paper.
MATERIALS AND TECHNIQUES

Most of the material used in this study
was sent, for identification, to this In-
stituteby the staff of the Health Depart-
ment Republic as part of the anti-
of the
bilharziasis programme in the Transvaal.
Snails were collected from the following
localities which will, be re-
in the text,
ferred to by the symbols indicated below.
Locality
lines were then measured using an opiso- SHELL
meter (rotameter).
In spite of the fact that the shells of
Radula preparations were mounted in
Bulinus species are " notoriously poly-
CMC 10 (a Turtox product), and a num-
morphic" (Cawston, 1938) yet, even in
ber of these were drawn with the aid of
this genus, the shell exhibits certain
a camera lucida. Whole mounts of 32
characters by which the various groups
penes were made using the technique of
and species are distinguished and which
van Eeden (1958). The remaining penes
necessitate some reference to it in any
were measured, using a micrometer eye-
morphological account of B. tropicus.
piece, in the following manner. The co-
The most complete descriptions of the
pulatory organ was removed together
shell of B. tropicus are those given by
with a small portion of the vas deferens.
Krauss (1848), Germain and Neveu-Lemai-
A pin was inserted through the distal end
re (1926), Connolly (1931, 1939), Cawston
of the preputium and the organ straight-
(1938), all for South Africa, Mozley (1939)
ened, taking care that the preputium was
for Tanganyika, Mandahl-Barth (1954,
not partially invaginated. The penis
1956) for Uganda and Southern Rhodesia,
sheath was straightened by placing pins
and de Azevedo et al. (1957) for Portu-
through the distal parts of the organ and
guese East Africa. These all lead to the
through the vas deferens close to the
conclusion that there is no single descrip-
point where it enters the penis sheath.
tion of B. tropicus which will embrace
For measurement of the penis and epi-
all the diverse forms of this species and
phallus the proximal half of the penis
one has only to be confronted with a
sheath was removed and the penis and
photographic series of shells like those
epiphallus exposed. (It is not necessary
published by Mandahl-Barth (1956), or
to open the penis sheath distally as the
indeed our own series (Figs. 1 and 2)
length of penis and penis sheath are the
most fully to realise this.
same in this region). The penis was
The indications of size given in the
then straightened by pinning down its
literature are generally not very useful.
proximal region, and then the epiphallus
Boettger's (1910) data are based on a
was unravelled, straightened and pinned
few specimens or on one only. Krauss
down by a small piece of the penis sheath
(1848) seems to have examined a number
left at the junction of vas deferens and
of shells while de Azevedo' s mean, based
epiphallus.
on a too small sample of 5 shells, is also
The radula, penis and the rest
shell,
unreliable. In his 1956 publication Man-
of the soft parts of each snail were given
dahl-Barth does, however, give mean
a symbol and a letter to allow correla-
measurements based on 25 specimens,
tion of any peculiarities occurring and to
which, unfortunately, are not always use-
allow comparison of snails from different
ful in comparative studies.
localities and of different sizes. Snail
The general appearance of the shell
specimens from localities and W were
varies so much that several authors em-
serially sectioned, usually after having
phasize this variability as a feature of
been fixed in formalin. To prevent the
the species. Thus the shell is described
sand in the gizzard from damaging the
as pointed ovate (Mandahl-Barth, 1954),
microtome blade it is necessary to re-
somewhat obese (Connolly, 1939) and
move the sand or entire gizzard, as sug-
oval and bulging (Germain and Neveu-
gested by Carriker and Bilstad (1946). A
Lemaire, 1926).
few specimens from Wwere fixed in
The spire varies from slightly elevated
Bouin's Serial sections were made
fluid.
(Mozley, 1939) to rather high, about half
of entire snails and, occasionally, of iso-
the length of the aperture (Mandahl-Barth,
lated organs. A few slides were stained 1956).
with Masson's, but Mallory's triple stain
Mozley (1939) notes that the shell con-
proved to be more satisfactory.
sists of three whorls increasing rapidly
while Krauss (1838) and Germain and is described both as narrow (Germain
Neveu-Lemaire (1926) found shells with and Neveu-Lemaire, 1926; Mandahl-Barth,
as many as five whorls, but four to four 1956) and as broad and deep (Mozley,
and a half convex whorls is the usual 1959; de Azevedo et aL, 1957).
number (Mandahl- Barth, 1954, 1956; de From the above summary the fact
Azevedo et al., 1957). The sutures may emerges these descriptions differ
that
be very deep (Mozley, 1939; de Azevedo so much that they are often completely
et al., 1957), deep (Germain and Neveu- inadequate in species discrimination and
Lemaire, 1926), or fairly deep (Mandahl- the need for reducing verbal descriptions,
Barth, 1954, 1956). wherever possible, to concise mathemati-
The colour of the shell is described as cal terms becomes clear. Although sta-
yellowish reddish, pale, shiny (de Aze- tistics cannot, intaxonomy, play the de-
vedo et al, 1957), yellowish brown (Ger- cisive role which it does in experimental
main and Neveu-Lemaire, 1926), lighter work, it is nevertheless indispensable in
or darker dull brownish (Mandahl -Barth, bringing out correlations, differences,
1956) or pale or dark horny brown (Man- indications of variability, etc. In the
dahl-Barth, 1954). According to Germain present work the tatest and analysis of
and Neveu-Lemaire (1926) the shell is variance were not applied because the
rather solid while Mandahl-Barth (1954) samples were collected by different peo-
and Mozley (1939) regard it as ''rather ple at different seasons. Bearing in mind
thin walled" and "thin walled" respec- that "it is the naturalis' s original sin
tively. The sculpturing may consist of to pay more attention to very small, or
minute regular growth lines (Mozley, extra large or otherwise exceptional
1939) or rather coarse growth lines (Man- specimens" (Boycott, 1928, p. 28) our
dahl-Barth, 1956), in either case closely samples are most probably not random,
set. Krauss(1848) records that stria- and, except for those from localities W
tions on the last three whorls may, in and B, are not large enough to compare
juveniles, be covered by a thin membrane classes of equal size, as was done by
through which the costulations show. The Schutte and van Eeden (1959a) for Biom-
nature of this membrane is not very phalaria pfeifferi (Krauss). It was more-
clear. over not possible to compare data from
The aperture is oval (Germain and modal classes since in some localities,
Neveu-Lemaire, 1926; Mozley, 1939; de e.g. (Fig. 2), there were no specimens
Azevedo et al, 1957) and the labrum re- large enough to fall in the modal class.
gularly curved (Mandahl-Barth, 1956) In any case Table I reveals that there
and simple (Krauss, 1848; Germain and is so little difference between the mean
Neveu-Lemaire, 1926; de Azevedo et al., of the modal class and the mean of the
1957). Krauss (1848) states that the la- total sample that the latter has been used
brum is joined to the body whorl by a wherever it might be expected to throw
clear white shiny lamella, which, accord- light on some aspect of the shell or soft
ing to de Azevedo et al. (1957), is some- anatomy.
times indistinguishable from the labrum. It has often been suggested and it has
The columella is straight (Mandahl-Barth, in some cases been demonstrated, among

1954; de Azevedo et aL, 1957) and ac- others by Simroth and Grimpe (1918),
cording to Connolly (1931, 1939) it may Baker (1938), Rotarides (1932), Wesen-
also be slightly concave. The columellar berg-Lund (1939), Boettger (1944), Huben-
margin is generally widely reflexed (Man- dick (1941) and Schwetz (1954), that eco-
dahl-Barth, 1956; de Azevedo et aL, 1957) logical factors influence the moUuscan
and may either cover the umbilicus which shell in a number Unfortunately
of ways.
is open from below (Krauss, 1848) or the very features known to be affected by
may only partially cover the umbilicus ecological conditions are those most com-
(Mandahl-Barth, 1954). The umbilicus monly employed in taxonomic discrimina-
shape, thickness of the same general facies as juveniles in sam-
tion, e.g. size,
W and the average length is
I
shell, colour and sculpturing and our own ple (Fig. 2)
observations can do no more than to only 7.2 mm. As they come from the
underline the necessity of a closer in- same locality as sample A (Fig. 1) and
vestigation of the relation between eco- were collected on the same day, the dif-
logical factors and conchological features. ference in size between the samples
Shells of B. tropicus from nine locali- seems explicable only by assuming that
ties (all but "L") were treated statisti- the juveniles, in this case, occupied a
cally to discover correlations of changes particular part of the habitat, different
in ratios and also in shell form and to from that occupied by the adults. This
find out which of these changes were de- supposition further strengthened by the
is
pendent on increasing age of the snail. fact that the juveniles of sample W
are
With regard to size it seems important thickly covered with sediment, a condition
to know the maximum length attained by rarely found in adults and this, too, in-
any shell in the population, the mode of dicates that the juveniles may live under
the sample, which is often of more use conditions slightly different from the nor-
than the mean, and also the size range mal adult habitat. In this case they seem
of the sample. Such data for one sample to prefer a more benthonic life.
(W) are represented in Fig. 4. The Series indicates that size, which
largest shell in all our samples is 13.3 must obviously be influenced by eco-
mm but the modes of all the localities logical factors affecting growth, has an
combined are only 9.2 and 10.0 mm. important effect on the general shape of
The size frequency, in a series of 171 the shell. In young snails (type I, Fig. 3),
shells, determined to the nearest 1/10 of such as sample K, (Fig. 2) the whorls
a millimetre is shown in Fig. 5. These overlap each other greatly, resulting in
dimensions cannot, however, be regarded the relatively large mouth aperture and
as truly representative of B. tropicus as a low spire. Here the base of the colu-
the size irregularity revealed by graphic mella is straight In the mature forms
representations should seemindicate
to (Fig. 3)two further types (types II and
that the sampling was inadequate. The III) of shell are distinguishable, depending
snails in sample (Fig. 2) are all re- on the tightness of the coiling of the
garded as young because they have the whorls. In these older snails the whorls
TABLE I. Shell proportions in Bulinus tropicus. Comparison of the means and modes of modal
classes of certain ratios for the 9 samples (W - G) separately and combined (T).
FIG. 1. Photograph of representative specimens from localities B, M, S, A, and G.

w
# ^ ^
D
4'
L
I
J
2»
H
àà è f)
10mm
^ ^ i^
FIG. 2. Photograph cf representative specimens from localities W, D, L, H, and K.

MORPHOLOGY OF BULINUS TROPICUS il
do not overlap as much as in the juve- TABLE . Means of the ratios shell length
niles so that the height of the aperture to aperture height (L/AH) and shell length to
(AH) is relatively smaller when compared breadth (L/B)
with the total length of the shell (L).

These proportions in the adult result in
a higher value for the ratio L/AH (Table
II). Type represented by sample H
II is
(Fig. 2). These snails have tightly wound

whorls which are generally not highly
convex. The shell is elongated and the
umbilicus narrow. Although types I and
II differ in details from figures 4 and 14
(Physa zuluensis and Isidora compta) of

Melvill and Ponsonby (1903), type I agrees
with their Fig. 4, and t5e II with their
Fig. 14 (see Fig. 3). Type II is repre-
sented by sample (Fig. 1). This last
type has whorls which are more convex
than in the former two, and are not as
tightly wound around the central axis,
resulting in a more globose form with a
wide umbilicus.
It is often found that as the age of a
snail increases, its shell assumes a more

elongated form (Lais, 1925; Franz, 1928;
Degner, 1930, quoted from Neuhaus, 1952).
This observation is fully borne out, for
B. tropicus, by Fig. 8 in which the ratio
L/B (length/breadth) is plotted against L
(length of the shell) and which also shows
that,with regard to this ratio, only shells
of approximately equal length should be
compared.
8á STIGLINGH, VAN EEDEN AND RYKE
4-7 SPECIMENS 7-5 lOO

5HEL^ Êi^TH IN MM.
4.0 4.5
running and stagnant water respectively. findings illustrate the need of detailed
The globose, widely umbilicate type III field studies,which were not possible in
is found in dams while the elongate, nar- the present investigation as snails were
rowly umbilicate type II is found in slow- sent to this Institute from various parts
flowing streams. These findings are in of the country.
accordance with the view held by Sim-
roth and Grimpe (1918), Baker (1928) TABLE m. Mean ratios of shell length to
and other authors, that the stream forms spire (L/S) and the spire angles.
of certain gastropods are more elongated

than those living in stagnant water. The
two exceptions which do not fit into this
picture are shells from locality L (pools)
which show an elongated" stream" shape,
and shells from locality M
(a stream),
which show a squat "pool" shape. In the
last mentioned case the habitat contained
many waterplants and thus the flow of
water may have been hindered to such an
extent that the habitat in which the snails
lived may in reality have been a "pool"
habitat
Our deductions as to the presumable
influence of ecological factorson the
shape of the shell of B. tropicus do not
match with the findings of Boettger (1944)
working on various Lymnaea species.
That author was primarily interested in
the difference between lake forms in
stagnant water, and those subject to wave
action. Snails under the latter conditions
were found to be short spired and obese,
thus agreeing with what, in B. tropicus,
seems to be the stagnant water form.
Boettger also found that stream forms
had rounded apertures, once more a char-
acter apparently correlated with a pool
habitat in B. tropicus. These apparently
conflicting results may, perhaps, be as-
cribed to the fact that the effects of run-
ning water and waves, on the snail, are
probably not the same. These diverse
both in the entire series and in each
sample separately. In spite of this vari-
ation there is an obvious negative corre-
lation (Fig. 9) between this ratio and the
length of the shell, proving that the small-
er forms have relatively shorter spires.
Mandahl-Barth (1956) gives the length of
the spire as about half the height of the
aperture. From Table I it can be seen
that, inour series, the ratio of spire to
height of the aperture, excluding samples
D and (Fig. 2) which have unusually
short spires, is 0.2, i.e. only about one
fifth of the aperture height. This dis-
crepancy between our own and Mandahl-
Barth's (1956) values may perhaps in
part be explained by the fact that it is
not quite clear how he measured the
spire. The angle of the spire, measured
as shown in Fig. 13 appears to be too
variable to be of any use in taxonomy.
From Table II it is apparent that shell
types with spires as different in shape
and length as and A have spire angles
of almost the same magnitude, while the
total range of the whole series is 58° -
1340.
TABLE rv. Labrum depression angles in the

shell of Bidinus tropicus.
Sample
cipal dam and from temporary pools

(D) margin is broadly reflexed and its lower
at the township (L).
native The shells half is generally free and either covers
from locality W
are elongated and fairly the umbilicus when it is narrow, as in
smooth, whereas those from localities D type II, or only partly covers the umbili-
and L are more squat and have coarse cus when it is wide and deep as in type
transverse striations or ridges; attention III. The callus is usually well developed
though, should also be focussed on the and of the same colour as the rest of the
sporadic occurrence of smooth forms in shell. It may also be either transparent
a sculptured population. The sculpturing and indistinct or white and sharply con-
in these three samples can be correlated trasting.
with the shape of the shell: the elongated In sample the labrum sometimes
stream forms (type II) are smooth, while meets the body whorl in a very rounded
the squat "pool" forms (type III) are angle, the aperture margin being slightly
ribbed. These findings agree with the expanded like a trumpet This condition
observations of Haas (1922) on various presumably represents that described by
Naiads and on the prosobranch snail Vivi- de Azevedo et al. (1957). The angle of
parus costatiis Quoy and Gaimard, in depression of the labrum was measured
which stream forms are smooth while by the method indicated in Fig. 14 and
the lake forms are sculptured. was found to be surprisingly
constant
The varying aperture shapes in our (see Table IV), the mean varying by
about
series can be classified into three types one degree only in all localities (57. 1° -
coinciding with the three general types 58.8°) except for H (61.7°) in which the
of shell referred to above (see Fig. 3, shells are large and elongated and the
I, II, III). Fig. 10 reveals that the ratio labrum is more depressed than usual,
aperture height to aperture breadth does thus producing a rather streamlined shell.
not change as the snail becomes older
and that the mean fluctuates between 1.4 TABLE
and 1.5 for all samples, except for the
juveniles from locality in which the
ratio is 1.3. The graph plotted in Fig.
II, which reflects the relation between
shell length and aperture height, plotted
against the entire length of the shell,
shows a correlation, indicating that as
the snail becomes older the mouth aper-
ture becomes relatively smaller. Two
additional ratios which might prove to
be of taxonomic value are (a) shell
breadth to aperture breadth (B/AB) and
(b) aperture height to shell breadth
(AH/B). The corresponding data for our
specimens of B. tropicus are tabulated
in Tables V and VI. Both the ratios
B/AB and AH/B are fairly constant (see
also Table I), the low value of B/AB for
snails from locality being due to the
relatively large mouth aperture of im-
mature forms. The shape of the aper-
ture is determined by the shape
partly
of the columella which is usually straight,
slightly concave or concave in the three
shell types respectively. The columellar
13 :
J4
15
16
the closer correlation. In both cases the tentacle (Fig. 19) and not at its outer
scatter was so great that correlation was side as mentioned by Mandahl- Barth
scarcely evident. This lack of corre- (1954). Lateral processes occur at the
lation must partially be ascribed to the base of the tentacle in the Bulininae, Pla-
fact that there were not enough of the norbinae and Ancylidae (Hubendick, 1948).
smaller shells. In B. tropicus there are two unequally
developed processes, the dorsal tentacular
process (D.T.P., Figs. 18 and 19) being
EXTERNAL MORPHOLOGY
more prominent then the ventral one
The animal has been described as blue- (V.T. P., Fig. 18). Between them they
grey by de Azevedo et al. (1957) and, enclose a ciliated groove, thus creating
although this is true of preserved speci- a condition resembling that found in spe-
mens, live animals are yellowish- or cies of Miratesta and Protancylus (Sim-
reddish brown as a result of the haemo- roth, 1928).
globin in the blood. Living specimens of The gill (pseudobranch) (G), anal lobe
B. tropicus have small white flecks due (A. L.)and pneumostome (PN) are situated
to concretions in the tissue such as occur on the left hand side of the animal in the
in other planorbids (Baker, 1945; Abdel- sharp angle formed by the junction of
Malek, 1952). The tentacles are filiform the labrum of the shell with the body
and circular in cross section although, whorL The natural relations of the vari-
in his 1946 publication on the anatomy ous structures are most readily under-
of Bulinus Hubendick regards the tentacle stood by referring to Fig. 19. The pneu-
as an outgrowth of the post-tentacular mostome is large and bounded ventrally
process and the latter as the homologue by a small lobe referred to as the infe-
of the flattened triangular tentacles of rior palliai lobe, mantle lobe or lower
the Lymnaeidae. Hubendick' s (1948) fig- mantle lobe by Hubendick (1946, 1955)
ure of a cross section of the tentacle of and as the pulmonary siphon by Watson
B. (Physopsis) africanus (Krauss) (Plate (1925) and pulmonary siphon and pneumo-
2, Fig. 8) differs from ours for B. tropi- stome by Baker (1945). The shape of the
cus (Fig. 15) in being more solid, perhaps siphon varies constantly in a living animal
due to contraction. Throughout the length where it is alternately protruded and
of the tentacle of the latter snail there retracted to collect air from the surface
is one central (C) and a number of peri- and then to convey it to the mantle cavity.
pheral cavities (P.C.) in the loose mus- The rectum opens at the tip of the anal
cular tissue which is surrounded by a lobe which is flattened dorso-ventrally
basement membrane (B.M.) supporting and lies mediad to and slightly in front
the external epithelium (E). The eyes of the base of the gill. This region
are situated medially at the base of each agrees more closely with Watson's (1925)
FIG. 12. Diagram illustrating the method of measuring the length (CD) of the spire.
FIG. 13. Diagram illustrating the method of measuring the angle (CA'D) of the spire.
FIG. 14, Diagram showing the method of measuring the angle (CFE) of labrum depression.
FIG. 15. Representative transverse section of tentacle at point anywhere between base and tip.
FIG. 16. Transverse section of a kidney lamella.

FIG. 17. Oblique section through the osphradium, situated on collar, outside mantle cavity.
FIG. 18. Transverse section of the post- tentacular processes.
FIG. 19. Dorsal view of the gill (pseudobranch) region.
FIG. 20. View of ventral surface of roof of mantle cavity with attendant structures.
See list of abbreviations p. 114

figure of Physopsis globosa (Morelet) of the same thickness throughout The
{-Bulinus globosiis) than with Hubendick's distal dilatation shown
Plate XXII by
in
(1955) figure of B. tropicus in that the de Azevedo et al. (1957) is probably un-
anal lobe seems narrower and the rectal usual. In the snails we examined the
ridge more pronounced. The gill (G) is a saccular part is more distinct from the
U-shaped structure thrown into a number tubular part than appears to be the case _
of large transverse folds alternately pro- in the figures of either Hubendick (1948) I
truding on the dorsal and ventral surfaces. or Mandahl- Barth (1946). The saccular
These could be called the primary folds portion possesses internal transverse
(P. F.) of which there are 4 or 5 on each ridges or lamellae (Fig. 16) which are J
surface. Each of these primary folds more pronounced in the tubular region. '
consists of a number of smaller second- The ureter is usually smooth internally

ary folds (S. F.) running in the same di- but a few small lamellae may occur.
rection so that the large surface area, The lamellae may be fairly large but are
the thin epithelium and large blood spaces never solid. Each lamella consists of a
allow for the maximum oxygenation of double layer of epithelial cells (E) which
the blood. As the mantle cavity of Buli- sometimes appear to be pseudostratified
nus is generally filled with air, an os- and are connected by fine, sparse con-
phradium (O, Fig. 20) in its primitive nective tissue fibres among which numer-
position would be functionally useless in ous pigment cells may occur. A base-
this species where, in fact, it lies out- ment membrane (B.M.) is just visible
side the mantle cavity on the collar, just underlying the epithelium. The whole
medially to the pneumostome, and not just structure is much less solid than that
within the opening of the mantle cavity figured by Abdel-Malek (1952) for Heli-
as claimed by Hubendick (1948). More- soma corpulentwn (Say), North American
over this organ is much smaller than planorbid. Histologically the proximal
the one figured on page 30 by that author region of the tubular portion of the kid-
and agrees better with his earlier de- new differs from the distal region. In
scription (1947) and withLacaze-Duthier's the former, cells are large, almost glo-
figure of the osphradium of Planorbarius bose, possessing conspicuous round bodies
corneus (Linné) reproduced in Boettger (U.C., Fig. 16)which are subsequently
(1944, Fig. 95). A cross-section of this set free into the lumen of the kidney, as
organ in B. tropicus is depicted in Fig. in certain Stylommatophora (Simroth
17. 1928), while distally the cells are much
smaller and narrower. We
found no trace
of a longitudinal muscle on the ventral
PALLIAL ANATOMY
surface of the kidney.
The mantle which only slightly overlaps Hubendick (1955) contends that the mea-
the edge of the labrum has
of the shell surement of organs is useless taxonomi-
conspicuous black patches concentrated cally as the size may be affected to a
mostly over the kidney region. varying degree by different types of fixa-
tive. Although it is true that slight
The renal system
changes most probably do occur, and
The kidney (Fig. 20) is well developed also that not all the snails are relaxed
and consists of a proximal saccular (S.K.) to the same degree, we cannot but support
and a distal tubular portion (..). The Schutte and van Eeden (1959a) in their
saccular part lies on the right hand side rejection of Hubendick's (1955) statement.
of the animal and posterior to the heart, The kidney (K) and ureter (U) of B. tropi-
while the tubular part roughly follows cus were measured as already described
the border of the mantle cavity to the under Material and Techniques and the
left hand side where it is reflected later- ratio K/U calculated for 153 snails. Al-
ally as the ureter (U), which is generally though the variation in each sample is
great, the values ranging from 2,0 - 4.6 thelium while the sides are clothed with
(Fig. 7), the different means show a fair cuboidal cells, we assume that the ridge
degree of constancy. The mean for all is in reality ciliated as it is in S. (Phys-
the snails was while the mean for
2.9, opsis) africanus (Hubendick, 1948). The
30 snails of 7.7 mmand less was 2.7, length of this ridge relative to that of
and for 25 snails of more than 10 mm the nephridium is used by Mandahl- Barth
it was found to be 2.8, In other words (1954, 1956) for distinguishing certain
there is a random variation of these three Bulinus species from each other. That
values which indicates that the ratio does author (1956) records that the intermedi-
not increase with age. ate mantle ridge is much shorter than the
kidney in the short spired species of
The mantle ridges
Bulinus s. s. Although we found this ob-
A feature of the mantle which is con- servation to be true, the ridge in our
sidered to be of systematic importance own series of snails is longer than that
in Planorbid snails (Mandahl- Barth, 1954, figured by Mandahl- Barth.
1956) is the presence of ridges which The different means of the ratio: length
occur on the ventral surface of the man- of kidney (excluding ureter) to intermedi-
tle and along the rectum. The confusion ate mantle ridge, calculated separately
resulting from the nomenclature employed for the nine samples studied, varies from
in describing these folds is dealt with in 1.4 - 1.9, while the mode and the mean
great detail by Schutte and van Eeden of all the localities are 1.6 In 1954
(1959a) and need not be recapitulated. Mandahl- Barth expressed the opinion that
In B. tropicus there are only two ridges the intermediate mantle ridge extends
the lateral rectal ridge (L.R.R., Figs. posteriorly as far as the mantle edge
19, 20) and the intermediate mantle ridge while he subsequently (1946) figures the
(I.M.R., Fig. 20), the renal and the median ridge as ending between the rectum and
rectal ridges being absent. A well de- kidney. In our series the former condi-
veloped lateral rectal ridge is described tion is always realised. The rectal ridge
for the Planorbidae in general (Baker, bears cilia, as does the intermediate
1945), for Physopsis globosa (=Buliniis mantle ridge, and the epithelium of these
globosas) (Watson 1925) and for various ridges, of the collar and of the sole of
bulinids (Hubendick, 1955). In 5. tropicus the foot resemble that shown inBoettger's
this ridge originates as the flattened edge (1944) Fig. 44 for the marginal dorsal
of the anal lobe (A.L., Fig. 19) and is epithelium of the foot of Plmwrbarius
continued posteriorly to terminate against CO meus (L.).
the roof of the mantle cavity (M. CR.,
The circulatory system
Fig. 20) near to its posterior margin.
At this point it meets the intermediate The blood vessels of the mantle are
mantle ridge (I.M. R.) which runs anteri- depicted in Fig. 20. An account of these
orly along the mantle between the rectum structures in the Planorbidae is given
and the kidney, to end near the thickened by Baker (1945) and Abdel-Malek (1952).
mantle collar (COL). The degree of de- The latter notes that the pulmonary vein
velopment of the intermediate mantle (P.V.) supplies the gill (G). This is
ridge, especially that part near the man- surprising since the blood in this vein
tle collar, is variable, but it is neverthe- should flow into the auricle (A) instead.
less always distinct. No cilia were seen From a dissection of B. tropicus it ap-
on this ridge in our specimens, but this pears that the gill is drained by the renal
fact may perhaps be attributed to the vein (R.V.) which proceeds posteriorly
prolonged preservation, because the ma- between the kidney and the ureter (U).
terial was a few years old when dissected. It seems logical to assume that blood
However, seeing that the top of the ridge from this vein flows through the large
is covered by a layer of columnar epi- vessels of the kidney into the pulmonary
vein (P.V.) so that it would be more true no rbinae the jaw and the radula are taxo-
to say that the latter drains the gill. nomically important (Baker, 1945) and
The blood containing corpuscles which the salivary glands are sometimes de-
become red in Mallory's stain passes scribed. The jaw (J, Fig. 24) of B.
from the pulmonary vein into the auricle tropicus is described and figured by
and thence into the less delicate ventricle Hubendick (1948) and de Azevedo et al.
(V) which has internal muscle strands (1957). It forms the dorsal bar of the
(M.S., Fig. 22) resembling those found T-shaped mouth and, in specimens exam-
in the prosobranch snail Thiara granifera ined by us, it differs markedly from that
(Lamarck) (Abbott, 1952). From here the described by either of the authors men-
aorta () supplies the structures indi- tioned. Whereas Hubendick (1948, 1955)
cated in Figs. 21 and 23. states that the jaw of S. tropicus is weak-
The pericardial cavity (PER, Fig. 22) ly developed and without lateral jaws
is frequently filled with round, white (L.J.)>we found it to be well developed in
structures thought to be the eggs of an all the specimens examined and composed
unknown organism (EG). Although, in of three elements as in other species of
this instance, they were observed to be Bulinus. The dorsal horizontal portion
free from one another, they are probably (superior jaw) is generally broader dorso-
identical with the strings of unidentified ventrally, than indicated in Hubendick' s
spherical bodies mentioned by Schutte and figure (p. 30, 1948) and snails from all
vanEeden (1959b) inBiomphalaria pfeifferi nine localities possess lateral jaws.
and also by Baker (1954). Cross-sections These are attached to the two extremities
of these eggs are depicted in Fig. 27. of the superior jaw and are folded behind
to meet in the midline whence they de-
INTERNAL ANATOMY scend vertically to form the upright por-
tion of the mouth. Admittedly the lateral
Jaw and R adula jaws may be poorly developed and they
In this system attention is focussed on may even appear to be absent in cases
only those features which are believed to where they are folded in by retraction
be of taxonomic importance. In the Pia- of the buccal mass, as occurs in snails
FIG. 21.
FIG.
killed in boiling water. De Azevedo et al. TABLE VIII. Height of the crown of the cen-
(1957) describe the jaw as biconvex, a tral and first lateral tooth inmm, calculated
from the published figures of other authors
shape assumed by none of the jaws of
and from two of our own specimens.
our series.
The length/breadth ratio of the radula
was found to be fairly constant, both for Author
snails from different habitats and of dif-
ferent sizes (Table VII). That this con-
stancy is of taxonomic importance has
TABLE Values of the ratios length to

VII.
breadth (L/B) of the radula in the samples
separately, combined (T) and for snails with
shells of 7.7 mm
and less and with shells
larger than 10.0 mm.
Sample n Range Interval Mean
W 46 2.1 - 3.0 0.9 2.5

D 11 2.2 - 3.1 0.9 2.5
31 2.1 - 2.7 0.6 2.4
S 6 2.3 - 2.6 0.3 2.4
7 2.3 - 2.9 0.6 2.6
A 9 2.3 - 2.7 0.4 2.4
H 8 2.2 - 2.9 0.7 2.6
M 10 2.2 2.7 0.5 2.4
G 9 2.2 2.8 0.6 2.4
L. 1.1 : 30 2.3 - 2.9 0.6 2.5
> 10 mm 24 2.2 - 3.1 0.9 2.5
134 2.1 - 3.1 1.0 2.5
n = number of specimens
not been proved and the data are given

merely facilitate future comparative
to
studies. In the case of Biomphalaria
pfeifferi this ratio was found to vary be-
tween 2.15 and 3.09 (Schutte and van
Eeden, 1959a), whereas inBidinus tropicus
it varies between 2.1 and 3.1. When
dealing with the Bulininae, Mandahl- Barth
(1956) emphasizes the importance of the
relative size of the teeth and, in certain
cases, the shape of the first lateral, while
he apparently finds the shape of the mar-
ginals more important as a means to
characterize the different species of the
Planorbinae. In describing B. tropicus
he refers to the radula merely by saying
that the teeth are rather small. From
the data in Table VIII it is seen that,
considering their small size, the size of
may also be a small denticle between the of the truncatus group which might indi-
two main cusps. In all the specimens cate either that the Grahamstown speci-
of 5. tropicus examined under oil immer- mens do not belong to the tropicus group
sion the sides of the crown have a more at all, or that the shape of the mesocone
or less fluted (Fig. 30) appearance, al- does not always constitute a reliable ba-
though it may not always be very evident, sis to distinguish between the tropicus
a circumstance which probably explains and truncatus groups of Bulinus as claim-
why this feature, which also occurs in ed by Mandahl-Barth (1956). However it
other types of tooth, has not yet, to our might perhaps be premature to suggest
knowledge, been recorded in the litera- the latter explanation. In all samples
ture. The crown may also be relatively the base of the lateral tooth is relatively
smaller when compared with the base and smaller than that of the centrals and the
this always appears to be the case when tooth is not like any figured in the litera-
the central is slightly out of focus on ture. It approximates that shown by
account of its having been pushed out of Mandahl-Barth (1956) but actually it is
position by pressure on the coverslip. closest to Schutte and van Eeden's (1959a)
The shape of the base differs from that figure for Biomphalaria pfeifferi. Pro-
shown by Cawston (1925, 1926). Nowhere ceeding laterally in the radula the teeth
did we see the rounded corners of the become progressively more obliquely
broad end of the base figured by that placed and the ectocone comes to lie
author. In some cases the base ap- lower than the endo- and mesocone.
proaches the trapezoidal shape shown by Gradually the last- mentioned two cusps
Mandahl- Barth (1956) although it is usu- are replaced by 5 - 9 denticles while
ally more horned, resembling the figures 1-5 denticles appear laterally at the
by Connolly (1939) and de Azevedo et al. base of the ectocone. At the same time
(1957), and yet not quite like either. the tooth becomes more elongate till it
The laterals (Figs. 30, 33) are larger assumes the shape typical of a marginal
than the centrals and almost invariably (Fig. 33). There is no clarity or uni-
have a tiny denticle at the base of the formity as to how the various authors
mesocone (ME), either on one side or distinguish between lateral and marginal
on both. The first lateral is a broad teeth, and as the transition is so gradual
tooth, fluted medially, while the cutting and variable, we have for the sake of
edges of the ecto- (EC) and endo cones objectivity, regarded the first marginal
(EN) adjacent to the mesocone may be tooth as one in which the endo- and meso-
concave, straight or convex. The mesocone are represented by at least three
cone is almost never simply triangular denticles of approximately equal size or
as is claimed by Mandahl-Barth (1956). four denticles which may be of unequal
Usually the two sides converge slowly size. In counting the teeth we therefore
from the base and then suddenly slope located the first marginal and then pro-
more sharply towards the apex, the la- ceeded medially counting the teeth be-
teral side of the triangle being longer tween this and the central, thus grouping
than the median. In this respect the the atypical laterals or so called inter-
mesocones differ from the mesocone de- mediate teeth of Schutte and van Eeden
scribed and figured by Mandahl-Barth (1959 a) and de Azevedo et al. (1957)
(1956) for the B. tropicus group and yet which, in B. tropicus, vary in number
by reason of the more acute apex of the from 0-2, with the typical laterals.
cusp and the longer lateral side of the This procedure is also reflected in the
triangle they are usually distinct from radula formula. A distinction between
those found in the B. truncatus group. what is typically lateral, intermediate
The radulae of snails from locality G and typically marginal is not feasible in
(Fig. 34) are an exception to this in that B. tropicus where the transition from the
,
their mesocones strongly resemble those first to the last mentioned is much more
31
FIG. 30. First lateral tooth illustrating the various parts.
FIG. 31 - 32. The two types of central tooth of the radula, with and without central denticle.
FIG. 33. Central tooth and half of a transverse row (1 - 29) of a specimen from sample B.
FIG. 34. First lateral teeth of specimens from sample G.

gradual than in Biomphalaria pfeifferi. in the formula has been proved to in-
The marginals (Fig. 33) are elongated crease with the age of the snail in the
and obliquely placed. The tooth is fluted case of Biomphalaria pfeifferi (Schutte
medially and the denticles occasionally and van Eeden, 1959a). Correspondingly,
have bifid tips. The denticle which rein B. tropicus the mean number of rows
presents the mesocone of the lateral teeth for snails having shells of 7.7 and mm
is broader and higher than the other den- less is 113, whereas the mean number
ticles. Sometimes it is evenly rounded of rows for snails having shells larger
laterally, but more typically it is angular. than 10 mm is 134. Bearing in mind
Lateral to the ectocone there are up to the shortcomings of any such formula,
5 small denticles which may increase in we nevertheless offer one as it may, in
number so as to produce a serrated la- certain respects, be useful. From our
teral margin. The crowns of the extreme own investigations of 92 radulae from 9
marginals have reduced denticles and localities and from a random selection
occasionally unicuspid marginals, similar of a few from each locality in which the
to those described forS, pfeifferi (Schutte relative numbers of teeth were counted,
and van Eeden, 19 59a), seem to occur. a representative formula for B. tropicus
On close inspection, however, these most- seems to be 1 6+2 24 x 123. In this
: :
ly turn out to be ordinary marginals seen formula the number 2 represents the
edge-on. The base of the marginal tooth intermediate teeth which are neither typi-
is more elongated than any figured in
cally lateral nor typically marginal.
current literature. Besides the teeth
described above, a variety of abnormally
The Salivary glands
shaped teeth may occur throughout a lon-
gitudinal row but these are easily recog- The salivary glands(Fig. 25) are very
nised as anomalies and merit no further uniform Planorbidae (Baker, 1945),
in the
attention. where they are usually long and cylin-
Several authors have thus far given drical Contrary to Hubendick's (1948)
radula formulae for Bulinus tropicus but account of the condition in B. tropicus,
these, unfortunately, are not always in in which he states that the glands do not
concurrence with each other. Thus, pass through the nerve ring, they do pass
Connolly (1939) gives the formula as through it, narrowing considerably as
1:8:22 x 90+n, and that given by de Aze- they do so and then becoming larger pos-
vedo et aL (1957) is 1:25 x 121. Cawston teriorly. The end of the loop is attached
(1925, 1926) gives no formula but sets to the oesophagus or body wall or both
the number of laterals at 8. The limi- of these, and at this point receives a
tations of these formulae and the possible blood vessel. Although they may some-
causes for the different results have al- times have slight constrictions they are
ready been dealt with by Schutte and van not moniliform as in the case of Physop-
Eeden (1959a) whose views are fully borne sis globosa (=B. globosus) (Watson, 1925).
out by our investigation of B. tropicus. Except for the fact that there is not much
While 8 laterals seem to be a constant more of the gland behind the nerve ring
mean, the actual number varies from one than there is in front of it. the lengths
radula to another, and even from one being about equal, Watson's (1925) de-
transverse row to the other. The number scription is applicable to B. tropicus.
of marginals to a transverse row like- Histological detail of the gland is shown
wise oscillates greatly and, moreover, in Fig. 26. It appears that the structure
increases with the age of the snail. In is more or less the same throughout the
B. tropicus the issue is further compli- length of the gland, except at the very
cated by the fact that there is no satis- thin part passing through the nerve ring,
factory criterion for distinguishing be- where the cells do not show signs of such
tween the various t5es of tooth. Finally active secretion as in the parts with
the number of transverse rows reflected larger cells.
Nervous system by Boettger (1944). That author reported
seven pairs of peripheral nerves, origi-
At the higher taxonomic levels the ner-
nating from the cerebral ganglia: nervus
vous system is generally considered reli-
opticus, n. staticus, n. tentacularis, n.
able for comparative studies. It was
fronto -labialis superior, n. labialis médi-
unfortunately not possible for us to study
the reports of the nervous system in vari-
us, .labialis minor, n. nuchalis and an
ous Basommatophora as presented by

additional unpaired .
penis on either the
right side in dextral forms, or on the left
Lacaze-Duthiers (1872), Pelseneer (1895),
side in sinistral forms. In the B. tropicus
and Elo (1938) but accounts or short
snails examined we recognised only six
notes are alsofound in Watson (1925),
(not seven) paired nerves from the cere-
Simroth (1928), Boettger (1944), Baker
bral ganglion, which did not arise in the
(1945) and Hubendick (1947, 1948). Some
order mentioned by Boettger (1944). The
of Pelseneer' s figures are reproduced
first nerve to arise from the dorsal lobe,
in Boettger (1944) and Simroth (1928)
(from front to back) is the stout tentacular
while the last mentioned author also re-
(T.N.) which proceeds to the base of the
produces figures by Lacaze-Duthiers.l
tentacle where it joins a ganglion which
Hubendick (1947) is of the opinion that,
Simroth (1928) describes as the cup-shaped
allowing for the differences resulting
ganglion. In B. tropicus the ganglion is
from being coiled sinistrally ordextrally,
elongated and grooved laterally in con-
the nervous system of the Planorbidae
formity with the post-tentacular groove.
shows a maximum agreement with that
The fine ocular nerve arises just posterior
of the Lymnaeidae and Physidae, in which
to the tentacular nerve and agrees entirely
the system consists of a concentrated ring
with Simroth' s description of it. A third,
of ganglia surrounding the oesophagus.
extremely delicate nerve originates at the
There are many designations for the pos- posterior edge of the dorsal lobe of the
terior three ganglia of the ring. However,
dorsal lobe of the cerebral ganglion and
the most commonly used appear to be
innervates the statocyst (ST) associated
parietal and visceral for the lateral and The
with each pedal ganglion (PE. G).
median ganglia respectively. inferior (ventral) cerebral lobe also gives
A semi-diagrammatic sketch of the rise to three nerves. There are two (not
circumoesophageal ganglionic ring in B. three) labial nerves (L.N.), of which the
tropicus, indicating the major nerves more slender anterior one branches in
issuing from the ganglia, is given in Fig. two. On the left side only, at the point
35. The ganglia are paired except for or origin of the more posterior labial
the hindmost, median visceral ganglion. nerve, and before the last of the paired
Each cerebral ganglion (CG.) is composed nerves, arises the penial nerve so that
of a large dorsal and a smaller ventral the left cerebral ganglion has seven
lobe, though variations occur not only as nerves. The last nerve is about as fine
regards the general shape of the ganglia as the one supplying the statocyst and
which sometimes possess numerous small proceeds mediad to innervate the walls
lobes, but also in the surface which may of the cephalic artery.
be quite smooth or have the appearance Finally the cerebro-buccal connective
of a mulberry. These ganglia and the (C.B. C., Fig. 25) arises from the inferior
nerves issuing from them differ in a cerebral lobe on either side. The inter-
number of details from the generalized buccal connective is so short that the
description for the Basommatophora given buccal ganglia (B.G., Figs. 25 and 29) may
^The nervous system among others, of a related species of Bulinus has been described in detail by
Demian (1960, Morphological studies on the Planorbidae of Egypt I. On the macroscopic anatomy
of Bulinus (Bulinus) trunca tus (Audouln). Ain Shams Sei. Bull. No. 5 (Monograph), 84p,, 9 pi..
Faculty of Science, Cairo). ED.
.
be contiguous. The origin and distribution literature, where it is stated that the gill
of tlie buccal nerves (Fig. 29) agree with is supplied by the left parietal ganglion,
Watson's (1925) findings. while only a thin connection leads from
The pleural ganglia (PL.G., Fig. 35) that ganglion to the combined gill and
agree with Boettger's (1944) account in collar nerve in B. tropicus. The uterine
being small and in having no peripheral nerve (U.N.) just antero-dorsal to the gill
nerves. nerve, runs between the uterus and colu-
The parietal ganglia (PA.G., Fig. 35) mellar muscle and sends small branches
are asymmetrical, the left being at least to both, as well as a small branch to the
twice the size of the right. As in Boett- prostate gland. It also has a very fine
ger's (1944) description for Platwrbariiis branch supplying the cephalic artery.
corneus (L.) (after Merker), the right The third nerve (B.R.N.), which is also
parietal ganglion in B. tropicus has only very fine, arises from the right side of
one nerve, the collar nerve (C.N'.). This the ganglion and supplies the right retrac-
nerve also splits into two, the anterior tor muscle of the buccal mass.
branch innervating the anterior part of The pedal ganglia (PE.G.) are almost
the collar and the other branch curving as large as the cerebral and agree with
around to that part of the collar which Boettger's (1944) account in having six
overhangs the foot posteriorly. Boettger' s nerves. Three of these (PE.N.) are stout
descriptions of the left parietal and the and innervate the anterior, median and
visceral ganglia do not coincide with the posterior portions of the foot, while three
conditions obtaining inS. tropicus, where more slender dorsal nerves (LAT.N.)
each of these ganglia has one nerve less; supply the body wall, generally at the
the left parietal ganglion has three nerves. level of the lateral muscle bands. In
The largest of these (OS.N.) corresponds addition to the ganglia already mentioned
to the single nerve of the right parietal there is, situated posterior to the inter-
ganglion and has two main branches, one pedal connective, a minute little ganglion
(PN.N.) innervating the dorsal lip of the which sends a few fibres to the foot. A
pneumostome and giving off a tiny branch conspicuous white statocyst (ST, Fig. 28)
to the ureter, while the more ventral containing highly refractile crystals is
branch (OS.N.) supplies the osphradium placed postero- dor sally on each of the
and the collar. The remaining two pari- pedal ganglia.
etal nerves are very fine, the median,
or body wall nerve (B.W.N.) arising be- Reproductive System
tween the parietal and visceral ganglia
and running backwards along the cephalic The general appearance of the genital
artery to supply the anterior body wall system is depicted in Fig. 41.
at the level of the base of the uterus,

Hermaphrodite System
while the lateral nerve joins a branch
(G.N.) from the stoutest and most pos- The histology of the hermaphrodite
terior abdominal nerve at the level of the gland has been described for various
collar. snails such as the Pulmonata generally
The single visceral ganglion bears three (Simroth, 1928), the Basommatophora
nerves. The stoutest of these passes (Boettger, 1944), some Planorbidae (Ab-
just posterior to the vagina and then splits del- Mai ek, 1952), Bulimis (B.) contortus
into three, a small branch (S.N.) going Michaud (=S. truncatus Audouin) (Laram-
to the siphon (ventral lobe of the pneu- bergue, 1939) and for B. (Physopsis)
mostome), a branch (G.N.) supplying the jousseaumei (Dautzenberg) (Wright, 1957)
branch The general morphology of the ovotestis
median part of the gill, and a third
(C.N".) supplying the collar posterior to of B. tropicus agrees with that of Phy-
the gill. In this respect the situation sopsis globosa i^-Bulinus globosus) fig-
differs from the descriptions seen in the ured by Watson (1925), and with Abdel-
AL.GL
M. GL.
Malek's (1952) description for Helisoma bodies, one slightly smaller and staining
trivolvis Say. The number of acini (AC, more deeply than the other.
Fig. 41) most commonly varies between Regarding the pulmonates, Simroth
14 - 16 in the longest of the longitudinal (1928) claims that as soon as the undif-
rows, of which there are 3 - 4 at the ferentiated cells of the germinal epithe-
widest part of the gland. Only the long- lium become differentiated into male ele-
est of the longitudinal rows extends as ments they leave the syncytium and com-
far as the apex of the organ. The acini plete their development in the lumen of
are arranged with their narrow bases the acinus. This observation is apparent-
opening into the tubular common atrium ly supported by Abdel-Malek (1952) who
(C.A.) which is continuous with the her- mentions sperm bundles occurring free
maphrodite duct (H.D.). in the lumen in Helisoma trivolvis. Our
As noted by Abdel-Malek (1952) the own impression is that, in B. tropicus,
ovotestis is surrounded by a pigmented the sperm bundles (S.B., Fig. 37) may
connective tissue layer which is most either be free in the lumen or attached
densely pigmented over the apices of the to the basal or Sertoli cells, a view also
acini. The germinal epithelium of B. put forward by Wright (1957), although
(B.) tropicus lines only the basal portion transverse sections through the basal
of each acinus (Fig. 37) so that in this part of the acinus reveal the basal cells
respect it agrees with B. (B.) contortus (B.C., Fig. 36) attached to its walls, thus
(Larambergue, 1939) and B. (P.) jous- also lending support to Boettger (1944)
seaiimei (Wright, 1957), but differs from who maintains that the basal cells never
both Planorbis planorbis (L.), a pale- float in the lumen. The spermatogonia
arctic planorbid, where the germinal (SO) spermatids (STD) and sperm cells
epithelium is restricted to the common (SP) are easily recognisable as such and
atrium (Boettger, 1944) and Arianta ar- present no noteworthy features.
bustorum (L.), a stylommatophoran snail, The hermaphrodite duct (H.D., Fig. 41)
where this epithelium seems to be present is tubular both proximally, i.e. nearest
in the apical portion of the acinus as to the ovotestis, and distally, but the mid-
well (Simroth, 1928, Fig. 205A). Like dle region has closely arranged irregular
Larambergue (1939) in B. contortus, we seminal vesicles (V.S.), and is convoluted,
can see no differentiation of the ovotestis so that the whole appears as a solid and
into male and female zones, the two kinds rather wide structure. Where the her-
of sexual element occurring intermingled. maphrodite duct splits into the male and
The oocytes (,
Figs. 36 and 37) have female ducts, which are completely sepa-
double nucleoli (N1), a condition also oc- rate in this case, it receives the duct
curring in species of the terrestrial pul- from the albumen gland (AL. GL.).
monates Helix 2ináArion{Ancel and Lams, The mutual relations of these ducts
quoted from Simroth 1928). Each nucle- are shown in Fig. 38. Basically they
olus consists of two opposed spherical agree with the figures for B. contortus
FIG. Dorso-lateral view of the circumoesophageal ganglionic ring and its main associated
nerves (diagrammatic).
Section showing oocyte and sperm in apical part of acinus.
Section of an acinus of the ovotestis.
Sketch of the carrefour region (dorsal view).

Section of a penis in the spermatheca.
Drawing of the dorsal surface of the uterus.
FIG. 41. Illustration of the reproductive system (excluding the penial complex).
.
by Larambergue (1939) and for Biom- the uterus as representing a part of the
phalaria pfeifferi by Schutte and van oviduct and designate it as the dilatation
Eeden (1959b), both belonging to Ethiopian of the oviduct, although most authors re-
planorbid subfamilies, but differ from gard it as part of the uterus. This last-
that given by Abdel-Malek (1952) for Heli- mentioned organ is broader proximally
soma trivolvis, a nearctic planorbid. In than distally and lies just below the floor
B. tropicus the duct of the albumen gland of the mantle cavity and, like the albumen
is short and opens into a pear-shaped gland, is slightly concave ventrally and
sac, the carrefour (CAR), embedded in convex dorsally. The oviduct which en-
the hilus of the albumen gland. This sac ters it at the left posterior margin is
is lined by ciliated cuboidal cells with surrounded at base by an irregular
its
large nuclei. The proximal end of the sac (S, Fig. 40) extending from the uterus.
oviduct (OD) is folded back on itself and We have not encountered any reference
lies lateral to the carrefour to which it to this structure in the literature but an
is connected by means of a short wide apparently similar, though less well de-
duct into which both the hermaphrodite veloped sac, appears to be figured for
duct and the sperm duct (SP.D.) open. B. contortus by Larambergue (1939) who,
Wright (1957) also mentions a carrefour however, does not refer to it in the text.
in B. (P.) jousseaumei which, although In B. tropicus the penis of the partner
not figured, seems to agree with that acting as the male may occasionally be
described by Abdel-Malek (1952) in being found inserted into this sac. A similar
a dilatation of the oviduct a short dis- interpretation is suggested by Laram-
tance from its point of origin. Even bergue' s (1939) Fig. 44. In one instance
though this structure does not seem to we found the penis to have been inserted
us to be homologous with that occurring into the spermatheca (Fig. 39).
in B. tropicus, we follow the example of Dorsally and running obliquely across
Larambergue (1939), who applied the term the posterior surface of the uterus, is
carrefour to a structure occurring in B. the large well developed muciparous
contortus, which is similar to that of B. gland (M.GL., Figs. 40, 48). This struc-
tropicus ture consists of tall, large glandular
The albumen gland (AL.GL.) is more cells arranged to form simple acini, the
or less kidney- shaped and is slightly con- openings of which can be seen in regular
cave on the ventral and convex on the longitudinal rows on the inside of the
dorsal surface. The posterior edge of uterus. On the ventral side of the uter-
the uterus fits into the hilus of the gland us the gland is restricted to the right
while the intestine is lodged in a groove hand posterior region. The cells secrete
(OR) along its posterior edge. The al- mucus and remain unstained except for
bumen gland consists of small lobules of their nuclei. The rest of the dorsal wall
large, closely packed cells filled with of the uterus is folded longitudinally, the
deeply staining secretory droplets, simi- region along the border of the muciparous
lar to those found by Wright (1957) in gland being a darker yellow than the rest
the same organ in B. (P.) jousseaumei. of the uterus. De Azevedo et aL (1957)
The duct of this gland is lined by ciliated designate this darker portion as the nida-
cuboidal cells containing large nuclei. mental gland (N.GL.) while Abdel-Malek
(1952) and Wright (1957) refer to the
The female system
corresponding structures as the oöthecal
The oviduct (OD, Figs. 38, 40, 41) is gland. The longitudinal folds (F) are
convoluted and proximally it is lined by covered with tall, glandular cells with
a very glandular, columnar, ciliated epi- almost basally placed nuclei. There are
thelium. Its walls are thickest towards large, round, lighter staining areas pre-
the uterus. De Azevedo et al. (1957) sent probably representing secretory
consider the dorsal glandular region of cells. The epithelium of this region is
The penlal complex; the usually subterminal junction of the vas deferens is obscured by
FIG. 42.
the penis sheath.
FIG. 43. Representative transverse section of the sperm duct.
FIG. 44. Representative transverse section of the vas deferens.
FIG. 45. Cross-section of the proximal region of the epiphallus.

papilla,
FIG, 46, Transverse section of part of penlal wall, through a
wall of the preputium sectioned to show the tissue lining the lumen.
FIG. 47. Portion of the
show the structure of the muciparous
FIG, 48. Semidiagrammatic cross- section of the uterus to
gland,
FIG. 49. Cross-section of the distal region of the epiphallus.

not richly ciliated. The rest
of the uterus region, representing the apices of the
is also folded but a lesser degree
to acini, and a central whitish and more
and the ventral surface may be almost translucent one, representing the bases
smooth. The cells in the latter region of the acini. The apical cells are full
are very tall, bear numerous cilia and of secretory droplets which readily take
are interspersed with goblet cells. up acid fuchsin when stained with Mal-
The spermatheca (STH, Fig. 41) is lory's triple stain. The translucent re-
round; its size depends on the amount gion stains very light blue and the cyto-
of sperm it contains and the age of the plasm appears to be less granular than in
snail. The spermathecal duct is short the apical cells. They contain large clear
The walls of the spermatheca are thin areas which probably represent masses
and covered with a few strands of con- of secretion. A narrow blue staining
nective tissue. On the inside they are zone intervenes between the peripheral
lined by an epithelium of tall ciliated and central regions.
cells with centrally to basally placed The vas deferens (Fig. 44) is lined by
nuclei. Both the spermathecal duct and a cuboidal ciliated epithelium with large
the vagina are lined by cells of this type nuclei; this lining is surrounded by a
but, although still ciliated, the cells of thick layer of circularly arranged muscle
the latter become more cuboidal toward fibres interspersed with a few connective
the female aperture. tissue fibres.
The penis sheath (P.S., Fig. 42) is
The Male System
club-shaped, the proximal end being
The male reproductive system (Figs. broader. The surface is usually fairly
41, 42) is of more use, taxonomically, smooth although it may be slightly con-
as it is not subject to such great sea- stricted or even markedly bulging as
sonal changes as those occurring in the figured by Hubendick (1948). It is thin
female system. The sperm duct (Fig. proximally where the lumen is wide and
41) arises as a fairly narrow tube and conversely it is thick distally where the
is of the same diameter throughout. It lumen is narrow. The walls of the proxi-
is lined by a cuboidal ciliated epithelium mal portion consist of a thin muscular
with large nuclei (Fig. 43) and is en- layer containing scattered pigment cells
sheathed by a thin layer of lightly pig- and a few strands of connective tissue
mented connective tissue. and are not always of the same thickness
The prostate gland (PR.GL., Fig. 41) is throughout. At the distal end of the sheath
flat ventrally and convex dorsally where where it joins the preputium (PP) the
it abuts against the uterus. The gland outer muscular layers are replaced by
consists of numerous fingershaped acini connective tissue and isolated muscular
radiating from a more or less central fibres. The preputium is of about the
point on its ventral surface. At this same diameter throughout and is straight
point the sperm duct enters and the vas whereas the penis sheath is always bent
deferens (V.D.) leaves the gland, thus or convoluted. Numerous small muscles
agreeing with the descriptions furnished (EX.M.) are attached along the anterior
by Larambergue (1939) and Wright (1957) and posterior margins of the preputium.
for related bulinid snails. Each acinus According to de Azevedo et al. (1957)
is deep and surrounded by a thin connec- these are extensor muscles. A single
tive tissue sheath containing pigment cells large retractor muscle (RE.M.) is in-
as described by Wright The simple serted at the junction of penis sheath
epithelial lining consists of large secre- and preputium. It has its origin in the
tory cells having large basal nuclei and columellar muscle as far back as the
conspicuous nucleoli. In gross dissection base of the uterus. Along the lines of
two zones may be distinguished in the attachment of the extensor muscles the
gland, viz. a peripheral opaque yellowish preputial wall is swollen out into two
longitudinal pilasters similar to those Since the length of the penis sheath
occurring in other species of the Planor- relative to that of the preputium has been
bidae, described by Baker (1945),Huben- employed to characterize certain species
dick (1946, 1955), Mandahl-Barth (1954, of the genus Bulinus (Mandahl-Barth 1954,
1956), de Azevedo et al. (1957), Wright 1956), we have calculated this ratio for
(1957), and Schutte and van Eeden (1959b). a number of specimens of B. tropicus
Contrary to Hubendick's (1948) statement (Table EX and Fig. 50). From the table
that there are no muscular pillars proxi-
mally inß. tropicus and Wright's (1957) TABLE EK. Ratios of the penis sheath to
the preputium (PS/PP) in each of the sam-
statement that only one of the pilasters
ples and for snails of different sizes.
extends as far as the junction of penis
sheath and preputium in ß. (P.) jous-
Sample
seaumei, they do in the specimens of
B. tropicus examined, both extend into
that region and determine the shape of
the lumenof the preputium.^
In B. tropicus the pilasters are situ-
ated opposite each other and are almost
contiguous so that in transverse section
the lumen isH-shaped. In certain cases,
however, probably through unequal con-
traction of the muscles, the pilasters are
not directly opposite each other and this
results in an S- shaped lumen.
The epithelium lining the lumen of the
preputium, which Hubendick (1948) de-
scribes as being cuboidal in B. tropicus
may be regarded as either cuboidal or
columnar. This was also found to be the
case in B. (P.) jousseaumei (Wright,
1957). Whereas the latter author found
only a few ciliated cells in the preputial
lining of B. (P.) jousseaumei, and Huben-
dick (1947) states that the preputium is
nonciliated in the Planorbidae, the proxi-
mal region, in B. tropicus, was found to
be richly ciliated (Fig. 47). The large
goblet cells of B. (P.) jousseaumei are
described as grouped together and pene-
trating into the muscular layer, but B.
tropicus differs in both these respects
(G.C., Fig. 47). The differently staining
cells by Wright (1957) are
mentioned
visible. The epithelium is underlain by
a distinct basement membrane and the
rest of the preputial wall consists of a
loose meshwork of pigmented connective
and muscular tissue interspersed with
blood spaces.
ment of the muscular tissue is weaker thus forming protuberances projecting
than in the latter. Distally the epiphallus into the lumen. Following Larambergue
loses its circular appearance owing to (1939), we refer to these protuberances
the accumulation of muscular and connec- as papillae. In surface view the arrange-
tive tissue on either side, which produces ment of these papillae resembles that of
a more or less oval shape. Scattered the protuberances of a pineapple. Proxi-
cells full of globules (GLO, Fig. 49) occur mally they are large while the muscular
in this connective tissue. From their part of the walls of the penis is not thick.
appearance they seem to be glandular Distally the papillae decrease in size and
cells and, as these globules do not stain the cells become more squat while the
at all, whereas the other secretions ge- muscular layer increases in thickness.
nerally take up stains readily, they are The large vacuolated cells described by
probably mucous cells. The cells lining Larambergue (1959) occur in the distal
the lumen of the epiphallus in this region region. At its extreme tip the penis it-
are taller than elsewhere and the nuclei self becomes thin-walled and the epithe-
are basally placed. The epiphallus usually lial cells cuboidal. In most cases a short
joins the penis subterminally. Neither ridge occurs on the inside of the penis in
this duct nor the penis proper contains line with the opening of the epiphallus.
any pigment cells. This ridge coincides with a groove on
The morphological and histological de- the external surface. In those cases in
tails of the walls of the penis are depicted which the ridge is absent its position is
in Fig. 46. From this illustration, it is generally betrayed by a slight concen-
apparent that the wall of the penis con- tration of longitudinal muscle fibres. To
sists of two layers, viz. an inner epi- our knowledge no ridge of this kind has
thelial layer and an outer layer consisting as yet been described for any of the
of connective and muscular tissue. The bulinids. In a few instances the penis
latter forms a series of projections (PAP) and sheath could not be distinguished
directed toward the lumen and clothed and the lumen of the sheath was filled
with greatly elongated epithelial cells, with loose, strongly pigmented connective
3-0
FIG. 50. Histogram of the length ratioof the penis sheath to the preputium (PS/PP); 155 specimens
measured.
FIG. 51. Histogram of the length ratio of the penis to the epiphallus (P/EP); 147 specimens meas-
ured.
tissue. As in the case of the ratio penis

sheath to preputium, (PS/PP, Table IX
and Fig. 50) the ratio penis to epiphallus
(P/EP, Table X) is very variable. In
TABLE X. Ratios of the penis to epiphallus

(P/EP) for each of the samples and for
snails of different sizes.
Sample
CAWSTON, F., 1922, Experimental infestation of 1955, Phylogeny in the Planorbidae.

,
freshwater snails with special reference to Trans. zooL Soc. London, 28: 453-542.
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, 1923, Occasional hosts of some South lusken. Ebner und Seubert, Stuttgart. 1-140.
African trematodes. Rep. S, Afr. Ass. Adv. *LACAZE-DUTHIERS, H. de, 1872, Du système
Sei., p. 351-353. nerveux des mollusques gastéropodes pul-
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hosts and a study of their radulae. J. trop. 437-500.
Med. (Hyg.), 28: 14-15. LARAMBERGUE, M. de, 1939, Etude de l'auto-
1925b, A contribution to the study of
,
fécondation chez les gastéropodes pulmones.
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Med. (Hyg.), 28: 365-366. chez Bulinus (Isidora) contortus Michaud.
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ZUSAMMENFASSUNG
BEITRAG ZUR MORPHOLOGIE VON BULINUS TROPICUS
Gegenstand der vorliegenden Studie ist Bulinus (Bidinus) tropicus (Krauss),

eine mit den Zwischenwirten der Schistosome der haematobium- Gruppe eng ver-
wandte afrikanische Süsswasserschnecke, die auch als Überträger der Trematoden
Paramphistomum und Cotylophoron von veterinärer Bedeutung ist. Drei taxonomisch
kennzeichnenden Partien, dem Gehäuse, der Radula und dem Penialkomplex, wurde
besondere Aufmerksamkeit geschenkt; andere Organe wurden jedoch gleichfalls
morphologisch und histologisch untersucht.
Das zur Untersuchung dienende Schneckenmateral wurde sowohl auf Ähnlichkeit
mit typischen B. tropicus Exemplaren hin gesammelt, wie auch in Hinblick auf seine
Herkunft aus der Gegend des locus typicus. Trotz dieser sorgfältigen Auswahl wurde
aber hier wieder einmal der notorische Polimorphismus dieser Art bestätig. Die
wichtige Rolle der Umweltseinflüsse in der Bildung von Schalentypen welche sich
sowohl ihrer Gestalt nach, wie auch ihrer Beschaffenheit nach, von einander unter-
scheiden, wird erörtert. Obwohl sich unter unserem Material bei ausgewachsenen
Formen 2 Haupttypen vorfanden, ein langgestreckter, eng genabelter mit fliessendem
Wasser verbundener und ein gedrungener, weit genabelter, mit stehendem Wasser
verbundener (Fig. 3), gehörten die jugendlichen Formen einem einzigen Typus an,
ein Hinweis darauf, dass, wie es auch bereits Schutte und van Eeden (1959a) für
Biomphalaria pfeifferi hervorgehoben haben, die Gestalt der jugendlichen Exemplare
weniger von der Umgebung abhängig ist als die der Erwachsenen. Dieses scheint
insofern folgerichtig als ja die letzteren den modifizierenden Einflüssen der Umwelt
länger ausgesetzt waren als die Jungschnecken. Konchologische Messungen wurden
an insgesamt 171 Schnecken aus 9 Fundorten vorgenommen, wobie die Länge
imd Breite des Gehäuses, Länge und Breite der Schalenmündung, und Länge
des Gewindes gemessen, sowie die verschiedenen Grössenverhältnisse dieser
Ausmasse zueinander berechnet wurden, um die Natur der Beziehungen zwischen
immer je zwei derselben zu erforschen. Die Gewinde- und Lippenabfallswinkel
wurden ebenfalls gemessen. Nur ein Vergleich mit bei anderen Arten erhaltenen
108 STIGLINGH, VAN EEDEN AND RYKE I
Werten dieser selben Messuiên wird zeigen, welche dieser Beziehungen sich als
genügend ausgeprägt erweisen werden um taxonomisch verwertbar zu sein. Obwohl
die einzelnen Messungen manchmal stark variieren, so sind doch die mittleren Werte
für jede der 9 Proben im allgemeinen ziemlich konstant. Eine Ausnahme bildet nur
das Längenverhältnis Gehäuse/ Gewinde, in welchem die mittleren Werte zwischen
6,7 und 15.2 schwanken. Dennoch war das Gewinde in kleineren Formen immer
verhältnismässig kürzer als in den grösseren. Der apikale Winkel des Gewindes
zeigte sich ausserordentlich veränderlich: 58 ° - 134° . Der Mttelwert der Mün-
°
dungslippenabfallswinkel hlnggen liegt für alle Posten bei 57° order 58 , ausser in
einem einzigen, wo er 61° betragt. In Bezug auf die Mündungsform zerfallen die
ausgwachsenen Gehäuse gewöhnlich ebenfalls in 2 Hauptgruppen: deren Gestalt
scheint nlch nur vom Alter abhängig zur sein sondern auch von verschiedenartigen
ökologischen Bedingungen. Obwohl die Gestaltung der Kolumellapartie innerhalb
der Fimdortsproben ziemlich einheitlich war, so erscheint sie doch von Gruppe zu
Gruppe etwas verändert.
Ein Osphradium wurde ausserhalb der Mantelhöhle auf dem Mantelwulst ge-
funden. Die Kieme (Pseudobranchie) erwies wich als normal. Die Analregion
hingegen war etwas anders als im Schrifttum beschrieben, nämlich mit schmälerem
Anallappen und ausgesprochenerer Rektalleiste. Die Mittelwerte der Längenver-
hältnisse Niere/ Ureter schwankten zwischen 2.0 und 4.6.
An der Decke der Mantelhöhle befinden sich nur 2 der in den Planorbiden
taxonomisch bedeutsamen Falten: es fehlen die Nierenleiste und die mittlere Rek-
talleiste. Die zwischen Niere und Rektum beûndliche intermediäre Mantelleiste
unterscheidet sich von der von Mandahl- Barth (1956) veröffentlichten Abbildung
durch ihre grössere Länge und dadurch, dass sie immer nach rückwärts bis zum
Rand der Mantelhöle reicht, wo sie der seitlichen Rektalleiste begegnet.
Eine Beschreibung der die Pseudobranchie entleerenden Blutgefässe ist gegeben.
Die sphärischen, von Schutte und van Eeden (1959b) im Pericardraum der afrikani-
schen planorbiden Schnecke Bio»iphaIaria pfeifferi angetroffenen Körper wurden in
einer grösseren Anzahl von Bulinus tropicus ebenfalls aufgefunden. Es handelt sich
möglicherweise um die Eier eines nicht identifizierten Wesens.
Im Verdauungstrakt entsprach der Idefer, im Gegensatz zu gewissen im Schrift-
tum gemachten Angaben, durchaus dem normalen Schema für Bulinus. Unter-
suchungen über die allgemeinen Proportionen der Radula zeigten, dass das Verhält-
nis der Länge zur Breite ziemlich konstant ist (2.4 - 2.6) und dass sich somit die
mittleren Verhältniszahlen möglicherweise zu Vergleichen mit verwandten Arten
eignen könnten. Die Seitenränder der Zahnplatten waren gewellt. Kleine Zäckchen
wurden zwischen den Zacken des Mittelzahnes und auch an der Basis des Mesokonus
der Seitenzähne gefunden. Dieser letztere ist beinahe nie einfach dreieckig, wie
von Mandahl- Barth (1956) als kennzeichnend fur die B. tropicus Gruppe angegeben;
manchmal nähert er sich sogar der Pfeilspitzenform die ihm zunach fur die B, trun-
catus Gruppe charakteristisch ist. Trotz der Begrenzungen die anerkannterweise den
Radulaformeln inneliegen, word hier eine allgemeine Formel für B. tropicus, wie
folgt, angegeben: 1: 6+2 24 x 123.
:
Die verschiedenen Bezeichnungen der Ganglien des Nervenschlundrings werden

erörtert und die daraus entspringendem Nerven beschrieben. Die Speicheldrüse
geht auch hier durch den Nervenring hindurch.
Die Zwitterdrüse ist nicht in männliche und weibliche Zonen gegliedert. Wie
auch bei gewissen anderen Lungenschnecken weisen die Oozyten doppelte Zellkerne
auf. Die "carrefour" Gegend unterscheidet sich von der bei der nordamerikanischen
planorbiden Schnecke Helisonm trivolvis beschriebenen, stimmt jedoch mit der
bei BiotHplialarin pfeifferi gefundenen überein. Die Anwesenheit eines unregel-
mässign, vom Uterus ausgehenden Sackes, welcher teilweise die Basis des Eileiters
umfasst, wird hervorgehoben. In der Prostata lässt sich eine opake, perlpherale,
gelbliche Zone von einer durchscheinenderen, weisslichen, zentralen, unterscheiden.
Im proximalen Teil des Penis wurde die Anwesenheit einer kurzen inneren Leiste
bemerkt. Die Pfeiler im Präputium erstrecken sich bis zu seiner Vereinigung mit
der Penisscheide, Das Längenverhältnis Penlsscheide/Präputium schwankt zwar im
einzelnen sehr, aber die Mittelwerte für die verschiedenen Gruppen variieren nur
zwischen 1,2 und 1,6,
RESUME
CONTRIBUTIONS A LA MORPHOLOGIE DE BULINUS TROPICUS
L'étude porte sur Bidinus (Bulinus) tropicus (Krauss) un mollusque fluviatile

africain prochement apparenté aux hôtes intermédiaires des schistosomes du
groupe haeniatobiii))!, qui a aussi une certaine importance vétérinaire comme vecteur
des trematodes Para»iphisto»m}}i etCotylophoron. Notre attention a été spécialement
dirigée sur 3 charactères d'importance taxonomique: la coquille, la radule et le
complexe pénial; cependant, les autres organes furent également examinés, mor-
phologiquement ainsi qu'histologiquement.
Dans la sélection du matériel pour cette étude la ressemblance de ces mollus-
ques aux spécimens typiques de B. tropicus ainsi que leur provenance de la region
approximative du lieu typique furent prises en considération. Malgré ce choix, la
présente étude de ces coquilles a démontré une fois de plus le polimorphisme notoire
de l'espèce. Le rôle important des facteurs écologiques dans la formation de types
conchologiques assez divergeants, différant en forme ainsi qu'en sculpture, est
discuté. Quoique 2 types principaux peuvent être distingués dans notre matériel,
l'un élongé, à ombilic étroit, associé à l'eau courante et l'autre trapu, à ombilic plus
large, associé aux mares (Fig. 3), il n'y avait qu'un seul type juvénile, fait qui
suggère, comme l'ont déjà observé Schutte et van Eeden (1959a) pour Biomphalaria
pfeifferi, que les formes jeunes sont moins affectées par l'ambiance que ne le sont
les adultes, ce qui est d'autant plus logique que les derniers sont exposés aux in-
fluences modifiantes plus longtemps que les premiers. Une analyse conchologique
fut faite sur 171 exemplaires provenant de 9 localités différentes. Furent mesurés:
longueur et largeur de la coquille et de 1' aperture, ainsi que hauteur des tours
de spire. Les divers rapports, l'un à l'autre, de ces mesures furent calculés afin
de découvrir la nature de leurs relations mutuelles. Les angles de la spire et de
la dépression lábrale furent également mesurés. Ce n'est pourtant que la com-
paraison de ces rapports avec les valeurs correspondantes dans les espèces
proches qui montrera lesquels sont suffisamment distincts pour servir taxonomi-
quement. Quoique les écarts individuels de ces quotients sont parfois consi-
dérables, les moyennes obtenues pour les 9 lots sont d'habitude assez constantes,
exception faite du rapport entre les longueurs des coquilles et des spires, où ces
moyennes varient entre 6.7 et 15.2. Néanmoins les spires étaient toujours relative-
ment plus courtes dans les petits specimens que dans les grands. L'angle apical de
la spire est très variable: 58° - 134°. L'angle moyen de dépression de la lèvre
aperturale mesurait 57*^ et 58° pour toutes les localities sauf une, où il était 61° .
Quant à la forme de aperture, les coquilles adultes appartenaient d'habitude à 2
groupes; ce caractère semble changer non seulement avec l'âge mais aussi avec
les diverses conditions écologiques. La région columellaire était assez uniforme
dans chacun des lots, mais varie d'un groupe à l'autre.
L'osphrade fut trouvé à l'extérieur de la cavité palléale sur le collier du man-
teau. La pseudobranchie est normale. La région anale diffère légèrement de cer-
taines descriptions antérieures en ayant un lobe anal plus étroit et une arête rectale
plus distincte. Les valeurs moyennes du rapport rein/urètre varient entre 2.0 et
4.6 dans les lots différents.
Seulement 2 des plis taxonomiquement importants dans les planorbes se
trouvent sur la paroi supérieure de la cavité palléale tandis que le pli rénal et le
pli rectal central manquent. Le pli palléal intermédiaire, situé sur le manteau
même entre le rein et le rectum, se distingue de l'illustration donnée par Mandahl-
Barth (1956) par sa plus grande longueur et du fait qu'il s'étend toujours postérieure-
ment jusquau bord du manteu où il rejoint le pli rectal.
Les vaisseaux sanguins évacuant la pseudobranchie sont décrits. Les objects
sphériques, déjà observés par Schutte et van Eeden (1959b) dans la cavité péri-
cardiale de Biomphalaria pfeifferi, furent également trouvés dans un grand nombre
d'individus de B. tropicus. Il est possible qu'ils soient les oeufs d'un organisme
inconnu.
Dans le système digestif, la mâchoire fut trouvée être différente des descrip-
tions présentées dans la literature et conforme au dessin général pour les bulins.
Une étude des proportions générales de la radule révèle que la moyenne du rapport
lio STIGLINGH, VAN EEDEN AND RYKE
longueur sur largeur est assez constante (2.4 à 4.6) et que ce rapport pourrait ainsi
se montrer utile à comparer les espèces voisines. Les bords des dents sont ondulés.
De petits denticules furent découverts entre les cúspides des dents centrales et à
la base du mésocone des dents latérales. Les mésocones ne sont presque jamais
simplement triangulaires, condition typique pour le groupe de B. tropicus selon
Mandahl- Barth (1956), et s'approchent même en certain cas, de la forme en pointe
de flèche caractéristique, selon cet auteur, pour le groupe de B. triincatus. Malgré
les limitations reconnues de toute formule radulaire, une formule généralisée est
présentée pour la radule de B. tropicus: 1 6 + 2 24 x 123.
: :
Les noms donnés aux divers ganglions de l'anneau circumésophagéal sont dis-
cutés et les nerfs y prenant origine sont décrits. Les glandes salivaires passent à
travers de cet anneau.
Dans la glande hermaphrodite il n'y a pas de division entre zones maie et
femelle. Les oocytes, comme dans certains autres pulmones, ont des nucléoles
doubles. La région du "carrefour" diffère de celle décrite pour le planorbe Helisoma
trivolvis de l'amérique du nord et conforme avec les conditions trouvées dans le
planorbe africain Bioniphalaria pfeifferi. La présence d'un sac irregulier provenant
de l'utérus et entourant partiellement la base de oviduct est relevée. Dans la
glande prostate deux régions principales peuvent être distinguées, l'une péri-
phérique opaque et jaunâtre, l'autre centrale, plus pellucide et blanchâtre. La
présence d'une arête intérieure et courte dans la région proximale du pêne est
signalée. Les pilastres du prépuce s'étendent jusqu'à la jonction de ce dernier avec
la gaine péniale. Le rapport gaine péniale/prépuce varie considérablement, mais
les moyennes pour les divers groupes ne varient qu'entre 1.2 et 1.6.
CONTRIBUCIÓN AL ESTUDIO DE BULINUS TROPICUS

RESEÑA
Estudio realizado sobre el Bulinus tropicus (Krauss), caracol dulceacuícola
africano que está estrechamente relacionado al huésped intermediario de los
esquistosomas del grupo hematobium, y que tiene también importancia veterinaria,
como trasmisor de los trematodes Paramphistomun y Cotylophoron. Se prestó
especial atención a 3 caracteres taxonómicos: la conchilla, radula y complejo
penial. Los otros sistemas fueron también examinados morfológica e histológi-
camente.
Las muestras del caracol fueron seleccionadas en base a su parecido a los
especímenes típicos de B. tropicus y a su distribución geográfica, en la región de
la localidad típica. A pesar de la prolijidad en la selección, el estudio concho-
lógico destaca una vez más el notable polimorfismo de la especie. Se discute la
importancia de los factores ecológicos en el desarrollo ampliamente divergente de
los tipos de concha en forma y textura. Aún cuando se encuentran 2 formas adultas,
una prolongada, estrechamente umbilicada y asociada con cursos de agua, y otra de
forma ampliamente umbilicada asociada con lagunas y aguas estancadas (fig. 3),
existe solamente un tipo principal juvenil sugiriendo, tal como fuera observado por
Schutte y Van Eeden (1959a) para la Biomphalaria pfeifferi que el tipo juvenil es
menos afectado por el ambiente que el adulto, lo que parecería lógico, ya que el
último mencionado ha estado expuesto a las únfluencias modificantes por un período
más largo que los jóvenes. Un análisis conchométrico se llevó a cabo en el largo,
ancho, y alto - ancho de la abertura, y largo de la espira, en 171 caracoles de 9
localidades. Se calcularon diferentes proporciones de modo de descubrir la
naturaleza de la relación en cualquiera de estas características. Los ángulos
espirales y depresión del labio también fueron medidos. La comparación con
valores correspondientes de otras especies podrán indicar cuales, de todas las
proporciones obtenidas, varían suficientemente para usarlas taxonómicamente.
Los límites de estos valores son grandes en algunos casos y la media de las 9
muestras es bastante constante. Una excepción es la comparación entre la longitud
total de la concha y la de la espiral, en la cual los valores de la media varían entre
MORPHOLOGY OF BULESfUS TROPICUS HI
6,7 y 15,2. Sin embargo se descubrió que la media espiral es relativamente más
corta en los ejemplares pequeños. El ángulo apical es muy variable: 58-134? La
media del ángulo de la depesiön del labio es 57-58°para todos los ejemplares, con
excepción de uno solo, 61" Con respecto a la configuración de la abertura, los
adultos se clasifican por lo general en dos grupos principales. Tal característica
parece variar no solamente con la edad del ejemplar, sino también con las con-
diciones ecológicas. Aunque la región de la columnilla es bastante uniforme en
cada muestra, varía de uno a otro ejemplar.
El osfradio se encuentra fuera de la cavidad del manto, en el cuello. La
branquia (pseudobranquia) es normal. La región anal se diferencia ligeramente de
las descripciones previas en que el lóbulo anal parece ser más estrecho y la
arruga renal más clara. La longitud reñón-uretra varía entre valores de la media
de 2,0 y 4,6 en los distintos ejemplares.
Sólo dos de las arrugas taxonómicamente importantes se presentan en la parte
superior de la cavidad paleal. Las arrugas renal y media rectal no se encuentran.
La arruga intermedia de la cavidad, entre el riñon y el recto se diferencia de la
figura de Mandahl- Barth (1956) en que es más larga y siempre se extiende poste-
riormente hasta encontrar la arruga lateral rectal al borde de la cavidad paleal.
Se mencionan las venas en que desagotan las branquias (pseudobranquias). Los
cuerpos esféricos mecionados por Schutte y Van Eeden (1959b) para la cavidad
pericardial de B. pfeifferi, se encontraron en gran número de especímenes; se
cree que sean los huevos de organismos no identificados.
En el sistema digestivo, la mandíbula parece diferir de las descripciones
ofrecidas en la literatura y concuerda a la organización general de Bulinus. El
estudio general de las proporciones radulares demuestra que el porcentaje de la
media largo/ancho es muy constante (2,4 - 2,6), de manera que este porcentaje
puede ser muy útil en la comparación con las especies mencionadas. Los lados de
los dientes son acanalados y aparecen pequeños dentículos entre los centrales y
en la base del mesocono de los laterales; este último casi nunca es simplemente
triangular como estableció Mandahl-Barth (1956) para el grupo de B. tropicus. En
ciertos casos se aproxima a la característica forma en punta de flecha del grupo
de B. truncatus. A pesar de las limitaciones que se conoce en la fórmula radular,
una fórmula generalizada puede ofrecerse para B. tropicus: 1:6+2:24x123,
Los nombres aplicados a los varios ganglios del anillo circumesofageo son
discutidos, asi como también la descripción de los nervios que de él se originan.
Las glándulas salivares pasan a través del nervio del anillo.
En la glándula hermafrodita no hay división de zona masculina y femenina.
Los oocitos tienen doble nucelolo como en el caso de ciertos otros pulmonados. La
región "carrefour" difiere de la definida para el planorbido norteamericano
Helisoma trivolvis y concuerada con la condición encontrada en el africano B.
pfeifferi. Llama la atención un saco irregular que se extiende desde el útero y que
rodea parcialmente la base del oviducto. En la próstata se distinguen dos regiones
principales, una periferial opaca de color amerillento, y otra mas traslucida, blan-
quecina. Se nota la presencia de una arruga interna corta, en la proximidad del
pene. Las pilastras del prepucio se extienden hasta la unión del pene con la capa
que envuelve as mismo. El porcentaje de la capa que envuelve al pene-prepucio varia
considerablemente, pero las medidas de los ejemplares analizados separadamente
varían sólo entre 1,2 y 1,6.
, BULINUS (BULINUS) TROPICUS (KRAUSS)

(MOLLUSCA BASOMMATOPHORA PLANORBffiAE)
. . . . .
, .,
,« ,¿
Bulinus (.) tropicus IKrauss)
Planorbidae, -
, -
íosowa wa
--,
S
, , -
- tomum .
; , , -
.
,
.
.
-
tropicus
, . -, ,, , -
,
-
-
ö 2
,
- , ,
, . ,.., ,-.
--
.
-
Biomphalaria pfeifferi
,
-
(. 3),
(1959)
. ,.,, ., , ,
. -
,., , --
-- .
2,0
-
-
hth--
, ,
, -.
(19Ç6);
. , ., --
, Biomphalaria pfeifferi
(199)
. . ,. -
Bulinus tropf cus.
Bulinus
- ,
(2,Í4. - 2,6)
(195^) tropicus Bul i-

,
nus
, .-
MORPHOLOGY OF BULINUS TROPICUS
truncatus .
-
113
. . tropicus 1:6 + 2 ; 2Í4. 123.
.,, ., -
,-
--
Pulmón at
: ... .--
Bulinus (Phy sopsis) jous seaumei
Helisoma trivolvis
-- Biomphalaria pfeifferi.
-
,,-1,2 - 1,6,
LIST OF ABBREVIATIONS
A - auricle M.C.F. - mantle cavity floor
AC - acinus M.C.L. - muscular and connective tissue layer
AL.GL. - albumen gland M.C.R. - mantle cavity roof
AL.GL.D. - albumen gland duct ME - mesocone
A.L. - anal lobe M.N. - nerves supplying muscles
- aorta M.GL. - muciparous gland
- base M.S. - muscle strands
B.C. basal or Sertoli cells
- MU. L. - muscular layer
B.C. buccal ganglion
- N - nucleus
BL - blood N.C. - nurse cell

B.M. - basement membrane N.GL. - nidamental gland
B.RET. - buccal retractor N1 - nucleoli
B.R.N. - buccal retractor nerve - osphradium
buccal mass - oocytes
BU.M. -
B.W. - body wall OD - oviduct
B.W.N. - body wall nerve OE - oesophagus
- central cavity O.N. - oesophageal nerve
C.A. - common atrivim OS.N. - osphradial nerve
CAE - caecum OV - egg
CAP - capsule PA.G. - parietal ganglion
CAR - carrefour PAP - papilla
... - cerebro-buccal connective P.C. - peripheral cavity

CG. - cerebral ganglion P. COL. - posterior part of collar
CIL. . - ciliated cells round aperture of PE.G. - pedal ganglion
osphradium PE.N. - pedal nerve
C.N'. - CQllar nerve from right parietal PER - pericardium
ganglion P.F. - primary fold
C.N". - collar nerve from visceral ganglion PLC. - pigment cell
COL - collar PL.G. - pleural ganglion
CR - crown PN pneumostome
-
CRY crystals
- PN.N - pneumostome nerve
C.T. connective tissue
- PP - preputium
D - denticle PR.GL. - prostate gland
D.T.P. - dorsal tentacular process P.S. - penis sheath
E - epithelium P.V.- pulmonary vein
EC ectocone
- R rectum
-
EG - eggs RE.M. - retractor muscle

EM.C. - large embryonic cells R.SH. - radula sheath
EN - endocone R.SH.N. - radula sheath nerve
EP - epi phallus R.V. - renal vein
EX.M. extensor muscles
- S - sac
EYE eye
- SAL. GL. - salivary gland
F - fluted edge S.B. - sperm bundle
FO - fold S.F. - secondary folds
FOO - foot SG - spermatogonia
G - gill S.K. - saccular part of kidney
GAN - ganglion S.N. - siphon nerve
G.C. - goblet cells SP - sperm
GLC. - giant cells ST - statocyst
GLO - globules STD - spermatids
G.N. - nervegill STOM -stomach
GR - groove accommodating intestine STH - spermatheca
H.D. - hermaphrodite duct T - tentacle
H.GL. - hermaphrodite gland .. - tubular part of kidney
LC.L. - inferior cerebral lobe T.N. - tentacular nerve
I.M.R. - intermediate mantle ridge - ureter
INT - intestine U.C. - urine concretion
J - jaw U.N. - uterine nerve
- kidney V - ventricle
LAT. N. - lateral nerves VA - vagina
L.J. - lateral jaws V.C. - vacuolated cells
L.N. - labial nerve V.D. - vas deferens
L.R.R. - lateral rectal ridge V.G. - visceral ganglion
L.UT. - lumen of uterus V.S. - vesiculae seminales
MA - mantle V.T. P. - ventral tentacular process
11, 1962, 1(1): 115-137
PUNCTATION OF THE EMBRYONIC SHELL OF

BULININAE (PLANORBroAE) AND SOME OTHER BASOMMATOPHORA
AND ITS POSSIBLE TAXONOMIC-PHYLOGENETIC IMPLICATIONS!
H. J. Walter
Liberian Institute of the American Foundation for
Tropical Medicine, Inc. Harbel, Liberias
,
ABSTRACT
Microscopic punctation of the embryonic shell is here reported for the first
time to be a character universal in Bulinus s.l. over its geographic range, and to
exist in the only other recognized genus of Bulininae, Indoplanorbis.
The author studied over 1,500 Bulinus s.l. shells of at least 14 species, "sub-
species," or varieties of the subgenera Bulinus s.s, Pyrgophysa and Physopsis;
,
these were represented by 106 different field and laboratory populations, particu-
larly from Liberia, but also from Sardinia and Iraq and from numerous other lo-
calities ranging through the length of Africa. Punctation was observed in every
specimen studied except in relatively few cases when damage to the shell or other
factors interfered. The punctae are well-defined pits consistently occurring in a
basic pattern that dominates the embryonic microsculpture and differs somewhat
for the 3 subgenera. Study of large numbers of shells of embryonic laboratory
reared snails provided basic information about the nature and arrangement of the
punctae and of other microsculptural elements and showed the constancy of punc-
tation in each of the 3 subgenerlc groups oi Bulinus. Apical punctation of the shell
has been reported by others for each of these subgenera, but only for a very few
species and pertinent previous literature suggests that even in these, some indi-
viduals or populations fall to develop the character. The commonly Inexact usage
of terms such as "dots," "nodules," "punctures" or "Impressions," along with
meagre data in past conchologlcal descriptions, show that in respect to micro-
sculpture, the species of Bulininae are yet poorly known, the same applying for the
Basommatophora In general. Punctation must have been overlooked In the past be-
cause it Is identifiable only In relatively undamaged, very clean shells, with ap-
propriate magnifications and critical illumination. Success In demonstrating puncta-
tion depended greatly on the author's technique of cleaning shells with sodium hypo-
chlorite, which permitted study of sculptural detail by transmitted light.
Among non-bulinlne Planorbidae embryonic punctation had been previously
known to occur in the "Se^menizwö -group" and in Platytaphius ('i=Taphius) and, in
other Basommatophora, in the "ancylid" Bumupia of Africa, and in the ellobilds
Melampus Phytia, and Pythia. Here, from the examination of a few species of non-
,
bulinlne planorblds, the author reports the existence of nuclear punctation in

Planorbarius corneus and its total lack in Planorbina (Australorbis and Biomr
phalario) and in a species of Gyraulus. The same absence of punctation was found
In various Lymnaeidae, In Physa and in Ferrissia.
Among the Basommatophora, apical microsculpture has been accepted as of
taxonomic value within the Ancylidae, but otherwise its possible taxonomic-
phylogenetic significance in relation to taxa of all levels within the order has
so far been given only the slightest attention. The author here raises some ques-
tions about phylogenetic possibilities within the order in relation to punctation.
As for group-relationships within the Planorbidae, It is considered that, on the
basis of known anatomical characters, the Bulininae, the palearctlc Planorbarius,
the nearctlc Helisoma and the neotropical Taphius andecolus are all mutually re-
lated in some way. Accordingly, punctation might be sought fgr In Helisoma in
which it Is as yet unknown. On the other hand, a near relationship between Bulininae
and "Segmentlnlnae," both punctate, seems unlikely because of the known anatomical
dissimilarities.
^This investigation was supported (in part) by a research grant, E-2409, from the National
Institute of Allergy and Infectious Diseases, U.S. Public Health Service.
^Present Address: Museum of Zoology, University of Michigan, Ann Arbor, Michigan, U.S.A.
(115)
116 H. J. WALTER
A special affinity between the Bullninae and the punctate ?ancylid Burnupia is
suggested by published anatomical data. The existence of punctation in the presum-
J
ably primitive Ellobiidae, aside from pointing to their possible relationship to
Planorbidae, might be taken as evidence that apical punctae represent a primitive
character, although arguments to the contrary might be made.
From the literature it is concluded that past investigation and description of
shell microsculpture has been insufficiently exacting for adequate application to the
complex biosystematics within basommatophoran groups such as Bulinus.
INTRODUCTION No attempt was made to have the speci-

mens oi Bulinus specifically identified by
In a program of study of snail vectors other authorities. Since this study is de-
of Schistosoma Republic of Liberia,
in the signed to test the prevailing and past tax-
West Africa, it was noted that a pattern onomy, it is intended, in part, as a chal-
of punctation was the dominant sculptural lenge to the validity of the various species
feature of the embryonic whorls of the and forms that have been recognized. For
shell of some available Bulininae. This the purposes of this paper it is only neces-
finding led to a systematic examination of sary to use an accepted system of classi-
many Bulininae and limited numbers of fication in order to demonstrate that many
other Basommatophora, from Liberia and conchological forms from a wide geograph-
elsewhere, to determine the presence or ical area were included that represent a
absence of apical punctation in these shells significant number of the probably valid
as well as to an inquiry into the litera- species of the various species groups. For
ture so as to find out what might be known this purpose, it will be expedient to follow,
generally about such sculpture in order to in the main, Mandahl -Barth's (1958) sys-
confirm or refute previous reports for the tem of classification and to cite the illus-
species I was able to investigate. trations of shells in it corresponding in
The present paper, which is part of a form to those of the specimens studied
broader investigation on the morphological (Table I). In following his system of
basis of classificationin Basommatophoran classification for the identification of buli-
groups, deals with the results of this in- nine species and " subspecies", attempts
vestigation of the punctate character in were made to apply given characters of
conjunction with the results from the lit- the soft parts, as well as of shell sculp-
erature inquiry. ture, and sometimes of the radula, but,
mainly because of inadequacy of the de-
MATERIALS
scriptive information, it proved necessary
In this study over 1600 shells were ex- to rely almost entirely on the form of the
amined for the punctate character of their shell in conjunction with geographical con-
nuclear whorl and of these the large ma- siderations. The decisions on the specific
jority were of the species of African status thus made are for the most part
Bulininae that represent the 3 subgenera uncertain, because rather few of the shells
Bulinus s.S., Pyrgophysa Crosse, and closely matched the illustrations. Also,
Physopsis Krauss, of the genus Bulinus some series of specimens from a given
O. F. M311er,as they are recognized here*^ geographical area best match with figures
(Tables I and II). of a variant that is supposed to be re-
stricted to a widely distant area. Some
3The African Bulininae are now generally
specimens do not correspond to any of the
placed in the single genus Bulinus, with the
former genus Physopsis occupying subgeneric venlent for the purposes of this paper to use
rank, and the former genera Pyrgophysa and the old group names as subgeneric designations,
Bulinus combined in the subgenus Bulinus s.s. with Bulinus s.S. embracing the '4runcatus and
(see Mandahl- Barth, 1958). This grouping is tropicus groups," Pyrgophysa embracing the
based in part on anatomical data which are in- ^^forskalii group," and Physopsis embracing
sufñcient in my opinion, and I Und it more con- the ^âfricanus*' group of Mandahl- Barth.
,
PUNCTATION OF EMBRYONIC SHELL IN BULININAE 117
descriptions in particular and some series Sudan

of varying forms could as readily be as- 2 Khartoum and Khartoum area.
localities:
One compares to B.t. truncatus of the
lot
signed to one "species" or "subspecies"
innesi form: 53a. The other lot conforms to
as another. Even in assigning specimens the "dybowskii^' form of B. truncatus. Con-
to their subgenus, the given system could tributed by LeRoux and van der Schalie,
not be followed consistently. However, Tanganyika
from my personal experience with conchol - 1 locality: Mwanzd, Variants resemble B.
truncatus trigonus (Martens) and B. tropicus
ogy and soft anatomy, I was able to place
zanzibaricus (Clessin) and B. tropicus mu-
all species in their respective "subgenera" tandaensis (Preston): 54a; 46 b, d; 47b. Con-
with confidence. In one instance only some tributed by LeRoux.
doubt existed about the subgeneric assign- Kenya
ment of a series of "Pyrgophysa " ( the 1 locality:Kisumu, Lake Victoria. Partly
" cane s cens' form in Table I), in consid-
^
grown in laboratory, in London. These
the
closely resemble B. t. truncatus of the innesi
eration of certain microsculptural charac- form: 53a. Contributed by LeRoux. .
ters (see p. 123). Northern Rhodesia

Although the number of valid forms 9 localities: Mazabuka, Mbawa, and Lundazi
treated here is debatable, most of the areas. Some variants match with or tend
toward B. truncatus trigonus and B. coulboisi
generally recognized African species of
(Bourguignat), while some resemble the East
Bulininae assuredly were among those African B. sericinus (Jlckeli)^ of the trun-
studied, and certainly included are a num- catus group; most other variants correspond
ber of other forms that some malacolo- with or tend to resemble the "subspecies"
gists would call good species. tropicus s.S. of B. tropicus Krauss, (includ-
ing the "depressus" form of Haas), mu-
tandaensis (Preston), allimudi (Dautzenberg)
and angolensis (Morelet): 54b; 57a,c; 50a,d;
44a,d,h; 47a,c; 48a,d; and 45c,d. Some lots do
TABLE I. Bulinus s.l. studied
not especially match any of the figured forms
and other "subspecies" might be considered
Bulinus s.S. (25 localities)
to exist among them. In form many of the
Sardinia Island, Italy Bulinus s.S. in some of these lots closely
1 locality: Field origin, Santa Teodora (lab- resemble Physopsis of the forms ugandae
oratory-bred in London). The specimens cor- (Mandahl- Barth), globosus (Morelet) (includ-
respond to B. truncatus truncatus (Audouln) ing karongensis Smith), and africanus s.s.
form innesi Pallary: 53a. 4^ Contributed by (Krauss): 41c, d; 42b,j; 38c, Not infrequently
P. L. LeRoux. these samples actually included Physopsis
Iraq species, but I could separate these out in all
1 locality: Baghdad. The series corresponds cases on conchological characters by very
to B.t. truncatus (iorm innesi): 53a. Con- careful inspection. All lots contributed by
tributed by LeRoux. LeRoux.
Egypt Southern Rhodesia
4 localities: Hallaba, Sanañr, and Qalyub, all 3 localities: Bulawayo and Salisbury areas.
in Qalyub Province and one laboratory-bred Most specimens are referred to B. tropicus
strain from "Egypt." Variants in the labora- tropicus; some resemble B. t. angolensis:
tory strain closely approximate all of the fig- 44c,d,f; 45a. Contributed by LeRoux.
ures of B. t. truncatus', most of this labora- Union of South Africa
tory material and 2 of the field series, how- 1 locality: Cape Province. The lot compares
ever more closely resemble the innesi form: well with B. tropicus tropicus: 44c,d,f. Con-
53a; the remaining lot is more like the form tributed by LeRoux.
dybowskii Fisher (see Demian 1960, PI. I). Ghana
Contributed by H. van der Schalle, except for 2 localities. Tamal a (laboratory-bred); "Gha-
the laboratory strain which was descended na". These specimens resemble B. trop-
from a laboratory colony at the Tropenin- icus tropicus and B. truncatus rohlfsi (Cles-
stitut, Hamburg, Germany and which was sin): 44g,h; 55a,b. Contributed by F. Wick-
reared by the author in Liberia. remasinghe.
5 Mandahl- Barth considered B. sericinus to be a
4 The Fig. numbers cited here refer to the fig- member of his '^tropicus group" (1958) but
ures in the plates in Mandahl- Barth (1958) later (1960) transferred it to his "truncatus
unless specifically stated otherwise. group".
118 H. J. WALTER
Pyrgophysa (28 localities) Physopsis (53 localities)

Egypt Tanganyika
1 locality: Qalyub Province. The specimens 2 localities: Mwanza area. Both collections
appear to be fairly typical B. /orsfeaí¿¿(Ehren- consisted of undoubted B. nasutus (Martens):
berg), but ribbing of the shell is not well- 40b, Contributed by LeRoux and McClelland.
developed, and the sculpture is weak, muchas Northern Rhodesia
in: 59f. The larger shells {8-12mm long) tend 9 localities: Lundazi, Lusaka, and Mazabuka
toward: 59a, or j. Contributed by van der areas. Many of the shell forms figured
Schalle. for B.globosus Morelet) and B. africanus
Tanganyika (Krauss) are represented in these collections,
2 localities:Mwanza area. The shells of both but most tend to resemble B. globo sus: '^21.
series are much like the Egyptian material of Contributed by LeRoux,
B.forskalii cited just above, and the above Southern Rhodesia
statements equally apply. Contributed by 2 localities: Salisbury area. Much as for the
Le Roux and F. W. J. McClelland. Northern Rhodesian material cited just above,
Northern Rhodesia various shell forms of B. globosus and B.
1 locality: Lochinwar, "in a cattle tank". africanus appear to occur in these two lots;
The shells in this series are rather smooth one large series especially conforms to B.
and rather thickwalled and correspond to the africanus africanus: 38a, b. Contributed by
"canescens Morelet" form of B. forskalii; LeRoux.
they apparently are from the same original Ghana
collection of LeRoux as those figured: 59k,l. 1 locality:near Kumasi. The one young speci-
Contributed by LeRoux. men resembles B. globosus and it is especially
Southern Rhodesia strongly sculptured: 42a, Contributed by Wick-
1 locality: Bulawayo. Shells of this large remasinghe.
series r<esemble the '^canescens Morelet" Liberia
variant of B. forskalii in form: 59k, However, 39 localities: At sites scattered widely over
the shells are not smooth, but are strongly the Central and Western Provinces. Much ad-
ribbed, with clearly developed "shoulder- ditional material was obtained from a labora-
keels" as in more typical B. forskalii, and tory-bred colony in Liberia that was started
they vary toward: 59c. Contributed by LeRoux. with specimens from one of the Central Prov-
Union of South Africa ince populations. The shells mostly conform
1 locality: Cape Province. This large series to B. globosus especially, but some appear
particularly resemble B.forskalii as ñgured closer to B. africanus africanus: 42c,h; 38c.
in: 59c. Contributed by LeRoux. Some variants from the field especially could
?Unlon of South Africa be taken for B. ugandae (Mandahl- Barth):
1 locality: Kanlayis. Most of the specimens 41c. Collected (or reared) by the author.
resemble the ^^ cane s cens*'* variant of B.
forskalii in form: 59k, The shells however
have well-developed ribbing instead of being TABLE IL Material studied, other than
smooth. Several small, short, and obese
Biilinus S.I.
specimens among these must be taken to be
B. reticulatus Mandahl- Barth: 58b,c. Con-
PLANORBIDAE
tributed by LeRoux.
Ghana Bulininae
1 locality. Ashanti region. The remarks made Indoplanorbus exustus (Deshayes): India, 1
above for the series of B. forskalii from locality, UMMZ6 No. 80584; Ceylon, 1 lo-
Egypt and Tanganyika also apply to this lot. cality UMMZ No. 80582; and Afghan fron-
Contributed by Wickremasinghe. tier, 1 locality UMMZ No. 80583.
Liberia Planorbinae
20 localities: Eastern, Central and Western Planorbina ( =Biomphalaria) pfeifferi gaudi
Provinces. Additional to these was abundant (Ranson)'^ : Liberia, W. Africa; several lo-
material from a laboratory colony started with calities in the Central Province, and 1 lab-
specimens from one of the Central Province oratory-bred colony descended from speci-
localities and reared through several genera- mens from one of these localities. Collected
tions in Liberia. Most of the field and labora- and reared by the author.
tory shells correspond especially to B.for-
skalii, and some variants seem more like B.
scalaris: 59a,b,d,g; 60a,b. The few quite large 6 University of Michigan, Museum of Zoology
specimens from the Ûeld (about 15 mmin catalog number.
length) closely resemble the figured shell ^There seems to be no doubt that, despite some
types of B. forskalii: 59i,j. Collected (or controversy, Planorbina Haldeman 1843 is the
reared) by the author. oldest valid available name for the Planorbinae
Planorbina (=Australorbis) glabrata (Say): 1 character was in its occurrence and rela-
strain, field origin, Paramaribo, Surinam; tive development among individuals of the
reared by the author in the laboratory in colonies, and to work out appropriate
Liberia; descended from an old stock at the
techniques of observation, for which large
Tropeninstitut, Hamburg, Germany.
Gyraulus costulatus costulatus (Krauss): Li- amounts of expendable material were re-
beria, 1 locality in the Central Province, quired. Each laboratory strain provided
and a laboratory-bred strain descended abundant eggs from which large numbers
from specimens therefrom. Collected and of protoconchs were obtained and each also
reared by the author.
provided numerous shells of all growth
HelisomatinaeS
stages up to adults. With these series it
Planorbarius comeus (Linné): England; 1 lo-
cality, outskirts of London. Collected by the was possible to observe how erosion and
author. encrustation of shells with algae and other
ANCYLTOAE environmental materials affected apical
Ferrissia (Walker) sp, or spp, Liberia;
:
microsculpture as well as observation of
several localities in Eastern, Central and the sculpture itself. Use of embryonic
Western Provinces. Collected by the author. shells facilitated study of sculptural de-
LYMNAEIDAE tails at high magnifications (see below)
Radix natalensis (Krauss): Liberia, llocality, with a compound microscope, and allowed
Central Province, and a laboratory-bred critical observations on the nature of
strain descended from specimens there-
punctae.
from; collected and reared by the author.
Sierra Leone, 1 locality; contributed by The second phase of the investigation
E. G. Berry. was concerned firstly with the examina-
Pseudosuccinea sp: Union of South Africa, 1 tion of much field material of Pyrgophysa
locality, Pokkraal. Contributed by Le Roux. and Physopsis, the two endemic bulinines
" Stagnicola palustris group" 1 strain,
of Liberia, that were collected at various
:
field origin, London, England, bred in the

places, and secondly with examination of
laboratory in London, Contributed by
Le Roux. material of many forms oiBidiniis s.l.
from many other places in Africa and from
PHYSIDAE
Physa sp: 1 lot, origin and collector uncer- some adjacent areas (see Table I). In
tain; ?Liberia. this phase a "spot -check" procedure was
Physa integra (Haldeman): U.S. A., 1 locality, followed: to reach conclusive decisions
Michigan, UMMZ No. 60492. regarding the general presence or absence
of punctation in the Liberian Bulininae,
the series investigated were so chosen as
METHODS to include specimens from the central and
peripheral portions of their known ranges
The investigation of the occurrence of in the country, and also the more extreme
punctation in Bulininae and its relatives conchological variants. In the available
was divided into three phases. The first collections from areas outside of Liberia
phase consisted of a concentrated study of it was only in rare instances impossible
many individuals from one colony each of to reach a decision for a particular series
the bulinine groups Bulinus s.S., Pyrgo- or variant, for reasons explained below.
physa and Physopsis kept on hand in lab-
, Some of the non-Liberian series were
oratory aquaria at the Liberian Institute laboratory colonies reared elsewhere, and
for Tropical Medicine in Liberia. The these contained shells of many juvenile,
purpose of this part of the study was to and sometimes embryonic, young; in these
determine how consistent the punctate samples the presence of punctation was
determined for many individuals.
acting as intermediate hosts of Schistosoma
The third phase of the Investigation cov-
mansoni including the nominal genera Biom-
ered at least several specimens of various
phalaria Preston 1910 and AMsíraZor6¿s Pilsbry
1934. species of non-bulinine Planorbidae, and
8f. C. Baker, 1928. at least a few shells of species of some
-
120 H. J. WALTER
Other basommatophoran families. Only terminations, it was necessary to illum-

a few collections were available for this inate the sculptural structures from many
phase of the work, the main purpose of different angles, and often the exact angle
which was to determine if embryonic of incidence of the light, arrived at by
punctae might ever be fully lacking in any repeated trial and error, proved to be
member of these groups. One breeding critical. In using this mode of illumina-
colony each of Planorbina (=Australorbis) tion the shininess and translucency of the
glabrata {Say),1} bina (=Biomphalaria) surface of the shells cause interference
pfeifferi gaudi (Ranson), G yrawZMS costu- with visual resolution of surface details,
latus (Krauss) and Radix (=Lymnaea) na- and even in the best material the presence
talensis (Krauss) were maintained at the or absence of punctation sometimes may
laboratory in Liberia and provided shells be left somewhat in doubt even after in-
of embryonic young for study. Part of tensive study at high magnification (see
this phase of the research was carried below). The method helped however, in
out in the Mollusk Division of the Museum determining the physical nature of micro-
of Zoology of the University of Michigan, sculptural elements. Attempts were made
in the United States. to stain shells darkly in hopes of reduc-
Early in this study it was found that ing the surface translucency that inter-
shells could be cleaned by brushing, after fered with observations, and aqueous solu-
they had been immersed for a few min- tions of basic fuchsin proved to be of
utes in full-strength commercial Clorox some aid in this respect. Also, the spire
(sodium hypochlorite), a technique used on occasion was filled with black ink to
routinely thereafter, for fresh or "alco- provide a dark opaque background against
holic" specimens. Even when exposed for which sculptural highlights might be more
weeks, the shells are not altered physi- readily seen.
cally. This treatment completely removes It was found that shells, even from em-
the periostracum, external dirt and in bryonic young, that had been cleaned with
ternal traces of the body, without affecting Clorox, readily turned white and opaque
the translucency of the shell. The thinner, when only briefly exposed to a variety of
smaller shells cleaned this way can be agents in aqueous solutions, which include
studied by transmitted light; in embryonic hydrogen peroxide in the ordinary weak
shells especially, the microsculptural de- concentration of commercial brands, a
tail can be seen with great clarity. Shells common detergent ("Tide") in low con-
of progressively larger sizes are in gen- centrations acids and potassium hydroxide
,
eral more thick-walled, especially in the in low concentrations, as well as sodium

region of the apical whorls, or of darker bicarbonate in high concentrations. Most
color, and will transmit less light, so that or all of these agents cause a general
microsculptural details are seenwith diffi- surface erosion that rapidly erases the
culty or not at all clearly. Although most fine microsculpture. The very whitened
of the field material consisted of adult shells, as would be expected, appear dark
material, in all but a few series there and featureless when placed between the
were at least a few shells sufficiently observer and a strong source of light.
small and thin to allow proper study by Sometimes parts of shells that had been
transmitted light; sometimes egg masses preserved with the animal proved to be
found preserved with the series yield- whitened and opaque, apparently as a re-
ed embryonic shells for study. In sult of chemical erosion of the internal
colorless shells, such as those of most surface of the whorls, and this on occa-
Pyrgophysa, microsculpture was demon- sion was so pronounced as to prevent de-
strated more readily. Adult shells that termination of microsculpturalcharacters
could not be studied by transmitted light in most specimens of a series. In some
had to be viewed by less satisfactory re- series the nuclear whorl itself was de-
flected light. In making the desired destroyed as a result of normal environ-
FIG. 1. Apex of shells of Bulinus s.l. showing punctation on the upper part of the embryonic
whorl (highly enlarged).
a) B. (Bulinus) truncatus (Audouin), Egyptian laboratory strain, adult specimen.
b) B. (Pyrgophysa) forskalii (Ehrenberg), Li berian laboratory strain, adult specimen.
c) B. {Physopsis) globosus (Morelet) , Liberian laboratory strain, young specimen.
mental erosion but in practically all cases especially in those having much expanded
post-embryonic sculpture sufficed for a post-nuclear whorls, the embryonic shells
demonstration of punctation in a number may be hidden almost entirely. In cases
of specimens of these series, when punctae like the latter, when it was desired to de-
were present. termine if punctae might be present on a
Microsculptural structures are much small and very early fraction of the first
easier to demonstrate in shells having whorl, it was necessary to break the post-
more scalariform (more "loosely coiled") nuclear whorls away to free the proto-
apical whorls (e.g., "Pyrgophysa" forms). conch for study. In such discoidal shells,
In more "tightly coiled" forms (such as the internal shell deposits on the first and
most "Physopsis") the second whorl hides second whorl are so laid down that they
the larger part of the nuclear whorl and cover and obscure much of the nuclear
its sculpture, while in discoidal forms, whorl and its sculpture, and cannot be re-
122 H. J. WALTER
moved; for this reason, embryonic speci- seemed to be elevated ''nodular" struc-
mens are of particular value in the study tures. However, close study of many
of the apical part of such shells, as was specimens at magnifications of 500X and
found in investigating the microsculpture sometimes up to 12 50X and observations
oi Planorbina send Gyraidiis. made at lower magnifications with re-
When critically illuminated by trans- flected light, showed that the punctae are
mitted light, nuclear punctation often can well defined pits in the shell substance.
be seen fairly definitely at magnifications These pits were seen to be arranged in
as low as 12. 5X, but then only in very an intersecting pattern of conspicuous
clean material that is in excellent condi- spiral rows and more or less definite
tion. It is most often necessary to use axial rows. Differences between the punc-
magnifications of 32X, and over, as was tation of Bulinus s.S., Pyrgophysa and
done routinely in this study, for clear Physopsis were observed but basically the
resolution of individual punctae when using patterns were the same. There was no
transmitted light. In using reflected light, tendency for the punctal pattern to vary in
on the other hand, determination of the degree of development, nor to tend toward
existence of punctae was often uncertain obsolescence, although a consistent differ-
even at magnifications as high as 80X. ence in its relative prominence among the
Magnifications of 500X and even up to three subgroups of Bulinus was noted. In
1250X (using an oil immersion objective) general it was easiest to discern puncta-
were used in checking sculptural micro- tion in the Pyrgophysa and most difficult
detail from time to time, sometimes while in the Bulinus truncatus, partly because
employing reflected light. Routinely the of the differences in the size, thinness,
shells were kept immersed in fluid, usu- color, and manner of coiling of the shells,
ally water, while being studied. but also on account of some difference be-
tween the 3 groups in the relative coarse-
ness of the punctae. For the very many
RESULTS
laboratory- bred adult and undamaged B.
During the firstphase of this study, in (Pyrgophysa) forskalii that were studied,
the examination of the hundreds of embry- the nuclear punctation was always clearly
onic shells oiBulinus (Bidimis) trimcahis demonstrated, even in shells which ex-
of the Egyptian strain and of the Liberian ceeded 10 mm
in length. In examinations
strains of B. (Pyrgophysa) forskalii and of the many laboratory-bred ß, (.) trun-
B. (Physopsis) globosus that were bred in catus and (Physopsis) globosus, alihowgh
the laboratory, punctation was observed in it was more difficult because of their
all instances (see Fig. 1, a, b, c, but note darker, thicker, broader and more tightly
that the shells illustrated have grown be- coiled shells (especially the larger ones
yond the proto conch stage to different which often reached a length of 12 mm),
sizes). The character proved to be con- nuclear punctation was always demon-
spicuous in unmarred and clean embryonic strated in specimens that were relatively
specimens when observed with adequate undamaged, when sufficient care and effort
magnification and when properly illumi- was used. Punctation was observed to
nated by transmitted light. continue onto the postembryonic whorls,
It was found that, depending on slight but to an extent that differed among the
differences in orientation of the specimen strains representing the 3 different sub-
with respect to the source of light and genera.
with respect to one's eye, eachpuncta ap- In the relatively few instances when
pears as a brilliant microscopic spot of punctae were unidentifiable in laboratory
light, or as a dark minute ring enclosing specimens, tell-tale whitening and opaque-
a clear spot, or when foreshortened on ness of the shells indicated that they had
curves, as a dark or bright short dash. been accidently exposed to an erosional
Commonly, at lower magnifications, they agent; or else, in older individuals, com-
píete destruction of the nuclear whorl onstrated in at least one or in a few in-
with punctation had resulted from the
its dividuals, and the punctate condition was
more usual kind of erosional process that definitely observed in over 1500 speci-
also occurs in field colonies. It was es- mens (material from the laboratory col-
tablished beyond a doubt then, that nuclear onies included), that represented many
punctation was a constant and always fully species or forms, including members of
developed character in the three strains Bulinus s.S., Pyrgophysa and Physopsis.
of laboratory-bred Btdinus s.S., P y rgo- As for the only remaining bulinine
physa and Physopsis. species, Indoplanorbis exustus, nuclear
In observations of the Bulininae of punctationwas definitely observed in some
strains of other geographical origin that of the few shells from India and Ceylon
were bred in other laboratories, and in that were studied. In these adult speci-
examinations of field collections from Li- mens the nuclear whorl was not in good
beria, and from other African countries, condition and was quite thickened; this
and from the Mediterranean and Middle condition and the discoidal shape made
Eastern areas, nuclear punctation was the desired observations difficult, so that
again identified in hundreds of specimens, the punctal pattern could not be made out
except in a few particular instances, in very clearly or be accurately compared
which interfering factors, corresponding with that found in Bulinus s.l. The punc-
to those described above were clearly tae seemed to be relatively fine.
involved. However, even when the nuclear Among the other Planorbidae studied, a
whorl was found to be lacking, the origi- conspicuous punctal pattern was observed
nal presence of nuclear punctae was usu- on the nuclear whorl of the several speci-
ally inferable from the presence of post- mens of Planorbarius corneus from Eng-
embryonic punctation quite like that found land that were examined. In contrast,
in uninjured shells. Small shells, par- nuclear punctation was found to be totally
ticularly of hatching size, and embryonic lacking in all of some dozens of the ex-
ones obtained from occasional preserved amined specimens of Planorbinai" Biom-
egg masses, were in all cases seen to omphalaria" and "Aiistralorbis") from
have the conspicuous basic punctal pattern. Liberia and from Surinam and in the
Again, forms of Pyrogophysa from the Gyraulus from Liberia. Of the non-plan-
field, exception of one series,
with the orbid snails that were studied, the embry-
were found have a somewhat more
to onic whorls of the Physa integra and
prominent and apparently coarser puncta- Physa sp., the Ferris sia spp., and the
tion than the other Bulininae. The ex- several species of Lymnaeidae, were in
ceptional series was that of the '^can- all cases observed to be quite devoid of
escens form oi B. (P.) for skalii" in which punctation.
the punctae appeared relatively fine (and During this study the observations on
in which the development of post -nuclear punctae became involved to some extent
punctation apparently was more like that with consideration of other microsculp-
which have observed in Bulinus). Other-
I tural elements of the nuclear and post-
wise, various forms oí Bulinus s.s. in- nuclear whorls. Some of these elements
cluding those of B. tropicus consistently might be referred to as nodules or as
appeared to have the least prominent and minute elevations and depressions, all of
somewhat the finer punctation, as was which occur in spiral series following the
found earlier in the laboratory -bred B. direction of the whorls and which are ar-
truncatus colony. ranged also in more or less definite trans-
In the study of Bulinus s.l., then, for verse or axial series. These elements
practically all series that were obtained show a considerable range of diffère ntation
from the many places distributed over in form,magnitude,complexityof arrange-
essentially the whole geographical range ment, and visual prominence at the micro-
of the genus, nuclear punctae were dem- scopic and macroscopic levels. Although
124 H. J. WALTER
these structures tend to obscure punctae, microscopic dots, arranged in spiral but
particularly on certain parts of the whorls, not radial rows", and of the whorls of
and may in part tend to be confused with '^ Bulinus (Physopsis) africanus " he re-
them, careful observations made at ade- marks (p. 511): "1st minute, bearing ex-
quate magnifications have repeatedly shown tremely faint, fine transverse microscopic
that the punctae are a special type of wrinkles, 2nd with stronger transverse
structure that is well differentiated from sculpture and microscopically engraved
other sculptural features of the shells of with punctate dots, arranged in regular
the snails that were investigated. It may radial and spiral lines, which usually dis-
be added that the relationship between the appear about the 3rd whorl. .
" In a ref-
.
punctal pattern and the other sculptural erence to "Bulinus (Physopsis) globo sus"
structures gives the impression that the (p. 512) Connolly gives no account of any-
post -embryonic sculpture, in a sense, thing like "punctate dots, " but implies
"evolves" from the punctae. that this species is similar to B. (Phys-
opsis) africanus in that respect. As for
the many other Basommatophora that are
LITERATURE ON NUCLEAR treated in the same monograph, specific
PUNCTATION mention of punctation of the shell was
This inquiry into the literature was in- found for one or more species of the
tended to reveal specific information on genus Segmentina of the Planorbidae, of
punctation of the embryonic whorl par- Burnupia of the Ancylidae, and oí Me lampus
ticularly in Bulininae. Secondarily, it was ¡Lnd Phytia of the Ellobiidae. For none
concerned with relating such information of these, except Burnupia is the puncta-
to any existing evidence of a taxonomically tion stated to occur specifically on the
and phylogenetically significant pattern in first whorl or embryonic shell.
the distribution of the character among Among 4 species of Segmentina that
other Planorbidae and among other Basom- Connolly describes, only the one repre-
matophora. It also became involved with senting the "subgenus" Hippeutis was
some other aspects of microsculpture, as stated (p. 496) to have punctation, which
well as with certain other taxonomical- was said to be 'dense and] microscopic
'
[
' ',
morphological matters, for reasons evident and which was said to be "probably of no
from the quotations cited below. [taxonomic] value". As for Burnupia,
As a means of bringing the results of Connolly, following Walker (1923) and
this study on the occurrence of nuclear others, refers to "radially punctate"
punctation into perspective in regard to its sculpture, usually if not always occurring
possible significance to current system- on the "extreme apex" of the shell, as
atics of the snails being considered here, one of the diagnostic characters of this
extensive pertinent quotations, mostly from genus of many nominal species. He says
the more comprehensive works of leading in regard to 5 species of his section
authors follow. Melampus of the genus Melampus that
In reviewing Connolly's (1939) mono- the " early postapical whorls" are punc-
graphic treatment of the freshwater fauna tate, but no account is given of puncta-
of South Africa for information on punctation descriptions of 2 other species.
in
tion of the whorls of the embryonic shells The apical whorl of one of these species
of Bulininae, a series of informative state- of Melampus and the "extreme apex" of
ments were found. He says in regard to the one species of /^/ that is described
"Bulinus tropicus" (p. 501): " 1st fwhorl] are said to be "smooth" (p. 467, p. 462),
smooth" and in regard to "Biilimis di- and the same is stated or implied for
aphanus" (p. 505) (both Bulinus s.S.) related species that are considered by
''apical fwhorl] smooth". In reference to that author. In speaking of his section
Bulinus [Pyrgophysa] forskalii" he says Melampus oi Melampus, he warns (p. 466)
(p. 508): "extreme apex engraved with that punctae are only identifiable in a
shell that is "fresh enough to show it," dick' s conclusion refers, in part, to ana-
and that they might erroneously be taken tomical considerations raised by him in
to be lacking in species described from the same paper. His account gives the im-
shells that are not 'well -pre se rved".
' Fbr plication that the nuclear whorls of Bulinus
Pythia scarabaeusihinnê), another ellobiid s.S., at least usually, have no "small dots",
H.Harry (1951) gives a clearly illustrated and it perhaps also implies that Physopsis
account of apical punctation also. may exceptionally have "striae" instead
In Pilsbry and Bequaert's monograph of dots. These quotations account for ail
(1927) on freshwater mollusca of another of the data on shell punctation found in
large African region, "spiral lines of these two papers. Shell characters are
punctures" on the embryonic whorl were dealt with quite briefly in these and other
described (p. 129) as a character of works of that author that are known to
Segmentina and of the "subgenus of Plan- me which deal primarily with interpreta-
orbis ", Hippeutis. The authors also actions of the taxonomic significance of
count for apical punctation in Burnupia. characters of the soft parts of the snails.
For the remainder of the many Basom- Mandahl -Barth (1954), in reference to
matophora (Planorbidae, Ancylidae, Lym- African Bulininae, says (p. 99): "The
naeidae, Physidae, and EUobiidae) that sculpture of the shell consists in most
are given attention in the monograph, no cases merely of delicate growth lines, but
other mention of punctation was found. in some forms ribs or spiral sculpture
In an extended systematic account of many may be present"; later (1958, p. 52) he
species and forms oí Bidinus s.l., the states: "A spiral sculpture consisting of
only statement found (p. 146) that might delicate lines or nodules may be present
pertain to punctae is a reference to "spiral in some forms, especially on the upper
impressions" in " Physopsis africatm half of the shell. '' The same author says
globosa ". oí "Bulinus (Pyrgophy sa) forskalii^' (1954,
Of the papers of Hubendick dealing with p. 110) that "The first whorl (the embry-
systematics within the Bulininae and Ba- onic shell) is smooth" and again later, of
sommatophora, two will be considered the whorls oí "Bulinus (Bulinus) forskalii"
here, one that treats Bulinus s.l. (1948) (1958, p. 85), that: "the first one is nor-
and another the Planorbidae as a whole mally smooth." He describes (1954, p.
(1955). In the former paper, after discus- 109) the first whorl of "Bulinus reticu-
sing "^^ senegalensis (Müller)", the
type species of Bulinus, and '' Bulinus
latus n. sp.", which he later assigns to
his forskalii group (= Pyrgophysa) , as
forskalii" as species having a particular "smooth" and restates this (1958, p. 82)
type of elongate shell in common (p. 33, later. Mandahl -Barth further states (1958,
35), Hubendick describes the sculpture of p. 59) that "Physopsis has, as a rule, a
that shell-type by saying (p. 43): " More- distinct sculpture consisting of spirally
over, the shell shows a sculpture crossing arranged rows of small impressions — a
the direction of the whorls." Also, in re- type of sculpture never found in Bulinus
ference to the shell of Bulinus s.S., he s.S." and, also in the same paper (p. 60) ,
states (1955, p. 523): " The nuclear whorls in reference to his "Africanus" group
are usually providedwith striae" and: "In {=Physopsis): "In most forms a charac-
Physopsis\he columella is generally trun- teristic sculpture, consisting of spirally
cated. The nuclear whorls are usually not arranged rows of small transverse im-
striated but have small dots spirally ar- pressions or nodules, may be visible on
ranged. Sometimes, however, the correla- the upper part of the shell at a magnifi-
tion between these characters does not cation of about 25X. This sculpture has
hold good. All this means that Bulinus not been found in any of the forms belong-
and Physopsis should be regarded as sub- ing to the other groups." Here "other
genera of one genus {Bulinus) rather than groups" refers to all African Bulininae
as distinct genera." By inference, Huben- except Physopsis species. Referring to
-
126 H. J. WALTER
Bulinus (Phy sop sis) globo sus (1954, p. 113 lengthy treatment of shell characters in
that author reports that "very delicate, Bulinus tnmcatus of Egypt, give any indi-
small transverse lines, arranged in spiral cation regarding nuclear punctation.
series" occur on the ''upper whorls". In F. C. Baker's (1945) most detailed
"B. globo sus ugandae n.subsp.", he says and comprehensive work on systematics
(1954, p. 114), "completely lacks the spiral in Planorbidae other than Bulininae, there
sculpture" that is found in typical B. is no mention of punctate sculpture in his
globosus. For the same form (as species account of the two "Bulinidae" he deals
B. ugatidae) he later says (1958, p. 59 ) with, Isidora (=Physopsis) globosa and
that it has "no particular sculpture at all" Indoplanorbis exustus. As for the 55
and, in the same work (p. 66), he reaf- genera and subgenera that he treats as
firms this by saying that the species Planorbidae, he gives a systematic ac-
shows a "complete absence of spiral counting of the diagnostic conchological
sculpture on the spire". In an addendum characters of each, and also devotes a
(1960) to his 1958 monograph there is one special section to the shell characters of
further pertinent statement referring to Planorbidae as a group. The only infor-
the newly recognized subspecies Bulinus mation referring to shell -punctation ap-
(Physopsis) nasutus nasutus (p. 568): "The plies to his subfamily Segmentininae, with
microsculpture consists of spirally ar- definite specific reference to punctation
ranged rows of minute nodules or someof the embryonic whorls for each of 6 of
times punctures, as a rule more coarse the 10 recent (non-fossil) genera he placed
on the upper whorls and becoming finer in that category: Segmentina, Hippeutis,
on the lower, but usually covering the en- Polypylis, Pvigiella, Drepa^wtrema (Dre-
tire shell. In other species of Physopsis panotrema s.s. and subgenus Fossul-
the microsculpture is restricted to the orbis) and Platytaphius, and ?Helicorbis .
spire." Mandahl-Barth (1954) also has He utilizes such terms as "spiral rows
described the apical punctation in the an- of small pits", "punctures", and "fine
cylid Burnupia. punctations" in the descriptions. He
C. A. Wright (1956) deals taxonomically treats the punctate character as if it were
with "6 species" of Bulininae from the one of the consistent features of each of
Senegambia in northwest Africa. For 5 these "genera" and, at one point (p. 107),
of the 6 bulinine forms, he reports the he seems to imply that it might be diag-
existence of a "punctate pattern" or a nostic for the "subfamily" but does not
"fine punctate pattern" on the "first account for that character in the 4 other
whorl" or "nuclear whorl." The five re- genera of Segmentininae. Since he gives
portedly punctate species are "Bulinus relatively close attention to conchological
guernei (Dautzenberg)" and "Bulinus seri- matters in his taxonomic treatise, one
cinus (Jickeli),"both members oi Bulinus would feel that his failure to mention
s.S.; "Bulinus ludovicianus (Mittre)" and shell -punctation for so many species and
"Bulinus forskalii (Ehrenberg), " both species groups means that the character
Pyrgophysa forms; a.nd "Bulinus jous is probably absent in most Planorbidae.
seaumei (Dautzenberg)" of the subgenus Further reports on the occurrence of
Physopsis. To my knowledge he is the punctation in "Segmentininae" (in addition
first ever to account for punctate sculp- to Connolly's (1939) ioT Segmentina (Hip-
ture in Bulinus s.S. peutis) have been recently given by Par-
)
Although general sculptural features of aense and Deslandes (1956a, b; 1958) for 3
the shell, including the presence of "nod- species of Drepanotrema. In dealing with
ules" in Physopsis, are described by a series of other mostly neotropical plan-
Amberson and Schwarz (1953) in their orbids, including other "Drepanotrema"
account of African Bulininae, there is no species, these authors have not otherwise
definite reference to apical punctation. reported the punctate character in their
Nor does Demian (1960, pp. 10-12) in a systematic and quite careful conchological
.
PUNCTATION OF EMBRYONIC SHELL BULININAE 127
descriptions. clear punctation can be overlooked readily

No records were found in the literature in shells unless they are relatively free
of the occurrence of punctation in Plan- of injury and unless very special care and
orbina (Australorbis and Biomphalaria) special techniques in cleaning and examin-
ing the specimens are applied. This cir-
cumstance seems to account in large meas-
DISCUSSION AND CONCLUSIONS ure for the lack of reports on punctation
The preceding inquiry into the literature in so many cases.
shows that punctation of the embryonic The literature inquiry has shown that,
whorl or apex of the shell has been previ- in general, characters of microsculpture
ously reported by some authors for some in Bulinus s.l. have not been determined
species of each of the three subgenera or applied in descriptive usage with suffi-
constituting Bulinus sA. Punctation was cient exactitude. Above we have seen
recorded earliest and most frequently for references to "striae, "to "small dots," to
Physopsis, but only very recently for "small nodules"and to "spiral rows of
Bulinus s.S. (where it is finest), its pres- small transverse impressions'* and the
ence in the former being sometimes con- like, which give no more than "impres-
sidered a diagnostic difference. For sions" about the nature and magnitude of
Pyrgophysa, in which I found the dots the sculptural elements. The ambiguity
easiest to see, some records were also of such terms is shown by my findings
found. Taken on the whole, the literature that the spiral rows of small (transverse)
considered seems to indicate that the pune - impressions which are not truly pits, co-
tation varies in degree of development, exist with spiral rows of punctae, which
even to the point of obsolescence, among are well defined pits, and that spiral rows
individuals of the species. The litera- of small more or less nodular raised
ture would imply also that in Bulinus s.s. "lines" (also transverse), coexist with
and in the Pyrgophysa group, the nuclear these. Authors, as quoted earlier in this
whorls usually lack punctation, and indeed paper, refer to shells of particular spe-
may lack sculpture (may be "smooth") cies of Bulinus as having small dots or
entirely. However, the results of my in- striae and minute nodules "or sometimes"
vestigation show that these reported ab- punctures, as if such characters were
sences of punctation in Bulinus s.l. are mutually exclusive. Statements to the
in error. My findings show that puncta- effect that spirally arranged rows of small
tion of the embryonic whorls must be impressions are a type of sculpture never
universal in Bulinus s.l. over its entire found in Bulinus s.S., that some species
geographical range, and that it even oc- completely lack spiral sculpture, and that
curs in the only other bulinine genus the microsculpture is restricted to the
known, Indoplanorbis . Wright's findings spire in species of the Physopsis group,
on Bulinus of the Gambia, (a corner of are erroneous. Despite the fact that many
Africa from which I had no material for bulinine shells seem to have smooth por-
examination), go along with my above con- tions when viewed at low or no magnifica-
clusion regarding 5mZ¿wms s.l. It has been tion and despite the fact that some forms
shown here that nuclear punctation in the (e.g. the "canescens " form of B. forskalii)
species of this genus is a character that may appear especially smooth, I have ob-
almost certainly is always fully developed served striation and spiral sculpture, at
and that does not vary towards obsoles- least at high magnifications, in all of the
cence. It was shown also, that in proper great many such examples in which I have
material of Bulinus s.l. (such as shells sought for it carefully. Also, apparently
from very young or embryonic snails) and ! Bulinus s. 1. have the nuclear punctae
with proper equipment and techniques, arranged partly in radial as well as spi-
nuclear punctae always can be observed ral rows, despite Connolly's (1939) con-
clearly. It was also shown here that nu- trary statement regarding B. forskalii.
.
128 H. J. WALTER
The relationship of these findings to and therefore one may expect that existing
the presently unsatisfactory state of tax- punctation has yet to be discovered in non-
onomy in ulinus s.l. is clear. The bulinine species that have been described.
difficulties I encountered in trying to My finding of well -developed nuclear pune -
identify my bulinine material illustrates tation in Planorbarius would support such
this situation. Although it may well be a probability.
that the genetic relationships involved are Difficulties in demonstrating conchologi-
so complex and of such a nature that a cal minutiae, as was attempted in the
convenient and consistent classification of usual ways and with the usual museum
the forms may be impracticable, it is collections of small series of large and
obvious that much thoroughly documented relatively opaque and often eroded and
finely delineated morphological detail in- dirty shells, seem to have prevented stu-
cluding that of the shell, which is not on dents of Planorbidae from giving cogni-
hand at present, is a prerequisite for the zance to the possible taxonomic - phylo-
making of a valid attempt at such a classi- genetic significance of embryonic micro-
fication. It should be supposed that exist- sculpture. Among other gastropods,
ing morphological differences between characters of the embryonic whorls have
species of snails, as for other organisms, been reported as of value in the system-
are in general of such slight magnitude atics of the streptoneuran subclass Pro-
that they need to be defined in explicit, sobranchiata (Iredale, 1911). In regard
refined terms. In this sense, and in re- to the euthyneuran Basommatophora, api-
gard to microsculpture, it can be said cal microsculpture of the shell has been
that the Bulininae are yet poorly known, recognized as of importance in discern-
despite the fact that a great many species ing taxa in the heterogeneous assem-
and their varieties (or ^'subspecies") were blage of freshwater limpets, all formerly
originally diagnosed on shell characters placed in the Ancylidae, which has al-
alone and then were subsequently rede- ready been subdivided on various grounds
scribed by later authors. It may be noted (Walker 1923; Burch 1961, 1962), al-
that my findings of nuclear punctation in though further revision is needed. Re-
Indoplanorbis brings conchological data, garding the Planorbidae, as cited earlier
for the first time, in direct support of in this paper, F. C. Baker (1945) has in-
the data on characters of the soft parts dicated that punctation of embryonic
on which that genus had been placed in whorls may be characteristic for genera
the Bulininae. of his subfamily Segmentininae, while
In respect to the occurrence of nuclear for Ellobiidae, as also cited ear-
the
punctation in Planorbidae other than the lier, Connolly (1939) implies that " post-
Bulininae, the literature reveals that sev- apical" punctation might be character-
eral workers have observed punctation istic of the shells of all species of the
(either on the nuclear whorl or in the ''section Melampiis" of the genus Me-
region of that whorl) in various species lampus
referable to F. C. Baker's subfamily As for the possible relationships be-
Segmentininae, and have failed to see tween particular punctate taxa within Plan-
punctae in other planorbids. Their nega- orbidae, a good case might be made for
tive observations, together with my find- a special affinity of Bulininae and the
ings of a total lack of punctation in Plan- palearctic Planorbarius. A comparison
orbina and Gyraulus of the Planorbinae of illustrations of the genital anatomy of
gives a strong implication that punctae Bulinus (Physopsis) and of Planorbarius
are lacking in most planorbids. There in F. C. Baker's monograph (1945) re-
are, however, reasons for assuming that veals obvious gross similarities, especi-
conchological work has not been more ally in the female system, notwithstanding
thorough in respect to these snails than the evident differences in the penial com-
it has been for Bulininae, as shown above. plex (also see Demian, 1960, on Bulinus
-
truncatus Plate VI, figs. 22, 12)? Ob- punctation should be subsequently found in
servations in more detail of my own con- the American " helisomatines".
firm that the resemblance is even greater A relationship of F . .
Baker's Segmen-
than might be inferred from these illus- tininae with Bulininae, on the basis of his
trations. Such a relationship would per- anatomical data and on the data of Paraense
haps have some bearing on the fact that and Deslandes (19 56a,b, among others) ap-
a snail considered to be a Planorbarius pears unlikely. Planorbina (Australorbis
(e.g. P. "dufoiirii " of the Iberian penin- and Biomphalaria) and Gyraiilus of the
sula) is the only non-bulinine species of Planorbinae, on published anatomical
planorbid that, like Bulinus, has been grounds, which I can confirm from per-
demonstrated to be a vector of Schisto- sonal observation, appear to be phylo-
soma haematobium A bulinine relation-
.
genetically distant from Bulininae (see es-
ship might hold also for the near tic pecially illustrations of the genital anatomy
Helisoma which seems to be closely al- of ^'Taphius glabratus" and T. liebmanni "
lied to Planorbarius on anatomical and in Paraense (1958), oí "Planorbis " sopé-
conchological grounds that F. C. Baker eles in Ranson (1953) -^^

and of Gyraulus
has advanced for including both of the in F. Baker, 1945). On the limited
genera in his subfamily Helisomatinae. current evidence given here these dis-
There is reason in turn to suspect some coidal members of the Planorbinae are
particular relationship of the Heliso- nonpunctate; this raises the question as
matinae with the neotropical Taphius to whether punctation is present or lack-
andecolus (d'Orbigny), on comparison of ing in the "physoid" ("bulinoid") species
the illustrations of the genital anatomy of of Planorbidae, placed by Hubendick (1955)
T. atidecolus in Paraense and Deslandes under Planorbinae. Future investigation
(1957) and Paraense (1958) with those of of this feature may provide evidence which
comparable organs (again excepting the could either weaken, or lend support to,
penial complex) of Helisomatinae in Baker that author's position in the matter.
(1945). Also, reference to the illustra- As for a possible genetic connection of
tions of Bulinus anatomy in Demian(1960) Planorbidae (or divisions of the family
should clarify this point. One may sup- that are characterized by punctation) with
pose therefore that Taphius might be other particular Basommatophora, such a
punctate, and in fact this might be taken connection may be indicated in the case
as demonstrated already, for according to of the punctate ?ancylid Burnupia. Man-
Hubendick (1955) and Paraense(1958), Pla- dahl-Barth's brief description (1954) of
tytaphius heteropleurus a species which
,
the prostate and penial complex of this
Baker (1945) describes as finely punctate freshwater African limpet would apply re
on the embryonic whorl, is but a synonym markably well to the known Bulinus spe-
of Taphius. These implied relationships cies. The recent reports of an ancylid as
would receive further support if embryonic a vector of a haematobium -like schisto-
some in India (Gadgil and Shah, 1955)
9lnmy opinion various workers have commonly might be considered in relation to this
overemphasized usage of characters of the type of data. If the primitive basom-
terminal male genitalia (penial complex) in con- matophoran EUobiidae, as their possession
nection with systematics of Planorbidae and of apical punctation might suggest, are
other Basommatophora, while disregarding the
not especially related to apically punctate
female genital structures; one may suppose
that evolutionary changes in the latter are likely Planorbidae, then perhaps they provide
to be of a more fundamental, conservative na-
10 This author places these 2 species of Plan-
ture, and that they therefore would provide the
orbina, which have been previously assigned
more weighty evidence in respect to broader
to Australorbis, in the synonomy of Taphius.
relationships. Attention is drawn here to the
11 Ranson by preference uses the old name
major glandular developments of the "uterus"
and "oviduct" of the female system, and to the Planorbis for the African species now usually
prostate as well. placed under Biomphalaria.
130 H. J. WALTER
evidence that punctation might be a prim- ways should be applied in connection with
itive or even perhaps ancestral character morphological data, and that a refined bio-
in Basommatophora. The possession of systematics of Basommatophora will de-
the character in Planorbidae might then pend on further increases and refinements
be taken as a mark of primitiveness of of our knowledge of, among other things,
Bulinus and its relatives. It would how- conchological morphology such as micro-
ever, be possible to make a case for rel- sculpture. Previous classifications of the
ative primitiveness of the nonpunctate species of Bulinus and other Planorbidae,
Planorbina on known anatomical grounds which have been treated with some urgency
(in this connection see Hubendick, 1955, in recent years because of their impor-
p. 533). One might relate this question tance to medical and public health matters
to my demonstration in this investigation as vectors of Schistosoma blood nukes,
of a total lack of punctae in some Physi- have been attempted on insufficient data;
dae and Lymnaeidae; at any rate the char- a revision of the species is needed, in
acter of nonpunctation should be pertinent which microsculpture of the shell may
toHubendick's(1947) argument that among prove to be of taxonomic importance.
the "higher limnic Basommatophora" (in-
cluding Planorbidae and Ancylidae) the
Lymnaeidae show primitive anatomical
ACKNOWLEDGEMENTS
characters, and that the opposite case By contributing most of the material of
might hold for Physidae. The apical non-Liberian Bulininae that is described
microsculpture of shells of Ancylidae has in this paper, the late Prof. P. L. LeRoux
been given sufficiently close attention of the London School of Hygiene and Trop-
(Walker, 1923) so as to leave no ques- ical Medicine, London, England, made this
tion that most ancylids are nonpunctate, investigation possible with warmth
; I recall
with which my limited data on Ferrissia the courtesy, enthusiasm, and spirit of co-
agrees. Thus it appears that most Ba- operation which he showed in helping me
sommatophora, including those of "higher choose samples from the very large and
limnic" categories, are nonpunctate. The valuable material of African Bulininae that
negative evidence in the literature cited he had amassed through conscientious and
here might be taken as evidence that the sustained effort in the field and labora-
Lymnaeidae are nonpunctate, in line with tory. For contributions of further speci-
my findings. As a speculation on a prac- mens my thanks go to Dr. Elmer Berry
tical matter it may be considered that if of the Natural Institutes of Health, Wash-
the Physidae are indeed entirely devoid ington, D.C., U.S.A. to Mr. W. F. J. Mc-
;
of punctation, and should it prove that Clelland of the East African Medical Sur-
the conchologically"physoid-bulinoid Plan- vey at Mwanza, Tanganyika; to Dr. F.
orbinae" of the Indo -Pacific region are Wickremasinghe of the Kintampo Medical
punctate, a means would be provided for Field Unit in Ghana; and to Prof. Henry
distinguishing between all conchological van der Schalle of the Museum of Zoology,
material of Planorbidae and Physidae that University of Michigan, U.S.A. Prof, van
yet (see Hubendick, 1948, p. 60) may be der Schalle deserves further grateful ack-
catalogued arbitrarily as to family in knowledgement for his support and aid,
museums. which included making arrangements for
The questions about relationships my use of the excellent facilities for mol-
among Basommatophora that are raised luscan study at the University of Michigan.
here may be understood when we have For aid in collecting in the field and in
advanced further in the basic knowledge rearing of snails in the laboratory in
on which systematics depends. It maybe Liberia, I wish to thank Dr. D. M. Levine
pointed out here that the so-called "more of the Liberian Institute for Tropical
biological approaches" to systematics al- Medicine, and my thanks also go to my
Liberian assistant of that Institute, Mr. BulL BioL, France et Belquique, 73 (1-2):
S. Vonleh and Mr. J. Gibson, for similar 19-213.
, 1939b, Remarques sur l'appareil
aid.
génital de l'Indoplanorbis exustus; affinités
I am
indebted to Mrs. Anne Gismann de cette espèce avec les Bulinidés. Bull.
and to Dr. J. B. Burch, of the Museum Soc. Soc. ZooL France, 64(5) :286-295.
of Zoology, University of Michigan, for MANDAHL- BARTH, G., 1954, The freshwater
much aid given with sustained interest, mollusks of Uganda and adjacent territories.
Ann. Kon. Mus. Belg.-Congo, Tervuren,in8°
in the preparation of this publication.
ZooL Wetensch., 32:1-206.
1958, Intermediate hosts of Sc/îisto-
,
LITERATURE CITED soma. African Biomphalaria and Bulinus.

Wld Hlth Org. Monogr, Ser, No. 37, Geneva,
1-89, pis. 1-40, reprinted from BulL Wld
AMBERSON, J. M, and SCHWARZ, E., 1953, On Hlth Org. 1957, 16:1103-63 and 17:1-65.
African Schistosomiasis. Trans, Rov. Soc. 1960, Intermediate hosts of ScAîsio-
,
Trop. Med. and Hyg,, 47 (6): 451-502, soma. in Africa. Some recent information.
BAKER, F, C, 1945, The molluscan family Bull, Wld Hlth Org. 22:565-73.
Planorbidae, Univ. 111. Press, Urbana, p.l- PARAENSE, W. L., 1958, The genera "Aus-
530. tralorbis^\ "Tropicorbis^, "Biomphalaria**,
BURCH, J. ., 1961, Some cytological aspects of "Platytaphius**, and "Taphius** (Pulmonata,
Acroloxus lacustris (Linnaeus), with a dis- Planorbidae). Rev. BrasiL Biol.,18(l):65-80.
cussion of systematics in fresh water lim- PARAENSE, W. L, and DESLANDES, N.. 1956a,
pets. Amer. Malacol. Union Ann. Reports The Brazilian species of "Drepanotrema**.
1961, Bull. 28:15-16. I, "D. anatinum** (Orbigny, 1835). Rev,
, 1962, Cyto-taxonomic studies of BrasiL BioL, 16(4):491-499,
freshwater limpets (Gastropoda: Basom- ,
1956b, The Brazilian species of
matophora). I. The European lake limpet, '^^rëpânotrema*' II, "D. melleum" (Lutz,
.
Acroloxus lacustris. Malacologia, 1(1): 55-72. 1918). Rev, Brasil BioL, 16(4):527-534.
CONNOLLY, M., 1939, A monographic survey of ,1957, A redescription of "TaM¿"S
South African non-marine Mollusca. Ann. S. andecolus" (Orbigny, 1835) (Pulmonata, Plan-
Afr. Mus., 33(l):l-660, pis. 1-19. orbidae), Rev, BrasiL BioL, 17(2):235-243,
DEMIAN, E. S., 1960, Morphological studies of , 1958, The Brazilian species of
the Planorbidae of Egypt. I. On the macro- "Drepanotrema**. V, "D. nordestense**
scopic anatomy of Bulinus (Bulinus) trun- (Lucena, 1953), Rev. BrasiL BioL, 18(3):
catus (Audouin). Ain Shams Sei. Bull. No. 5 275-281.
(Monograph): 1-84, pis. 1-9. PILSBRY, H. A. and BEQUAERT, J., 1927, The
GADGIL, R. K, and SHAH, S.N., 1955, Human aquatic mollusks of the Belgian Congo, with
Schistosomiasis in India, Part II. Infection of a geographical and ecological account of
snails with Schistosoma haematobium. Ind. Congo malacology. Bull. Amer. Mus. Nat.
J, Med. Res., 43:695. Hist. 53(2):69-602.
HARRY, H. W., 1951, Growth changes in the HANSON, G., 1953, Observations sur les Planor-
shell of Pythia scarabaeus (Linné), Proc. bidae Africains. BulL Soc. Path. éxot.,46(5):
Calif. ZooL Club, 2(2):7-14. 783-810.
HUBENDICK, ., 1947, Phylogenetic relations RANSON, G. et CHERBONNIER, G., 1952, Note
between the higher limnic Basommatophora. sur trois planorbes Africains: Planorbis
Zool. Bidr. Uppsala, 25:141-164. pfeifferi Krauss, Planorbis adowensis Bour-
1948, Studies on Bulinus. Ark. 1
,
guignat, Planorbis ruppellii Dunker. Bull.
ZooL, 40(16):l-63, pis. 1-2. Mus. 2e ser., 24(2):206-212.
, 1955, Phylogeny in the Planorbidae.
RAO, H. SRINIVASA, 1923, On the anatomy of
Trans. Zool. Soc. London, 28(6):453-542. Indoplanorbis exustus (Mollusca Pulmonata).
IREDALE, T,, 1911, On the value of the gastro- Rec. Ind. Mus. 25:199-219.
pod apex in classifications. Proc. Malac. WALKER, ., 1923, The Ancylidae of South
Soc. London, 9:319-323. Africa. 82 p. London,
LARAMBERGUE, M. de, 1939a, Étude de l'auto- WRIGHT, A., 1956, The anatomy of the six
fécondation chez les gastéropodes pulmones; species of the molluscan genus Bulinus
recherches sur l'aphallie et la fécondation (Planorbidae) from Senegambia, Proc. Malac,
chez Bulinus (Isidora) contortus Michaud. Soc, London, 32(3):88-104.
132 H. J. WALTER
ZUSAMMENFASSUNG
PUNKTIERUNG DER EMBRYONALEN SCHALE IN DEN
BULININAE (PLANORBIDAE) UND EINIGEN ANDEREN
BASOMMATOPHOREN UND DEREN MÖGLICHE
TAXONOMISCH-PHYLOGENE TISCHE
BEDEUTUNG.
Das Vorkommen einer mikroskopischen Punktierung des embryonalen Gehäuse-

umganges wird hier zum ersten Male als beständiges Charakteristikum der Gattung
Bulinus s.l. in ihrer gesamten geographischen Ausbreitung angegeben, sowie auch
für Indoplanorbis die einzige andere zu den Bulininae gehörige Gattung, gemeldet.
,
Es wurden die Gehäuse von über IbQQ Bulinus s.l. aus 106 verschiedenen Labo-
ratoriums- order Freilandskolonien untersucht, insbesondere solche liberischer
Herkunft, aber auch viele aus über die Länge Afrikas verstreuten Fundorten, sowie
einige aus Sardinien und Irak. Das Untersuchungsmaterial umfasste zumindest 14
verschiedene Arten, Unterarten oder Varietäten der Untergattungen BmZî'wms s.S.,
Pyrgophysa und Physopsis. Punktierung konnte durchwegs nachgewiesen werden,
ausser in seltenen Fällen wo der Nachweis durch Beschädigung der Schale oder son-
stige imgünstige Umstände verhindert wurde. Die^^punctae" sind scharf umrissene
Grübchen die ständig in ganz bestimmter, für jede der 3 subgenerischen Gruppen
etwas verschiedener Anordnung auftreten. Die grundlegenden Beobachtungen über die
Natur und Anordnung dieser punctae sowieüber andere Elemente der Feinstskulptur
wurden an zahlreichen embryonalen Gehäusen aus Laboratoriums kolonien gemacht,
wobei auch die Beständigkeit der Pimktierung in allen 3 subgenerischen Gruppen
festgestellt wurde.
Apikale Punktierung des Gehäuses ist zwar schon im Schrifttum für jede dieser
Untergattungen angeführt worden, jedoch nur für sehr wenige Arten und im all-
gemeinen in so wenig aufschlussreicher Weise dass man sogar bei diesen auf ihre
Abwesenheit in einigen Individuen oder Populationen schliessen könnte. Auch ge-
stattet die gewöhnlich wenig genaue Ausdrucksweise, wie z.B. Beschreibungen von
"TupfChen", "Knötchen", "Perforierungen" oder " Einprägimgen" vereint mit den
,
nur spärlichen Angaben früherer konchologischer Beschreibungen, den Schluss, dass

in Bezug auf Feinskulptur des Gehäuses, die Bulininae noch ungenügend bekannt sind,
was auch für die Basommatophoren im allgemeinen zutrifft. Zweifelsohne wurde
diese Punktierung deshalb so oft übersehen weil sie nur in verhältnismässig un-
beschädigten, sehr gut gereinigten Schalen bei angemessener Vergrösserung und
kritischer Beleuchtung erkennbar ist. Ihr erfolgreicher Nachweis beruht haupt-
sächlich auf des Verfassers Verfahren die Gehäuse mit unterchlorigsaurem Natrium
zu reinigen, wodurch Beobachtung der Einzelheiten des skulpturellen Reliefs bei
durchfallendem Licht ermöglicht wurde.
Unter den nicht bulininen Planorbiden ist embryonale Punktierung bei der
Segmentina Gruppe und bei Platytaphius (? Taphius) bekannt und, innerhalb der an-
deren Basommatophoren, bei der "ancyliden" afrikanischen Napfschnecke Bumupia
und bei den Ellobiiden Melampus, Phytic, und Pythia. Eigenen Untersuchungen an
einigen Arten nicht bulininer Planorbiden zunach, war sie ebenfalls in Planorbarius
corneus vorhanden, fehlte jedoch völlig in Planorbina (Australorbis und Biompha-
laria) und in einer untersuchten Gyraulus Art. Ihr Nichtvorhandensein wurde eben-
falls in verschiedenen Lymnaeiden, in Physa und Ferrissia festgestellt.
Bei den Basommatophoren wurde bisher apikale Mlkroskulptur nur in der Fa-
milie Ancylidae als taxonomisch wertvolles Kriterium erkannt. Ihrer möglichen
taxonomisch-phylogenetlschen Bedeutung Inder Bestimmung von den innerhalb dieser
Unterordnung zwischen verschiedenen Rangstufen herrschenden Beziehungen wurde
hingegen nur sehr wenig Aufmerksamkeit geschenkt.
Hier werden nun einige Fragen über die möglichen phylogenetischen Zusam-
menhänge in Bezug auf Punktierung innerhalb der Unterordnung aufgeworfen. Was
Beziehungen zwischen einzelnen Planorbidengruppen zueinander anbelangt, so wird
darauf hingewiesen, dass den Folgerungen des Autors nach, auf grund einiger anato-
mischer Merkmale, ein gewisses Verwandtschaftsverhältnis zwischen den Bulininae,
dem paläarktischenPZanorbarzMS, dem nearktischen HeZî'sowa und dem neotropischen
Taphius andecolus zu bestehen scheint. Es wäre daher Punktierung noch in
Helisoma, der einzigen dieser Arten bei der sie unbekannt ist, zu suchen. Dagegen
erscheint eine engere Beziehung Zwischen den Bulininae und " Segmentininae" ,
welche beide punktiert sind, auf grund bekannter anatomischer Unterschiede un-
wahrscheinlich.
Ferner wird ein näherer Verwandtschaftsgrad zwischen den Bulininae und der
punktierten ?ancyliden Burnupia durch veröffentlichte anatomische Befunde na-
hegelegt. Schliesslich könnte das Vorhandensein einer Punktierung in den mut-
masslich primitiven EUobiiden - abgesehen davon, dass sie auf eine mögliche Ver-
wandtschaft mit den Planorbiden hinweist- auch vielleicht als Beleg dafür angesehen
werden, dass apikale Punktierung einursprüngliches Merkmal darstellt, obwohl auch
eine gegenteilige Beweisführung geltend gemacht werden könnte.
Durchsicht des Schrifttums führt zu dem Schlüsse, dass die bisherigen die
Feinskulptur der Schale betreffenden Untersuchungen und Beschreibungen nicht
genügend genau sind um hinreichende Rückschlüsseauf die verwickelte Biosystema-
tik innerhalb solcher Basommatophorengruppen, wie z.B. Bulinus sie darstellt, zu
gestatten.
RESUME
PONCTATION DE LA COQUILLE EMBRYONNAIRE DANS
LES BULININAE (PLANORBIDAE) ET QUELQUES AUTRES
BASOMMATOPHORES AINSI QUE SA SIGNIFICATION
TAXONOMIQUE ET PHY LOGE NE TIQUE POSSIBLE
Une ponctation microscopique de la coquille embryonnaire est ici décrite pour
la première fois comme caractère constant du genre Bulinus s.l. dans toute sa dis-
tribution géographique et se trouvant aussi dans V Indoplanorbis le seul autre genre
,
reconnu dans les bulininés.

L'auteur étudia plus de 1500 coquilles de Bulinus s.l. appartenant à au moins 14
espèces, sous-espèces ou variétés des sous-genres Bulinus s.S., Pyrgophysa et
Physopsis et provenant de 106 différentes populations soit de laboratoire soit
d'ambiance naturelle, originaires tout particulièrement de la liberie, mais aussi
de nombreuses autres localités africaines disséminées sur la longueur du continent
africain, ainsi que de la Sardelgne et de l'Iraq. La présence d'une ponctation fut
observée dans tous les spécimens étudiés, excepté quand ils étaient endommagés ou
quand certains autres circonstances en entravaient la détermination. Nous entendons
par "ponctation" un système de "/)M«ctee" ou fossettes très bien définies, arrangées
en un dessin constant dominant la microsculpture et ne différant que de peu dans
,
chacun des 3 sous-genres de Bulinus. L étude d'un grand nombre de bulinins em-
bryonnaires cultivés au laboratoire fournit l'information fondamentale au sujet de
la nature et l'arrangement de cette ponctation, ainsi que des éléments autres de la
microsculpture et en démontra la constance dans les 3 groupes sub-génériques. La
présence d'une ponctation apicale de la coquille a déjà été relevé au préalable dans
la littérature pour chacun de ces sous-genres, mais elle ne se trouve indiquée que
pour un nombre très restreint d'espèces et ce de manière à faire croire qu'elle
pourrait manquer même dans celles-ci en certains individus ou populations. D'autre
part Tusage d'habitude peu exact de termes descriptifs tels que "points", "nodules",
"ponctures" ou "empreintes", joint aux bien maigres indications se trouvant dans
les descriptions conchyliologiques antérieures, démontre, qu'en ce qui concerne la
microsculpture, les espèces bulininés sont encore peu connues, ce qui d'ailleurs
s'applique également aux basommatophores en général. Le fait que si fréquemment
cette ponctation n'a pas été remarquée est certainement dû à ce qu'elle n'est visible
que dans des spécimens relativement peu endommagés, très propres, examinés à la
juste magnlflcatlon et illuminés critiquement. Le succès obtenu à la démontrer dé-
pend largement de la tecnique ici employée à nettoyer les coquilles à hypochlorite
de soude, ce qui permet l'étude des détails du relief sculpturel en transparence.
Parmi les planorbidés non-bulinins, la ponctation embryonnaire était déjà
connue dans legroupede "5^^" etdans le Platytaphius (7=Taphius) et, parmi
les autres basommatophores, dans l'"ancylide" africain Bumupia et les ellobiides
Melampus, Phytia et Pythia. Ici, d'après examen de quelques espèces planorbidés
non- bulininés, la présence d'une ponctation nucléaire de la coquille est rapportée
134 H. J. WALTER
pour lePlanorbarius comeus, ainsi que son absence totale en Planorbina {Austra-
lorbis Biomphalaria ) et dans une espèce de Gyraulus. Elle était également
et
absente dans un nombre d'espèces lymnéides, dans Physa et Ferrissia.
Parmi les basommatophores, la microsculpture apicale a été reconnue avoir
und valeur taxonomique au sein des ancylides; mais, pour le reste, son importance
taxonomique et phylogénétique n'a reçu que l'attention la plus restreinte. Quelques
questions se posentmaintenant au sujet des connections phylogénétique s possibles en
rapport à la ponctation. En ce qui regarde les groupements à l'intérieur des plan-
orbidés, il existe, à l'avis de l'auteur, en vue de caractères connus, une certaine
serait à rechercher /,
parenté entre les bulinlnés, le Planorbarius palear etique, le Helisoma néartique
et le Taphius andecolus néotropical. En conséquence la présence d'une ponctation
,
le seul de ces genres d'où on l'ignore. Par contre,

un proche rapport entre les Bulininae etles "Segmentininae," tous les deux pourvus
de ponctation, semble improbable en vue de leurs différences anatomiques connues.
En ce qui concerne les relations entre bulininés et groupes basommatophores non-
planorbidés, une affinité particulière avec l'{?) ancylide Burnupia est suggérée par
les données anatomiques publiées.
Quant au caractère même de ponctation, sa présence dans les ellobidés sup-
posément primitifs - apart indice qu'elle pourrait fournir pour une relation
possible avec les planorbidés - pourrait être interprêtée comme indice de sa pri-
mitivité, quoiqu'une argumentation contraire soit possible.
Finalement nous concluons de étude de la littérature que ni l'examen ni la
description de la microsculpture du test n' ont encore été exécuté assez rigour-
eusement pour être appliqués adéquatement à la biosystématique complexe prévalant
parmi les groupes basommatophores tels que Bulinus.
RESEÑA
LA PUNTUACIÓN DE LA CONCHILLA EMBRIONAL EN BULININAE
(PLANORBIDAE) Y ALGUNOS OTROS BASOMMATOPHORA Y SUS
POSIBLES IMPLICACIONES TOXONOMICAS Y FILOGENETICAS
La puntuación microscópica de embrional es indicada por primera

la conchilla
vez como una característica constante de Bulinus s.l. sobre su área geográfica, y
también existiendo en el solo otro género reconocido del Bulininae, Indoplanorbis .
El autor estudió más de 1,500 individuos conchxWdiS áe Bulinus s.l, de por lo

nienos 14 especies, subespecies, o variedades de los subgéneros Bulinus s.S.,
Pyrgophysa y Physopsis, representando 107 poblaciones diferentes de laboratorio y
en sus habitats naturales, particularmente de Liberia, así también de Sardinia e
Iraq y de otras localidades numerosas a través de Africa. La puntuación fué
observada en cada una de las especies menos en aquellas en que las conchillas
estaban destrisuidas o anuladas por otros factores. Los puntos "punctae'' son hoyos
bien difinidos que ocurren en una estructura básica que domina la microéscultura
embrionaria, modificados ligeramente en los 3 grupos. El estudio de un gran número
de conchillas embrionarias, de poblaciones criadas en laboratorio dio información
básica sobre la naturaleza y distribución de los puntos y de otros elementos y
confirma la constancia de puntuación en cada uno de los 3 grupos subgenéricos
de Bulinus. La puntuación apical de las conchillas de cada uno de estos subgéneros
se ha reportado por otros investigadores, pero solamente para algunas especies,
la literatura previa sugiere que hasta en estas especies algunos de los individuos
o poblaciones dejan de desarrollar esta característica. El uso commun pero in-
exacto de los términos como "manchas, módulos" y datos insuficientes en previas
descripciones, demuestran que las especies de Bulininae, así como todos los Basom-
matophora en general son muy mal conocidas con respecto a la microéscultura. La
puntuación debió de haberse pasado por trascurrido porque se puede indentificar
solamente en conchillas que estén limpias e intactas con aumento e ílluminación
apropiados. El éxito en demostrar la puntuación dependió mayormente en la técnica
usada por el autor al limpiar las conchillas con hipoclórito de sodio, el cual per-
mitió el estudio del detalle escultural por medio de luz transmitida.
Entre Planorbidae no-bulininae la puntuación embrional se sabía había ocurrido

en el "grupo Segmentina" y en Platytaphius {?=Taphius), y entre otros Basom-
matophora en el "ancylid" Bumupia de Africa; y en los ellobidos Melampus,
Phytia, y Pythia. Examinando varias especies de planorbidos no-bulininae el autor
reporta la existencia de puntuación nuclear en Planorbarius corneus, y totalmente
ausencia en Planorbina {Australorbis y Biomphalaria) y enlas especies de Gyraulus.
La misma ausencia de la puntuación se notó en varios Lymnaeidae, en Physa y en
Ferrissia.
Entre los Basommatophora, la microéscultura apical ha sido aceptada con valor
taxonómico entre los Ancylidae, pero de otro modo, su posible significación
taxonómica-filogenética en relación con otros niveles de taxonomía entre el sub-
orden, se le ha dado hasta ahora muy poca atención. El autor, aquí, da lugar a
prequntas sobre laposibilidadfilogenética entre el suborden en relación-grupal entre
los Planorbidae, se considera que los Bulininae, los paleárticos Planorbarius , los
neártlcos Helisoma y los neotrópicos Taphius an<¿ecoZMS parecen estar relacionados
de alguna manera. Por consiquiente, la puntuación en Helisoma se pudiera explorar,
ya que tadavía no ha sido observada en ella. Una relación cercana entre Bulininae
" Segmentininae", ambas puntuadas, parece improbable por lasdesimejanzas ana-
y
tómicas conacidas.
Una afinidad especial entre los Bulininae y la punteada ?ancylid Bumupia se
sugiere por medios de datos anatómicos y publicados. La existencia de puntuación
en el Ellobiidae, a parte de indicar su relación posible con los Planorbidae, se
puede tomar como evidencia que las puntuaciones apicales representan un carácter
primitivo, aunque se pudiera discutir lo contrario.
De la literatura se concluye que las investigaciones y descripciones pasadas de
la microéscultura han sido insuficientemente exactas para hacer aplicaciones ade-
cuadas en algo tan complejo como la biosystemática dentro de los grupos Basom-
matophora tan como Bulinus,
0^100 BULININAE (PLANORBIDAE)
,
Indoplanorbis
, ,
-
Bulinus
, ""
1
.
,500
s.l.
Bulininae ,
, -
, ,
14
,
Bulinus s.S., Pyrgophysa Physopsis;
.
-
, -
106
.
.
136
,
. >1
-,. H. J. WALTER
"" ,"" "",-

, -
,-
--
(pmictae)
"" "", , ., ,
, ,--
.
.
,, ,-
"
", ""
111
Platytaphius
Planorbidae,
( ?
Na cl
Taphius),
,
Melampus, Phytia Pythia. -
,
Planorbarius corneus Planorbina
(Australorbis Biomphalaria) Gyrauliis,
Lyra-
, -
-
naeldae,y Physa Ferrissia.
. , . Ancylidae,
bidae,
,
andecolus
, --
,-
.
Planor-
Taphius
,.-
Bulinlnae
Segmentininae qQq
, ,
.
^
,,
, ., -
-
- Bulininae
.
?
,
--
,, Bulinus.
2
11, 1962, 1(1): 139-161
DISTRIBUTION OF SPHAERHDAE (PELECYPODA) IN MICHIGAN, U.S.A.

"^
William H. Heard
Museum of Zoology, Ann Arbor, Michigan, U.S.A.
ABSTRACT
About half of the 31 species (and forms) of sphaeriid clams now inhabiting
Michigan are of a more or less general distribution in the state, while the re-
mainder have a geomorphologically or ecologically restricted range. During the
Pleistocene Epoch those sphaeriids present in the basin of the Mississippi River
presumably colonized the Michigan region from the south, as did the unionid
mussels, by upstream migration through the post-glacial streams that drained the
enormous water bodies occupying the region of the present Great Lakes, The
major routes of migration from the Mississippi drainage were: (1) into Michigan's
Upper Peninsula, the Fox River Valley in eastern Wisconsin and (2) into the Lower
Peninsula, the niinois-Des Plaines channel, which drained glacial Lake Chicago (the
southern basin of the present Lake Michigan) and the glacial Maumee River which
drained glacial Lake Maumee (the basin of the present Lake Erie), The subsequent
formation of the present Great Lakes, with new watersheds and an eastward drain-
age, interrupted the former confluences and created a discontinuous distribution
by isolating some species and preventing the progress of others at different times
and with varying effectiveness. A striking example of such an obstacle was the
glacial Grand River whose course transsected the area of the Lower Peninsula;
after the southern part of that Peninsula had been repopulated with sphaeriids, it
effectively blocked the northward spread of three species: Pisidium cruciatum,
P. punctiferum and Sphaerium transversum. likewise, this stream formed the
southern boundary for P. insigne, which did not enter the Peninsula directly from
the south, but from the north by more devious routes. The glacial Grand River later
divided into two streams running in opposite directions, the present easterly- flow-
ing Saginaw River and the present westerly-flowing Grand River, before P. cruci-
atum and P. punctiferum could enter the Saginaw drainage from the west.
The distribution, restricted largely to the Great Lakes bordering the state,
of P. conventus, P. idahoense and S. nitidum, which are species of deep and cold
waters, can be explained on an ecological basis; that of P. henslowanum, P.
amnicum and S. comeum which are restricted to the Great Lakes and their down-
stream drainage, by their probably only recent importation from Europe. The
immediate causes for the localized occurrence of various other species or forms
are, however, less apparent.
In general it is believed that both active migration during periods of alternate
flooding and low water levels, which ultimately disrupted previous confluences, as
well as passive transportation, partly in these waterways by other aquatic animals
such as crayfishes, frogs and flshes, partly overland by aquatic birds, have ob-
scured the original distribution of many of the sphaeriids in the inland waters of
Michigan. The patterns of original distribution are still clearly evident only for
P. cruciatum, P. punctiferum and S. transversum. while they are partially
,
masked in P.fallax, P. insigne, P. obtusale, P. walkeri and S. fabule.
INTRODUCTION drainage system. These watersheds com-

prise Upper (Northern) Peninsula streams
At present all of the streams of the flowing into (1) Lake Superior and (2) Lake
six principal watersheds in the two pen- Michigan, and Lower (Southern) Peninsula
Ínsulas of the State of Michigan belong to streams draining into (3) Lake Michigan,
the Great Lakes - St. Lawrence River (4) Lake Huron, (5) the St. Clair River,
iThis investigation was supported (in part) by a

research grant, 2E-41, from the Nationallnsti- 2present address: Department of Biological
tute of Allergy and Infectious Diseases, U.S. Sciences, Florida State University, Tallahas-
Public Health Service. see, Florida, U.S.A.
(139)
140 W. H. HEARD
TABLE Major streams in the six water
Lake St. Clair, and the Detroit River, and 1.
sheds of Michigan
(6) Lake Erie. The present account of
the sphaeriids inhabiting Michigan is based WATERSHED MAJOR STREAMS
primarily on the nearly 5000 lots in the
collections in the Museum of Zoology, UPPER PENINSULA
University of Michigan (UMMZ) from the Lake Superior Ontonagon River
Sturgeon River
major streams (Table 1) and their con-
Tahquamenon River
nected lakes in these watersheds.
The extensive list of 73 species and 36 Lake Michigan Menominee River
Ford River
ing 5/)
varieties of iingernsiil {Sphaerium includ-
,
and pill clams (Pisidiimi)

recorded for Michigan by Winslow (1926)
Escanaba River
Manistique River
LOWER PENINSULA
reflects the redundant taxonomy developed Lake Michigan Manistee River
by Victor Sterki (1916). The present re- Muskegon River
port reduces that list to 32 species and Grand River
primarily follows the specific nomencla- Kalamazoo River
St. Joseph River
ture of H. B. Herrington (1962); his as-
sistance with identifications is gratefully Lake Huron Cheboygan River
acknowledged. In addition to good species Thunder Bay River
Au Sable River
several common "forms" are mentioned.
Saginaw River
These are at present considered to be
St.Clair River Black River
ecological, but since they are incom-
Lake St. Clair Belle River
pletely known they are included here in
Detroit River Clinton River
the event that some of them might be River Rouge
raised to subspecific or even specific
Lake Erie Huron River
rank in the future. The generic and sub- River Raisin
generic classification of the Sphaeriidae St. Joseph River of
presented here is that which is currently the Maumee River
accepted by most malacologists. How-
ever, this classification is much in need nephridia united; constant retention of juvenile
characters.
of critical re-evaluation and will be the
subject of a future report. Pisidiiim conventus Clessin. The cir-
cumpolar P. conventus i^^byssorut)! Sterki)
SYSTEMATIC POSITION OF SPECIES occurs in the Great Lakes (Lakes Su-
AND THEIR DISTRIBUTION perior, Michigan, and Ontario; Heard,
1962) and in the deep cold lakes of Isle
Species and Habitats Royale, an island in Lake Superior.
Subfamily Pisidiinae F. Baker, 1927 Pisidiuni criiciatum Sterki (Fig. 2).
Genus Pisidum C. Pfeiffer, 1821 This minute sphaeriid (usually less than
2.0 mm
in length) is known only from the
Only the anal siphon developed, the branchial
lower portions of the Grand River (Ottawa
siphon either rudimentary or represented by a
mantle cleft; shell inequipartite: anterior end and Kent Counties) and from the River
of shell longer than posterior end; beaks occa- Raisin and its tributaries (Washtenaw
sionally terminal. County) . An
explanation of this peculiar
Q distribution attempted in the section
is
Subgenus Neopisidium Odhner, 1921 on Routes of Dispersal.
Complete absence of branchial siphon and of Pisidiiim insigne Gabb (Fig. 3). A rare
posterior gills; dorsal loop or lobe of the sphaeriid found in lakes, ponds, bogs, and
morphology of Pisidium cruciatum Sterki and P. insigne Gabb is incompletely known. How-
ever, preliminary observations suggest that both species are members of Neopisidium , and they
are placed here provisionally.
,
SPHAERIID DISTRIBUTION IN MICHIGAN 141
QUEBEC
Fig. 1. Present drainage pattern in Mchigan and surrounding areas.
Streams, P. insigne is absent from the Subgenus Eupisidium Odhner, 1921

southern drainages of the Lower Penin-
Partially fused mantle slit containing a short
sula.
slit representing the branchial siphon; small
Pisidium punctiferum (Guppy) (Fig. 2).
posterior gills present in addition to larger
The disjunct distribution of this sphaeriid anterior gills; posterior gills with inner lamel-
in Michigan (i.e., Brown Lake, Dickinson lae only (outer lamellae entirely lacking); dorsal
County (Baker, 1922) and the North Branch loop or lobe of the nephridia cleft.
of the Paint River, iron County, in the
Lake Michigan watershed of the Upper Pisidium adamsi Prime. This com-
Peninsula, and the lower Grand River in mon species occurs in the state of Michi-
Ottawa and Kent Counties in the western gan as the typical P. adamsi and as the
part of theLower Peninsula) will be dis- form sargenti Sterki. Both types are
cussed later under Routes of Dispersal. more representative of lakes than streams
142 W. H. HEARD
Pisidium aequilaterale Prime. "... Pisidium ferrugineum Prime. Rarely

reported from Michigan and northward taken in quantity, P. ferrugineum and
and westward, but I have seen no speci- its form medianum Prime are typically
ments from these regions" (Sterki, 1916). inhabitants of standing waters. This spe-
This species is typically found in the cies occurs throughout the state.
northeastern United States (Herrington, Pisidium hensloivanum (Sheppard). Pre-
1962). However, reliable records for sumably introduced from Europe, this
Michigan are lacking. Although I have sphaeriid is restricted to the Great Lakes
examined three different lots of P. aequi- in Michigan and occurs more abundantly
laterale from museum collections label- in the lower lakes. Its presence as far
led Reed's Lake, Grand Rapids, Kent inland as Lake Erie is well documented.
County, Michigan, the validity of this Its extended range has been recorded for
locality recordmust be subject to ques- Lake Michigan (Heard, 1961), and recent
tion because many other Grand Rapids dredging by Dr. Frank F. Hooper, Insti-
citations are quite obviously mislabeled tute for Fisheries Research, University
(some even include marine species) It . of Michigan, has also turned up living
would seem that, in the active exchange animals of P. henslowatium from Saginaw
of samples conducted by the early na- Bay of intervening Lake Huron.
turalists, certain lots were tagged with Pisidium lilljeborgi Clessin. Predomi-
the addresses of the sender and not the nently a lake dweller, this species ranges
true place of origin. widely in Michigan and occurs only in-
Pisidium casertanum (Poli). The most frequently in streams. The typical P.
common and widespread sphaeriid in the lilljeborgi is found in all drainages but
state, this variable species is composed the Lake Michigan watershed of the Upper
of several dozen forms from a wide range Peninsula; the form cristatum Sterki does
lakes, ponds, bogs, swamps,
of habitats: not inhabit the southern streams of the
temporary woods pools, beach pools, and Lower Peninsula.
streams of all sizes. Pisidium milium Held. This species
Pisidium compressum Prime. Also of is uncommon but has a wide range. It
widespread range, this species is com- inhabits lakes and small streams in all
mon in lakes and streams of all sizes. watersheds.
Specimens from streams usually exhibit Pisidiwu nitidum Jenyns. Typically
a pronounced diagonal ridge on the beaks; occupants of lakes, P. nitidum and the
shells from lakes are stunted, have an form paupercidum Sterki occur through-
atypical shape, and the ridge is incon- out Michigan. However, the form contor-
spicuous if it is present at all. Several tiim Sterki is found only in the Muskegon,
forms are represented in the state: P. Saginaw and Rouge drainages of the Lower
compressum arrosum Sterki (streams), Peninsula.
P, c. confer turn Sterki (lakes), P. c. lae- Pisidium obtusale C.Pieiiier. This fre-
vigatum. Sterki (a very common form in- quently globular species is found through-
habiting both lakes and streams), P. c. out the state, inhabiting lakes, ponds, and
pellucidum Sterki (primarily a stream sluggish, protected areas of streams.
form), and P. rostratum Sterki (lakes). The typical P. obtusale, however, is ab-
Pisidium fallax Sterki. P. fallax oc- sent in the Northern Peninsula where it
curs throughout the Lower Peninsula but is replaced by the forms rotwidatum
is found only in the Lake Michigan water- Prime and ventricosum Prime, the for-
shed of the Upper Peninsula. More com- mer occuring only in the Lake Michigan
monly found in streams than lakes, this watershed and the latter only in the Lake
species frequently exhibits tubercular Superior watershed.
beaks, a feature in which the beaks appear Pisidium subtnmcatum Malm. A spe-
to have been pushed down, creating a con- cies found in few but widespread locali-
centric pseudo -ridge or bar at their base. ties, P. subtruncatum occurs in all
DISTRroUTION IN MICHIGAN 143
Fig. 2. The distribution of Pisidium (Neopisidium) cruciatum Pisidium (Neopisidium)

punctiferum in Michigan.
144 W. H. HEARD
Flg. 3. The distribution of Pisidium (Pisidium) dubium and Pisidium (Neopisidium) insigne
in Michigan.
SPHAERIID DISTRIBUTION IN MICIDGAN 145
Fig. 4. The distribution of Pisidium (Pisidium) idahoense and Sphaerium transversum in

Michigan.
,
.
146 W. H. HEARD
Michigan watersheds, inhabiting lakes Keweenaw County; Sturgeon River and
and small streams. Douglas Lake, Cheboygan County: Hunt
Pisidiwn variable Prime. This spe- Creek, Ogemaw County; Bass Lake, Li-
cies is commonly encountered in all state vingston County.
watersheds in both lakes and streams.
Subfamily Sphaeriinae F. Baker, 1927
Pisidium walkeri Sterki. Occupying
Genus Sphaerium Scopoli, 1777
lakes and streams throughout the Lower
Peninsula, P. walkeri and its form mai- A distinct anal and branchial siphon present,
nense do not occur in the Upper Penin- either fused only at their base or for the greater
sula. part of their length; shell nearly equipartite,
anterior end of shell shorter than posterior end,
Subgenus Pisidium s.s. C. Pfeiffer
Sphaerium corneum (Linnaeus). An-
Branchial siphon rudimentary (P. dubium) or other sphaeriid introduced from Europe
represented only by a slit in the partially fused and restricted to the lower Great Lakes -
mantle (P.amnicum and P. idahoense); large
St. Lawrence drainage, S. corneum is
posterior gills present in addition to large an-
presently found only in waters outside the
terior gills; posterior gills with inner lamellae
as well as outer lamellae; dorsal loop or lobe boundaries of the state: Lake Erie.
of nephridia cleft, Sphaerium fabale Prime. This species
is very widespread in the streams
of the
Pisidium amnicum (Mnller) . This large Lower Peninsula watersheds but does not
species (length greater than 5 mm) is occur in the Northern Peninsula.
known only from certain waters bordering Sphaerium lacustre (Müller). Although
the state in the east: Lake Erie, the De- the typical S, lacustre ranges throughout
troit River, and Saginaw Bay of Lake all watersheds of Michigan, the form
Huron. Introduced from Europe and at ryckholti (Normand) is not found south of
present common only in the Great Lakes - the Grand- Saginaw Valley, while the form
St. Lawrence River drainage, P. amnicum jayense (Prime) does not occur north of
has advanced into Lake Huron and may it.
eventually extend its range upstream into Sphaerium nitidum Clessin. Typical of
Lake Michigan, as did P. henslowanum deep and cold waters, S, nitidum occurs
and possibly into Lake Superior. in Lake Michigan and Lake Huron, and
Pisidium dubium (Say) (Fig. 3). Typi- the inland lakes of Isle Royale (Lake
cally living in very small colonies, this Superior) in Keweenaw County (Heard,
widespread species rarely found in
is 1961).
lentic habitats. large size (length
Its Sphaerium occidentale (Prime) A char-
.
more than 5 mm) may lead to some con- acteristic part of the fauna of woods
fusion with P. amnicum and P. idahoense pools, S. occidentale ranges throughout
However, the coarse striae of P. dubium Michigan.
are absent from the beaks while remain- Sphaerium partumeium (Say) (Fig, 5).
ing prominent in P. amnicum; the striae This species is very widespread in tem-
in P. idahoense are fine. In addition the porary woods ponds and muddy substrates
beaks are more terminal in P. dubium, of lakes and sluggish streams of Michi-
and the hinge teeth are different from gan.
those of the two other species as de- Sphaerium rhomboideum (Say) (Fig. 6).
scribed in detail by Herrington (1962). Of wide range in all Michigan watersheds,
Pisidium idahoense Roper (Fig. 4). this peculiar species, which has a rhom-
This large species (length greater than boid shape, inhabits muddy areas in lakes
5 mm; see P. dubium) is typical of cold and streams.
and deep waters such as Lake Superior Sphaerium securis (Prime). Widely
and Lake Michigan (Heard, 1962), although ranging throughout the state, S. securis
it occurs as well in suitable ''inland" is found in lakes and ponds with muddy
localities: Isle Royale (Lake Superior), substrate, swamps, and woods pools.
Sphaerium striatinum (Lamarck) This . The sphaeriid species or "forms" with

variable species is the most commonly restricted range can be grouped into the
encountered of all the sphaeria in the following categories:
state. It has many forms which are found (a) Species present in all drainages
in lakes and streams of all sizes. except '(1) the Lake Superior Water-
Sphaerium sulcatum (Lamarck). This shed of the Upper Peninsula in the
is one of the largest of all sphaeriids, very north: P.fallax and P.ob-
sometimes reaching one inch in length. tusale s.S. -wiihiis iormrotwidatiim,
It has a more rectangular shape and more or except (2) the Lake Michigan
consistent striae (in spacing and height) Watershed of that Peninsula: also
than S. striatinum with which it is fre- P. obtusale s.s. and its form ven-
quently associated. Although S. sulcatum tricosum, and P. lilljeborgi.
includes several forms, it is considerably (b) Species of northern ocurrence, ab-
less variable than most sphaeriid clams. sent only from the southern drain-
It occurs commonly in lakes and streams ages of the Lower Peninsula: P.
throughout Michigan. insigne (Fig. 3); also P. lilljeborgi
Sphaerium transversum (Say) (Fig. 4). form cristatuma.nd S lacustre form
.
This sphaeriid occupies streams rather ryckholti.

than the usual habitats of its more closely (c) Species restricted to the Lower Pen-
related fellow species, i.e., S. lacustre, insula: P. walkeri, S.fabale and P.
S. partumeium and S. securis. In Michi- obtusale s.s. which are found gen-
gan S. transversum occurs only in the erally throughout the Lower Penin-
southern streams of the Lower Penin- sula, and P. cruciatum (Fig. 2), S.
sula. transversum (Fig. 4), P. nitidum
form contortum and S. lacustre form
jayense which occur only in the
Distribution Patterns southern drainages of this peninsula,
In summary, a review of the locality south of the Grand-Saginaw Valley.
records available to me reveals that the (d) Species occurring only in water-
Sphaeriids in Michigan fall into two cate- sheds draining into Lake Michigan
gories: species of general distribution from both the Upper and Lower
of which 3 representatives have been Peninsulas: P. punctiferum and its
mapped, and species with a restricted forms (Fig. 2).
range. (e) Species common to the Great Lakes
Sphaeriids with general distribution bordering the State: P. amnicum,
(1. e. found in all watersheds of both
,
P. henslowanum and S. corneum.
peninsulas) are, as follows: Pisidium conventus and S. nitidum.
Pisidium adam s i are also characteristic of the Great
P. casertanum and its forms Lakes and in addition occur in cer-
P. compre ssum tain lakes on Isle Royale in Lake
P. dubium (Fig. 3) Superior. Pisidium idahoense is
P. ferrugineum typical of these habitats and also
P. milium persists in highly localized relict
P. nitidum s.S. and form pauperculum populations in a few inland lakes
P. subtruncatum and streams in the Lower Penin-
P. variabile sula (see Fig. 4).
Sphaerium lacustre s.s. The observed distribution for both spe-
S. occidentale cies and forms can, to a certain extent,
S. partumeium (Fig. 5) be explained by habitat requirements and
S. rhomboideum (Fig. 6) particularly by the post-glacial history of
S. securis the territory and the paths of invasion
S. striatinum that were used by these and other fresh
S. sulcatum water clams, as discussed below.
W. H. HEARD
,
*^-.
SPHAERIUM PARTUMEIUM
Fig. 5. The distribution of Sphaerium partumeimn in Michigan.

SPHAERiro DISTRroUTION IN MICHIGAN 149
Fig. 6. The distribution of Sphaerium rhomboideum in Michigan.

150 W. H. HEARD
ROUTES AND MEANS OF DISPERSAL

General effect of former Drainage
Confluences
Glaciation in the Great Lakes region
not only created a large number of lentic
and lotie habitats, but the subsequent
formation of the present drainage basins
played an instrumental role in the spread
of numerous aquatic mollusks. When the
peculiar distribution patterns of certain
Michigan sphaeriids are considered, one
can find in the patterns a reflection of
the original drainage basins and evidences
of some of the later changes in them.
Fresh water clams were able to migrate ^
from one drainage to another during peri-
ods of glacial confluence. Low water
levels, following the retreat of the gla-
ciers, interrupted many of these drainage
systems so that (1) barriers to further
Fig. 7. Migratory routes of sphaeriid clams into
migration were created and (2) some Michigan, a: the Fox PU ver Valley In
faunal elements were isolated. eastern Wisconsin; b: the Illinois— Des
The relation of the distribution of Plaines drainage of glacial Lake Chicago;
unionids or naiades to the post-glacial c: the Maumee River draining glacial
Lake Maumee. The extent of glacial ice
history of Michigan has been studied by
cover Is indicated by stippling, and the
several investigators. The major avenues present Great Lakes are shown by slant-
of northward migration into this region ing lines.
from the Mississippi River drainages
were outlined by Walker (1898, 1913), The relationship between present day hy-
Ortmann (1924), van der Schalie (1938, drographical patterns in the Great Lake
1945) and Goodrich and van der Schalie area and those of post-glacial times can
(1939). The three major routes that served be seen by comparing Figs. 1 and 7, and,
for the dispersal of mussels were evi- for greater detail in the Fox River Val-
dently also used by the sphaeriid clams. ley, Fig. 8.
Briefly they are (Fig. 7): (a) the Illinois-
Des Plaines outlet of glacial Lake Chi-
cago (the present Lake Michigan), (b) the
Routes of Penetration
Maumee River, draining Lake Maumee The role of the Illinois -Des Plaines
(the present Lake Erie), which served for Lake Chicago and the
outlet of Glacial
the colonization of the Lower Michigan Maumee River draining glacial Lake
Peninsula from the east, and (c) the Fox Maumee The interpretation of the marked
.
River Valley in eastern Wisconsin, which similarity of the Mississippi naiad fauna
served as a path of entry into the Upper with that of the central Great Lakes is
Peninsula. After these pelecypods suc- based on the direct connection, already
cessfully invaded Michigan waters, the referred to, of glacial Lake Chicago (the
confluences between the Mississippi and present Lake Michigan) and glacial Lake
Great Lakes drainages were eventually Maumee (the present Lake Erie) with the
broken and further dispersal of certain Mississippi drainage during the Pleisto-
species was prevented by the formation cene Epoch.
of certain other barriers, such as the The same explanation may be used to
glacial Grand River, discussed below. interpret the distribution of certain
observed among several unionids.

Pisidium cruciatum reached the lower
regions of the Grand River through the
Illinois-Des Plaines drainage, and gained
access to the River Raisin through the
glacial Maumee River. The River Raisin
evidently flowed directly into the Maumee
at that time (van der Schalle, 1938).
Sphaerium trans versum {Fig. 4) oc-
curs in Michigan only in the Lower Pen-
insula south of the glacial Grand River
Valley, but ranges a little farther north
in Wisconsin (Baker, 1928) and much
farther north in Canada (Great Slave Lake;
Herrington, 1950). This species also
reached Michigan through the Illinois-Des
Plaines and Maumee River channels. Its
northward spread in Michigan was blocked
by the glacial Grand River barrier (see
below), and in Wisconsin by the rupture
of the glacial Fox River and subsequent
stream -capture of part of this stream by
the Great Lakes-St. Lawrence drainage
system.
Fig. 8. The present drainage systems in the Fox Pisidimn punctiferum s.s. and P.p.
River Valley in eastern Wisconsin, a: the form armatum Sterki (Fig. 3) are local-
Wisconsin River, still in the Mississippi ized in Michigan in the lower reaches of
River drainage; b: the Fox River, now in the Grand River, a colonization which may
the Great Lakes drainage; c: Lake
again be interpreted as post-glacial in-
Winnebago; d: the Menominee River in
the Upper Peninsula of Michigan. The vasion through the Illinois-Des Plaines
Mississippi River drainage is indicated drainage.
by stippling. Role of the Fox River Valley. During
the late Wisconsin stage of glaciation the
sphaerlids in Michigan, particularly that Mississippi (Wisconsin River) and Great
of the very localized species Pisidimn Lakes drainages (the present Fox River
cruciatum, Sphaerium trans version, as and Green Bay) were also connected
well as of Pisidium punctiferiim s.s. and through the Fox River channel in eastern
its form armatum Sterki, which are re- Wisconsin (Goodrich and van der Schalle,
stricted to the southernmost drainages 1939), and this route was employed by
of the Lower Peninsula and whose fur- many of the species of sphaeriids pres-
ther advance was quite evidently blocked ently inhabiting the Upper Peninsula of
by geographical changes. Michigan. The present drainages of this
Examination of museum specimens and former confluence are shown in Fig. 8.
of the literature (Sterki, 1895, 1916; Her- The sphaeriids of Wisconsin (Baker, 1928;
rington, 1962) reveals that Pisidium cru- Morrison, 1932) and of the Lake Michigan
ciatum presently inhabits streams of the Watershed of the Upper Peninsula are
extensive Mississippi River drainage sys- compared in Table 2. Most sphaeriids
tem and two Michigan streams, the Grand are found throughout the remnant drain-
and Raisin Rivers (Fig. 2), which are ages of the glacial Fox River confluence,
tributaries of the Great Lakes - St. Law- although there are some significant ex-
rence River drainage. This pattern is ceptions.
similar to the discontinuous distribution Baker's (1928) records of P. punctiferum
152 W. H. HEARD
TABLE 2. Sphaeriids of the Fox River Valley in eastern Wisconsin and the Lake Michigan
Watershed of the Upper Peninsula of Michigan. The records from the present Fox River
drainage include those from Lake Winnebago (see Fig. 3)
gan from the north are not accurately TABLE 3. Sphaeriidae of the Grand - Saginaw
known. An extension of the Fox River Valley, Michigan^
Valley route would, however, explain Grand Saginaw

Species
the distribution of Pisidium insigne, P. River River
lilljeborgi form cristatiim and Sphaerium Pisidium
lacustre form ryckholti which do not adamsi Prime X
occur in the southern part of the Lower form sargenti Sterki
Peninsula, These species evidently did casertanum (Poli)
f. roperi Sterki
not penetrate the Lower Peninsula through
compressum Prime
the Illinois -Des Plaines or Maumee cruciatum Sterki
routes, but were presumably able to mi- dubium (Say)
grate through the Fox River Valley, fallax Sterki
colonize the Upper Peninsula, and pass ferrugineum Prime
f. medianum Sterki
southward into the northern area of the
insigne Gabb
Lower Peninsula. lilljeborgi Clessin
milium Held
nitidum Jenyns
f. contortum Prime
f. pauperculum Sterki
Barriers: Effect of the Glacial
obtusale C. Pfeiffer
Grand River rotundatum Prime
f.
ventricosum Prime
f.
During the Pleistocene Epoch, the lower punctiferum (Guppy)

peninsula of Michigan was transsected by f. armatum Sterki
variabile Prime
the glacial Grand River (Bretz, 1953)
walke ri Sterki
which channeled the waters of the Erie f .mainense Sterki
'
and Huron basins through the Grand-Sagi- Sphaerium

naw Valley into glacial Lake Chicago fabale Prime
(Hough, 1958). The present remnants of lacustre (Müller)
this glacial stream are the Grand River, occidentale Prime
partumeium (Say)
flowing westward into Lake Michigan, and
rhomboideum (Say)
the Saginaw River, draining northeastward securis Prime
into Saginaw Bay of Lake Huron. Walker's striatinum (Lamarck)
(1898) Zoogeographie study of the unionid sulcatum (Lamarck)
clams of Michigan revealed that the great transversum (Say)
majority of species are essentially con-
fined to the Grand-Saginaw Valley and to
the streams south of it. In contrast, how-
ever, the Grand-Saginaw Valley was a bar-
rier to only a few sphaeriids (3 species
and 3 forms), as most species (18 spe-
cies, including 6 forms) exhibit a general
range throughout the state. The highly
varied assemblage of sphaeriid clams
occupying the Grand and Saginaw River
drainages is presented in Table 3.
Pisidium cruciatum Sphaerium lacustre
,
îorm jayense and S. transuersum do not

extend north of the Grand-Saginaw drain-
ages. S. lacustre s.S., has a general
range in North America; in Michigan,
however, the form jayense is not found
north of the Grand-Saginaw Valley.
154 W. H. HEARD
Peninsula; only the form cristatum has a migration through confluent drainage pat-
limited range in the Lower Peninsula. terns. Transportation by other animals
From the material at hand, it would has doubtlessly followed and in most cases
seem that several more species and has effectively masked the original dis-
forms inhabit the Grand River than oc- tribution patterns. The original distribu-
cur in the Saginaw River. It is however tion patterns of P. cruciatum, P. puncti-
suspected that further collecting in the forum, S. transversum and S.lacustre
Saginaw Valley might also reveal the form jayense in Michigan are clearly
still
presence of the more widespread species discernible, those of P.fallax, P. insigne,

Pisidium insigne and P. milium; P. crii- P. walkeri, P. lilljeborgi form cristatum,
ciatiim and P. punctiferum, of limited P. obtusale s.s. and its forms rotundatum
range, probably do not occur there. ventrico sum, S.fabale, and S. lacustre
2iná
Should their absence be confirmed, this form ryckholti are evident to a lesser
distribution pattern would indicate that a degree, while passive dispersal has pre-
rupture in a single drainage creates an sumably disguised to varying extents the
effective barrier to dispersal; i.e., that original ranges of the remaining species
glacial Grand River must have given rise and forms.
to the easterly -flowingSaginaw River be-
fore P. cruciatum and P. punctiferum DISCUSSION
were able to populate this area from the
west. It must be understood that not all
species disperse at the same time or at
Means of Dispersal the same rate. The same obstacles may
not exist at ail times and the same ob-
It has often been suggested that various stacle may be overcome by some species
aquatic animals are responsible for the but not by others in a given period of
dispersal of sphaeriid clams. Among time. While Pisidium, obtusale s.S., P.
these are various aquatic insects (Kew, ivalkeri and Sphaerium /abale, for exam-
1893; Fernando, 1954), crayfishes (Kew, ple, presently extend throughout the Lower
1893), fishes (Odhner, 1951), frogs, sala- Peninsula of Michigan, P. cruciatum , P.
manders and aquatic birds (Kew, 1893). punctiferum and S. transversum have not
Published accounts of sphaeriids attached as yet been able to spread north of the
to the exterior of other aquatic and ter- Grand-Saginaw Valley (either by active
restrial animals indicate that the spread migration or through adventitious trans-
of these bivalves is linked to the move- port by other animals), and P. insigne,
ments and dispersal of their transport present in the north, is still absent from
hosts, i.e., largely to the water connec- the region of Michigan south of that Valley.
tions. Odhner (1951) also mentioned the The geographical range of a species is
possibility of endozoic dispersal: unborn also confined by the limitations imposed
juveniles, protected in the gills within the by the ecological tolerances of the ani-
shell of the parent, may occasionally be mals. Thus, dispersal takes place not
able pass through the intestine of a
to only through time but also through an
fish without injury. Such a mode of dis- ecological continuum in space. Pisidium
persal is still dependent on the move- idaJioense is infrequently found south of
ments of the host animal in the water- the North American Great Lakes and is
ways. only rarely found "inland" from the Great
While overland carriage as a means of Lakes in that area. The disjunct inland
dispersal of sphaeriid clams is not alto- localities of P. idahoense in Michigan
gether discounted, it is relegated to a may represent suitable habitats which
secondary role. It seems reasonable to have persisted locally, enabling relict
assume that the distribution of sphaeriids populations to survive.
was originally accomplished by active Pisidium fallax presumably migrated
.
through the glacial Fox River Valley (see fauna cannot yet be adequately correlated
Routes of Dispersal) into the Lake Michi- with that of surrounding territories be-
gan Watershed of the Upper Peninsula cause records for these areas are only
which is easiest of access from the Fox fragmentary.
River, but it was apparently unable to The apparent absence of species and
invade and populate the Lake Superior forms from individual watersheds may be
Watershed, The apparent absence of P. due to the lack of sufficient collecting in
ivalkeri from the entire Upper Peninsula those areas. This is expected to apply
is unexpected, for Baker (1928) reports to P. lilljeborgi s.S., a species presently
this species to inhabit the Fox River unknown in Michigan only from the Lake
drainage and other localities in eastern Michigan Watershed of the Upper Penin-
Wisconsin. Comparing the distribution of sula, yet common throughout eastern Wis-
other pisidia, one would expect that P. consin. It is anticipated that with more
ivalkeri also migrated from the Missis- intensive collecting it will be found in the
sippi River drainage through the Fox Lake Michigan Watershed of the Upper
River Valley into the Upper Peninsula Peninsula because this drainage system
(see Role of Fox River Valley). Further must have been utilized in populating the
collecting in the Lake Michigan Water- Lake Superior Watershed from the Fox
shed, may reveal the presence of the River. Pisidium insigne is widespread
species in that area. in both watersheds of Michigan's Upper
A surprisingly extensive sphaeriid fauna Peninsula but has not been recorded from
is localized in the waters of the Isle Wisconsin at all. It, too, will probably
Royale, and island in northern Lake Su- be discovered in eastern Wisconsin
perior (Walker, 1909; UMMZ specimens) throughout the Wisconsin and Fox drain-
which is much larger than that found in ages of the Fox River Valley migratory
the Lake itself (Heard, 1962): route
Pisidiwn adamsi The peculiar distribution of "forms"
P. casertanum of certain species (see Distribution Pat-
P. conventus terns) is difficult to interpret. It is fre-
P. ferriigineum quently found that the typical species has
P. idahoense a different distribution than its forms as
P. lilljeborgi shown below for P. lilljeborgi and P.
P. milium ob túsale.
P. nitidum Pisidium lilljeborgi s.s. is more com-
P. ob túsale mon in eastern Wisconsin (the Fox River
P. punctiferum? Valley migratory route) and occurs
P. subtnmcatum throughout Michigan's Upper and Lower
P. variabile Peninsulas (being expected in the Lake
Sphaerium nitidum Michigan Watershed of the Upper Penin-
S. securis sula, as previously mentioned). The form
S. sulcatum. P. I. cristatwn is widely distributed in
This assemblage represents an isolated Wisconsin, occurs over the Upper Penin-
segment of the fauna of the ''mainland" of sula, and penetrates only into the northern
Ontario, Canada. The routes by which portions of the Lower Peninsula of Michi-
these sphaeriids colonized Isle Royale gan.
are not accurately known. Presumably The typical P. obtusale has not been
they migrated to the island from western recordedfor either Wisconsin or the Upper
Ontario after having passed northward up Peninsula of Michigan but is common in
the Mississippi River, bypassing the Fox the Lower Peninsula and is present on
River Valley outlet, and around the basin Isle Royale (Lake Superior). Baker (1928)
of the present Lake Superior. reports two widely separate localities for
On the whole, the Michigan sphaeriid the form P. o. rotundatum in Wisconsin,
156 W. H. HEARD
whose hydrographie al connections cannot Aquatic moUusks of the Upper Peninsula of
be determined. In Michigan the form ro- Michigan. Misc. Publ. No. 43, Univ. Mich.
Mus. Zool., 45 p.
timdatimi is found over the Lower Penin-
HEARD, W. H., 1961, Pisidium henslowanum
sula and replaces the typical P. obtusale (Sheppard) in Lake Michigan. Nautilus,
in Lake Michigan Watershed of the
the 74(3):123.
Upper Peninsula. The form P. o. ventri- ,1962, The Sphaeriidae (Mollusca:
cosum is not listed for Wisconsin but Pelecypoda) of the North American Great
Lakes. Amer. Midi. Nat., 67(1):194-198.
occurs throughout the Lower Peninsula
HERRINGTON, H. ., 1950, Sphaeriidae of
and the Lake Superior Watershed of the
Athabaska and Great Slave Lakes in north-
Upper Peninsula of Michigan. The com- western Canada. Canadian Field- Naturalist,
mon Wisconsin form is P. o. vesiculare 64(l):25-32.
which Baker (1928) states to occur in the , 1962,A revision of the Sphaeriidae
eastern part of the State. of North America (Mollusca: Pelecypoda).
Misc. Publ. No. 118, Univ. Mich. Mus. Zool.
The number of so-called "forms" with
p. 1-74, pis. 1-7.
limited range in a distribution pattern HOUGH, J. L., 1958, Geology of the Great
parallel to the geological history of the Lakes. Univ. 111. Press, Urbana, 313 p.
area raises the question of whether these KEW, H. W., 1893, The dispersal of shells.
forms are not really true subspecies (i.e., Kegan Paul, Trench, and Trubner, London,
291 p.
geographical varieties) which arose before
MORRISON, J. P. E,, 1932, A report on the
the present distribution of their parent Mollusca of the Northeastern Wisconsin
species was determined or which have Lake District. Trans. Wise. Acad. Sei.,
appeared the formation of the pres- 27: 359-396.
ent drainage systems and subsequent es- ODHNER, N. H., 1921, On some species of
Pisidium in the Swedish State Museum.
tablishment of a restricted range for the
J. Conchol., 16(7):218-223.
parent species (or are appeariiig non). 1951, The mountain fauna of the
,
Unfortunately, too little is known con- Virihaure Area in Swedish Lapland. Lunds
cerning the overall distribution and gen- Univ. Ârsskrift. N. F. Avd. 2, 46(2):26-50.
eral biology of sphaeriid clams (both ORTMANN, A. E., 1924, Distributional features
of Naiades in the tributaries of Lake Erie.
species and "forms") to permit further
Amer. MidL Nat., 9(3):101-117.
conjecture at the present time. It is STERKI, v., 1895, Two new Pisidia, Nautilus,
hoped, however, that this report will serve 8(9):97-100.
to stimulate other attempts to define the , 1916, A preliminary catalogue of
distribution and Zoogeographie al relation- the North American Sphaeriidae. Ann. Car-
negie Mus., 10(3/4): 429-474.
ships of the Sphaeriidae.
VAN DER SCHAUE, H., 1938, The Naiad fauna
of the Huron River, In southeastern Michigan.
Misc. Publ. No. 40, Univ. Mich. Mus. ZooL,
LITERATURE CITED 83 p.
BAKER, F.C., 1927, On the division of the , 1945, The value of mussel distribu-
Sphaeriidae into two subfamilies; and the tion in tracing stream confluence. Pap. Mich.
description of a new genus of Unionidae, Acad. Sei., Arts and Lett., 30:355-373.
with descriptions of new varieties. Amer. WALKER, ., 1898, The distribution of the
Midi. Nat., 10(7):220-223. Unionidae in Michigan. Privately printed by
1928, The fresh water Mollusca of
, the author, 23 p.
Wisconsin,II. Pelecypoda, Wise. Geol. Nat. , 1909, Annotated list of the Mollusca
Hist. Surv. Bull. 70, 482 p. of Isle Royale, Michigan. In C.C.Adams: An
BAKER, H. ., 1922, The Mollusca ofDicklnson ecological survey of Isle Royale, Lake Su-
County, Michigan, Occ. Pap. No. Ill, Univ. perior. Ann. ilep. Geol. Surv. Mich, for 1908,
Mich. Mus. Zool. , 44 p. Wynkoop, Hallenbeck and Crawford, Lansing,
BRETZ, J. H., 1953, Glacial Grand River, p. 281-298.
Michigan. Pap. Mich. Acad. Sei., Arts and , 1913, The Unione fauna of the Great
Lett., 38:359-382. Lakes. Nautilus, 27(2-5):18-23, 29-34, 40-
FERNANDO, C. H., 1954, The possible dispersal 47, 56-59.
of Pisidium by Corixidae. J. Conchol. ,24(1): WINS LOW, M. L., 1926, A revised check list of
17-19. the Michigan Mollusca. Occ. Pap. No. 181,
GOODRICH, CandH. VAN DER SCHAUE, 1939, Univ. Mich. Mus, ZooL, 28 p.
ZUSAMMENFASSUNG
DIE VERBREITUNG DER SPHAERIIDEN (PELECYPODA) IN MICHIGAN
Ungefähr die Hälfte der 31 gegenwärtig im Staate Mchigan, in den Vereinigten
Staaten von Amerika vorkommenden Arten von sphaeriiden Muscheln mit ihren
"Formen" sind in diesem Staate allgemein verbreitet während die übrigen von
geomorphologisch oder ökologisch beschränkter Verbreitug sind. Während des
Pleistozäns besiedelten vermutlich jene Spheriiden die sich im Mississippibecken
befanden (ebenso wie die Unioniden Muscheln) die Gegend vom Süden her indem sie
stromaufwärts in den post-glazialen Flüssen wanderten welche die ungeheuren Seen
entwässerten die den Raum der heutigen Grossen Seen einnahmen. Die Hauptein-
wanderungswege vom Mississipibecken her waren (1) für Michigans obere oder
nördliche Halbinsel das Tal des Foxflusses im östlichen Wisconsin und (2) für die
untere oder südliche Halbinsel der Illinois - Des Piaines Wasserweg welcher den
glazialen Chicagosee (das heutige südliche Becken des Michigansees) entwässerte
und der eiszeitliche Maumeefluss, welcher den damaligen Maumeesee (das Becken
des jetzigen Eriesees) entleerte. Die darauffolgende Bildung der heutigen Seenkette
mit ihren neuen Wasserscheiden und östlichem Abfluss unterbrach die vorherigen
Konfluenzen und schuf ein diskontinuirliches Verbreitungsbild indem einige Arten
isoliert wurden und der Ausbreitung anderer, zu verschiedenen Zeiten und mit
minderem oder grösserem Erfolge, Grenzen gesetzt wurden. Ein schlagendes Bei-
spiel für ein solches Hindernis stellt der eiszeitliche Grandfluus dar, welcher das
Gebiet der unteren Halbinsel durchquerte. Nachdem der südliche Teil dieser
Halbinsel von Sphaeriiden besiedelt worden war, versperrte dieser das weitere
Vordringen dreier Arten: Pisidium cruciatum P. punctiferum und Sphaerium
,
trans versum nach dem Norden. Gleicherweise bildete er die südliche Grenze für
P. insigne, eine Art welche nicht unmittelbar vom Süden sondern auf Umwegen vom
Norden her in die Halbinsel eingedrungen war. Später teilte sich der Grandfluss in
2 nach entgegengesetzte Seiten fliessende, den nach Osten verlaufenden heutigen
Saginawfluss und den nach Westen strömenden heutigen Grandfluss, und zwar noch
bevor P. cruciatum undP. punctiferum vomWestenher in den Saginaw eingedrungen
waren. Das grösstenteils oder ausschliesslich auf die den Staat umgebenden Grossen
Seen beschränkte Vorkommen vonP. conventus P. idahoense und S. nitidum, welche
,
Bewohner tiefer und kalter Gewässer sind, lässt sich auf ökologischer Grund-
lage erklären; das gleichfalls auf die Grossen Seen sowie deren Abflussgebiet be-
schränkte von P. henslowanum P. amnicum und S. comeum durch die wahrschein-
,
lich erst in der Gegenwart erfolgte Einschleppung aus Europa. Die unmittelbaren
Ursachen für die örtliche Begrenzung verschiedener anderer Arten oder Formen
1st jedoch weniger klar erkenntlich.
Im allgemeinen lässt sich sagen, dass einerseits durch aktive Wanderung in
Zeiten abwechselnder Überschwemmungen und niederer Wasserstände welche
schliesslich die ehemaligen Zusammenhänge zerstörten und andrerseits durch
Verschleppung, teils innerhalb der Wasserwege mittels anderer Wassertiere wie
Krebse, Frösche oder Fische, teils Überlands durch Wasservögel, das ursprüngliche
Verbreitungsbild vieler Sphaeriiden in den Binnengewässern Michigans verdunkelt
wurde. Ein solches ist nur noch bei P. cruciatum P. punctiferum und S. trans-
,
versum klar erkenntlich während es bei P. /aZZax, P. insigne, P. obtusale, P. walkeri

•\iíá S fabale teilweise verschleiert
. ist.
158 W. H. HEARD
RESUME
LA DISTRIBUTION DES SPHAERIIDES (PÉLÉCYPODES) AU MCHIGAN
A peu près la moitié des 31 espèces de bivalves sphaeriides et de leurs

"formes" habitant présentement l'état de Michigan aux États Unis d'Amérique y ont
une distribution généralisée, tandis que la répartition du reste des espèces est
géomorphologiquement ou écologiquement restreinte. Pendant l'époque pleistocene
les sphaeriides présents dans le bassin du Mississippi ont vraisemblablement colonisé
la région du Michigan de par le sud (tout comme les bivalves unionides) en re-
montant le cours des rivières post-glaciales par lesquelles s' évidaient les immenses
pièces d'eau occupant la région des grands lacs présents. Les routes principales
d' immigration du bassin du Mississippi étaient: (1) pour la péninsule supérieure ou
septentrionale, la vallée du Fox, dans le Wisconsin oriental et (2) pour la péninsule
inférieure ou méridionale, la voie Illinois -Des Plaines, par laquelle s'effectuait le
drainage du lac Chicago glacial (correspondant au bassin méridional du présent Lac
Michigan) et le fleuve glacial Maumee par lequel s' écoulaient les eaux du lac Maumee
glacial (le bassin du Lac Erie présent). La formation subséquente des Grands Lacs
actuels, avec les nouvelles lignes de partage des eaux, et leur écoulement vers
l'est, rompit les confluences antérieures et créa une distribution sphaeriide dis-
continue, isolant certaines espèces et entravant plus ou moins efficacement en di-
verses périodes le progrès de certaines autres. La rivière Grand glaciale.traversant
la région de la péninsule inférieure, fournit un exemple frappant d'un tel obstacle:
suivant la colonisation de l'extrême sud de cette péninsule par les sphaeriides, elle
limita effectivement la diffusion vers le nord des espèces Pisidium cruciatum, P.
punctiferum et Sphaeriwn trans ve rsum . De même, cette rivière constitua la limite
sud pour le P. insigne qui tí a. pa.s pénétré dans la peninsula directement par les voies
méridionales, mais l'a envahie du nord par des routes plus indirectes. La rupture
du Grand glacial, donnant naissance à deux fleuves courant en sens opposé, le
présent Saginaw prenant cours vers l'est et le présent Grand s'écoulant vers l'ouest,
eut lieu avant que les espèces?, cruciatumet P. pune tifemtu eussent pu parvenir de
1' ouest dans le bassin du Saginaw. La répartition, restreinte largement ou exclu-
sivement aux Grands Lacs bordant l'état, de P. conventus, P. idahoense et S. nitidum,
espèces d' eau froide et profonde, s' explique à base écologique; celle de P. hen-
slowanum, P. amnicum et S. corneum ,espèces d'origine européenne, qui sont
localisées dans les Grands Lacs et leur ligne de drainage, par leur invasion pro-
bablement récente. Les causes immédiates de la distribution limitée de certaines
autres espèces sont moins apparentes.
En général l'on peut dire que la distribution orginale de beaucoup de sphaeriides
dans les bassins riverains du Michigan a été obscurcie d'une part par une migration
active pendant les périodes alternantes d'inondations et de niveaux bas qui finalement
rompirenet les confluences antérieures et, d'autre part, par un transport passif,
soit à l'intérieur de ces mêmes voies d'eau au moyen d'autres animeaux aquatiques
tels qu'écrevisses, grenouilles ou poissons, soit à travers la région par l'entremise
d'oiseaux aquatiques. Le tracé de la distribution originale n'est encore clairement
visible que pour les espèces?, cruciatum, P. punctiferum et S transversum , tandis
.
qu'il est partiellement masqué pour les espèces P.fallax, P. insigne, P. walkeri et
S. /abale.
SPHAERIID DISTRroUTION IN MICHIGAN 159
RESEÑA
LA DISTRIBUCIÓN DE SPHAERIIDAE (PELECYPODA) EN MICHIGAN, E.E.U.U.
Como la mitad de las 31 especies y sus formas de esféridos que habitan hoy
en Michigan son más menos de distribución general en el estado, aunque el resto
tienen un área geomorfológicamente y ecológicamente restricta. Durante el
Pleistocene, estos esféridos presumiblemente repoblaron la región, desde el sur,
por migración activa remontando las aguas a través de las confluencias de los
cursos posglaciales. Las rutas mayores de migración desde el río Mississippi
hacia el interior de Michigan fueron: (1) Hacia la Alta Península de Michigan, el
Valle del Río Fox en Wisconsin oriental, y (2) Hacia la Baja Península el canal
niinois--Des Plaines que drenaba el lago glacial Chicago (la cuenca sur del presente
Lago Michigan) y el Rio glacial Maumee que drenaba el Lago Maumee (cuenca del
presente Lago Erie).
Después que la parte sur de la Baja Península de Michigan fué repoblada, su
dispersión fué diversamente obstaculizada. El Rio Grande glacial limitó la dis-
persión hacia el norte de Pisidium cruciatum, P. punctiferum y Sphaeriuní trans-
,
versum, y más tarde se dividió en el presente Rio Saginaw de curso oriental y el

Rio Grande que corre hacia el oeste antes que P. cruciatum y P. punctiferum
pudieran entrar en la corriente del Saginaw desde el oeste. La distribución,
restricta mayormente a los Grandes Lagos fronterizos del estado, de P. conventus,
P. iâahoense,, y s. nitidum, las cuales son especies de aguas profundas y frias,
puede explicarse ecológicamente; lade P./îêwstowanMw, P. amnicum and S. comeum,
que son restrictas a los Grandes Lagos y sus drenajes, puede explicarse por su
reciente importación de Europa. Las causas inmediatas de las ocurrencias locales
de muchas otras especies y formas, sin embargo son menos aparente.
La migración activa durante periodos que alternaban entre inundaciones y aguas
bajas, que terminó en la desuniónde las confluencias, el transporte pasivo mediante
otros animales acuáticos como langostas de agua dulce, ranas y peces, y el trans-
porte aereo por aves acuáticas, alteraron la distribución original de los esféridos
en Michigan. El tipo original es todavía evidente en la distribución de P. cruciatum,
P. punctiferum, y S. transversum, pero está parcialmente disimulado en P. fallax,
P. insigne, P. obtusale,P. walkeri, y S. fabale.
SPHAERIIDAE (BIVALVIA) ,.
,, , -
.
X.
31
, , , --
, ,, . --
160
,
2)
p.
,) W.
: H.
(
-
HEARD
)
,
(, -
,, , ).-
-
-
.; ,: , ,
-
-
-
Pisidium cniciatum,
.
,
ciatunj,
,
P. punctiferum
, , . .
Sphaerium
,
piinctiferiim
transversum.
, .
-
insigne,
.
-
-
-
Oepa^
S. nitidmn
., , ,, --
-
. conventum, .
.
idahoense
henslo-
wanum, P. amniciim
.
, , , -.-
S. corneiim,
-
,
, . , , -
--
-
, criiciatiwi, . puncti-
ferum S. transversum,
. fallax, . insigne, . obtusale, . walkeri
S. fabale.
DIRECTIONS TO AUTHORS
Malacologia will publish original monographs and mendations of the "Style Manual for Biological Journals"
longer papers devoted primarily or exclusively to the obtainable from the American Institute of Biological
study of mollusks. It aims to provide a common med- Sciences, 200 P Street, N.W., Washington 6, D.C. In
ium for such different aspects of malacology as anat- particular, simplified practices, such as the following
omy, ecology, medical malacology, paleontology, physi- are favored: numbers should not be written out except
ology, and taxonomy. The journal will try to combine at the beginning of a sentence; percentages following
scholarly standards with prompt publication. All manu- a number are expressed as %; abbreviations of meas-
scripts will be reviewed by at least two editors. ures (after a number): mm, ml, kg, etc. have no
Manuscripts (for the time being) may be in English, period, nor an s in the plural.
French, German, Russian or Spanish. They must con- Bibliography. See current number of Malacologia
tain a concise but adequate abstract for translation for desired form of citing.
into the other languages. Manuscripts should be type- Reprints. Authors will receive 25 gratis; additional
written and double-spaced and are to be submitted in copies may be obtained at cost price if ordered at the
duplicate (carbon copy on thin paper). Figures should time off-set proof is returned. Later orders cannot
be carefully executed and with figure numbers and be considered.
labels large enough to be read easily after reduction. Correspondence Address manuscripts to an ap-
.
Style will be changed as little as possible, and then propriate Editor according to area of specialization or
mainly in places of possible ambiguity or where econ- geographic location. Subscription requests, payments
omy of space may be achieved. Contributors in the and enquiries should be sent to the Managing Editor,
English language are advised to follow the recom- J. B. Burch, Museum of Zoology, University of Michi-
gan, Ann Arbor, Michigan, U.S.A.
ANWEISUNGEN
Malacologia nimmt Originalmonographien oder län- Styl. Die Schriftleitung wird die Schreibweise der
gere Arbeiten die sich hauptsächlich oder ausschliess- Abhandlungen möglichst wenig abändern und zwar
lich mit dem Studium der Mollusken befassen zur hauptsächlich um Platz zu sparen oder um allfällige
Veröffentlichung an. Unsere Zeitschrift bezweckt den Unklarheiten zu vermeiden. Es wird gebeten Zahlen
verschiedenen Gebieten der Weichtierkunde wie z.B. nur am Anfang eines Satzes voll auszuschreiben und
der Anatomie, Ökologie, Paläontologie, Physiologie, sonst in Ziffern auszudrücken; Hundertsätze (nach
Taxonomie oder der medizinischen Malakologie ein einer Zahl) werden mit dem Zeichen % wiedergegeben:
gemeinsames Ausdrucksmittel zu schaffen und ist be- Abkürzungen der Masseinheiten wie mm, ml, kg,
strebt strenge wissenschaftliche Anforderungen mit U.S.W, sind ohne Punkt zu schreiben.
raschester Veröffentlichung zu vereinen. Beiträg-. Schrifttitni . Die gewünschte Form ist aus jedem
werden von zumindest zwei Redakteuren geprüft. beliebigen Heft von Malacologia ersichtlich.
Beiträge können gegenwärtig in englischer, fran- Sonderdrucke. Die Verfasser erhalten zumindest
zösischer, deutscher, spanischer oder russischer 25 Stück unentgeltlich. Zusätzliche Exemplare können
Sprache abgefasst sein; ein kurzer, jedoch genügend zum Selbstkostenpreis erworben werden; Bestellungen
vollstandigerAí<s2í<§^ wird verlangt, zwecks Übersetzung sind gemeinsam mit der Korrekturfahne einzuschicken;
in die anderen Sprachen. Manuskripte sind mit Schreib- spätere Bestellungen können nicht berücksichtigt werden.
maschine, doppelzeilig auszuführen und in zwei Ex- Korrespoiulenz. Manuskripte sind an einen der
emplaren (Durchschlag auf dünnem Flugpostpapiei^ sprachlich sowie im Gebiete zuständigen Schriftleiter
zu unterbreiten. Zeichnungen müssen sorfältigst aus- zu senden. Abonnementsbestellungen, Zalilungen und
gearbeitet sein und Bezeichnungsziffern, — Buchstaben sonstige Anfragen sind an den Managing Editor, J. B.
oder sonstige Inschriften gross genug, um nach Ver- Burch, Museum of Zoology, University of Michigan,
kleinerung immer noch deutlich lesbar zu bleiben. Ann Arbor, Michigan, U.S.A., zu richten.
DIRECTIVES
Malacologia accepte à publier monographies orig- autres inscriptions suffisamment grands pour demeurer
inales et mémoires longs, dévoués principalement ou clairement lisibles après réduction.
exclusivement à l'étude des mollusques. Ce journal Style. En principe, les éditeurs ne changeront le
a pour but de fournir un organe d'expression commun style des articles que pour éviter ambiguïté possible
pour divers sujets en malacologie, tels quel'anatomie, ou pour économiser de l'espace. Les auteurs sont
l'écologie, la paléontologie, la physiologie, la taxon- priés de mettre les nombres en chiffres et de ne les
omie ou lamalacologie médicale, en s'efforçant à écrire en lettres qu'au début d'une phrase; pour-
unir des critères d'excellence rigoureux à une publi- centages, après nombres, seront représentés par le
cation prompte. signe % et les abbreviations des mesures (après un
Manuscrits. Les contributions pourront être, à nombre) tels que mm, ml, kg, n'auront ni point ni
l'instant, en langue anglaise, française, allemande, "s" au pluriel.
espagnole et russe; un abstrait bref, mais suffisam- Bibliographie. N'importe quel numéro courant de
ment complet y sera joint, pour traduction. Les Malacologia montrera la présentation désirée.
manuscrits seront écrits à la machine, à double es- Séparés. Les auteurs recevront gratuitement 25
pace, et seront soumis en deux exemplaires (double tirés à part. Des copies additionelles pourront être
en papier carbone, sur papier fin). Les dessins sont acquises à prix coûtant et les commandes y relatives
à exécuter soigneusement, avec lettres, chiffres ou seront placées au retour des épreuves. Les com-
mandes placees plus tard ne pourront plus être prises spécialisé. Les commandes d'abonnement, payements
en considération. et correspondances diverses seront adressées au Man-
Correspondence Les manuscrits seront addresses
. aging Editor, J. .
Burch, Museum of Zoology, Uni-
à un éditeur approprié quant à la langue et au sujet versity of Michigan, Ann Arbor, Michigan, U.S. A.
DIRECCIONES PARA LOS AUTORES

Malacologîa publicará monografías originales, asi los siguientes factores simplificados: los números
como estudios dedicados primera y exclusivamente al no deben escribirse, excepto al principio de una
estudio de los moluscos. Se propone proporcionar un oración; los porcentajes (símbolo) que siguen de-
medio común para los diferentes aspectos de la Mala- spués de un número deben expresarse asi: %; las
cologîa, tales como Anatomía, Ecología, Malacologîa abreviaturas de medidas (después de un número): mm,
Médica, Paleontología, Fisiología, y Taxonomía. Esta ml, kg, etc., no debiendo usarse punto después de la
publicación tratará de combinar ambos, los artículos abreviatura o eses en los plurales.
de rigurosa excelencia con la rapidez en la publica- En lo que se refiere a la bibliografía, véase un
ción. Todos los manuscritos serán revisados por lo número reciente de Malacologîa a cerca de la forma
menos por dos editores. de referencia.
Los manuscritos, por ahora, pueden ser en inglés, Reimpresos. Los autores de artículos recibirán
francés, alemán, ruso o español. Deben ser con- veinticinco ejemplares gratis. Copias adicionales pue-
cisos y adecuamente reseñados para su traducción en den obtenerse a precio de costo si se ordenan al tiempo
otros idiomas. Deben ser escritos a máquina, a de que se retorna la prueba al autor. Copias orden-
doble espacio, debiendo enviarse dos copias (la copia adas más tarde no serán consideradas.
en papel carbón debe ser en papel delgado) Las
. Correspondencia. Dirija o envíe los manuscritos
figuras deben ser claramente ejecutados, en números y a un Editor apropiado de acuerdo con el área de
leyendas suficientemente grandes para que puedan especialización o posición geográfica. Pedidos de
leerse fácilmente aún después de reducidos. suscripciones, pagos e información deben ser enviados
El entilo se cambiará tan poco como sea posible, al Director- Editor, J. B. Burch, Museo de Zoología,
principalmente en lugares que se presten a ambigüe- Universidad de Michigan, Ann Arbor, Michigan,
dades o donde la economía de espacio pueda asegurarse. Estados Unidos de América.
Los autores de artículos deben remitirse al uso de
MALACOLOGIA is published at irregular intervals MALACOLOGIA wird vom Institute of Malacology,

by the Institute of Malacology, Inc., 2415 South Circle Inc., 2415 South Circle Drive, Ann Arbor, Michigan
Drive, Ann Arbor, Michigan, U.S.A. The members of U.S.A. in unregelmässigen Zeitabständen veröffentlicht.
this Institute, also serving as editors, are: Die ebenfalls als Schriftleiter fungierenden Mitglieder
dieses Institutes sind:
MALACOLOGIA se publicará a intervalos irregu-
MALACOLOGIA est publié a intervalles irréguliers lares por el Institute of Malacology, Inc., 2415 South
,
par le Institute of Malacology, Inc., 2415 South Circle Circle Drive, Ann Arbor, Michigan, U.S.A. Los mi-
Drive, Ann Arbor, Michigan U.S.A. The sponsor mem- embros adhérentes de este instituto, quienes también
bers of this Institute, also serving as editors, are: servirán como editores, son:
D. W. TAYLOR, President
,
Circle Drive,
--:
,,.
M. R. CARRIKER, President-Elect
2i(.15 South
ROBERTSON, Vice President

R.
U.S. Geological Survey Systematic s- Ecology Program The Academy of Natural Sciences
U.S. National Museum Marine Biological Laboratory of Philadelphia
Washington 25, D.C., U.S.A. Woods Hole, Massachusetts, U.S.A. Philadelfáiia 3, Pennsylvania, U.S.A.
A. G. SMITH, Vice President J.B. BURCH, Secretary-Treasurer E. G. BERRY

Department of Invertebrate Zoology Museum of Zoology Laboratory of Parasitic Diseases
California Academy of Sciences The University of Michigan National Institute of Allergy
San Francisco 18, California, U.S.A. Ann Arbor, Michigan, U.S.A. and Infectious Diseases
National Institutes of Health
Bethesda 14, Maryland, U.S.A.
N. F. SOHL
U.S. Geological Survey
U.S. National Museum
Washington 25, D.C, U.S.A.
Vol. 1, No. 1 MALACOLOGIA October 1962
CONTENTS
Page
MALACOLOGIA, An International Journal of Malacology 1
D. W. TAYLOR and N. F. SOHL

An outline of gastropod classification 7
H. W. HARRY
A critical catalogue of the nominal genera and species of
neotropical Planorbidae 33
J. B. BURCH
Cytotaxonomic studies of freshwater limpets (Gastropoda:
Basommatophora) I. The European Lake Limpet,
Acroloxus lacustris 55
I. STIGLINGH, J. A. VAN EEDEN and P. A. J. RYKE

*
Contributions to the morphology of Bulinus tropicus
(Gastropoda: Basommatophora: Planorbidae) 73
H. J. WALTER
Punctation of the embryonic shell of Bulininae (Planorbidae)
and some other Basommatophora and its possible taxonomic-
/- phylogenetic implications 115
W. H. HEARD
Distribution of Sphaeriidae (Pelecypoda) in Michigan, U.S.A 139
VOL 1 NO. 2 JULY 1963
MALACOLOGIA

ANNE GISMANN,
MALACOLOGIA
General Editor B.BURCH, Managing Editor M. HUBER, Business Manager
19, Road 12 Museum of Zoology Museum of Zoology
Maadi, Egypt The University of Michigan The University of Michigan
U. A. R. Ann Arbor, Michigan, U. S. A. Ann Arbor, Michigan, U, S. A.

SCHRIFTLEITUNGSRAT CONSEIL DE RÉDACTION
P. , AGOCSY N. A. HOLME W. L. PARAENSE

Magyar Nemzeti MÚzeum Marine Biological Association Centro Nacional de Pesquisas
Természettudományi Múzeum of the United Kingdom Malacológicas
Baross U, 13 The Laboratory, Citadel Hill Caixa Postal, 2113
Budapest, VIII., Hungary Plymouth, Devon, England Belo Horizonte
Minas Gérais, Brasil
. H. BARNARD
G. P. KANAKOFF
South African Museum J. J. PARODIZ
Los Angeles County Museum
Cape Town Carnegie Museum
900 Exposition Boulevard
Republic of South Africa Pittsburg 13, Pennsylvania
Los Angeles 7, California
R. BOETTGER U. S. A.
, S. A.
Technischen Hochschule R, D. PURCHON

A. M. KEEN Dept. of Botany and Zoology
Braunschweig
Department of Geology
Braunschweig Chelsea College of Science
Stanford University
Pockelsstrasse 10a and Technology
Stanford, California, U. S. A.
Germany London, S. W. 3, England
A. H. CLARKE, JR. Y. KONDO S. G. SEGERSTRÂLE

National Museum of Canada Bernice P. Bishop Museum Zoological Museum of
Ottawa, Ontario, Canada Honolulu 17, Hawaii, U. S, A. Helsinki University
P.-Rautatiekatu 13
J. DUNCAN N. MACAROVICI Helsinki, Finland
Department of Zoology Laboratoire de Géologie
J. STUARDO
University of Liverpool Université "Al. I. Cuza"
Departamento de Zoología
Liverpool, England Ia§i, Romania
Instituto Central de Biología
E. FISCHER-PIETTE Universidad de Concepción
Museum National d'Histoire D. F. McMICHAEL Cas. 301, Concepción, Chile
naturelle The Australian Museum
W.S. S. VAN DER FEEN - VAN
55 rue de Buff on College Street
Sidney, Australia
BENTHEM JUTTING
Paris V^, France
Zoologisch Museum
A. FRANC Amsterdam, The Netherlands
J. E. MORTON
Faculté des Sciences
Department of Zoology M. YONGE
55 rue de Buff on
The University of Auckland Department of Zoology
Paris Ve, France
Auckland, New Zealand The University
K. HATAI Glasgow, Scotland
Institute of Geology and W. K, OCKELMANN A. ZILCH
Paleontology Marine Biological Laboratory
Senckenberge-Anlage 25
Tohoku University Gr(íínnehave, Helsingdr
6 Frankfürt am Main 1
Sendai, Japan Danmark
Germany
MALACOLOGIA was established with the aid of a grant (NSF-G24250) from the National Science Foundation,
Washington, D. C, U. S. A.
MALACOLOGIA wurde unter Beihilfe einer Unterstützung (NSF-G24250) der National Science Foundation,
Washington, D. C, U. S. A., gegründet.
MALACOLOGIA fut établi avec l'aide d'une subvention (NSF-G24250) de la National Science Foundation,
MALACOLOGIA
Foundation, Washington, D, C, U. S. A.
,
fue establecida con la ayuda de una subvención (NSF-G24250) de la National Science
. (NSF - G24250) -
MALACOLOGIA is prepared in the Mollusk Division, Museum of Zoology, The Univerisity of Michigan and
published at irregular intervals by the Institute of Malacology, 2415 South Circle Drive, Ann Arbor, Michigan,
U. S. A. The Sponsor Members of this Institute, also serving as editors are:
MALACOLOGIA wird in der "Mollusk Division, Museum of Zoology" der Universität von Michigan fertiggestellt
und in unregelmässigen Zeiträumen vom "Institute of Malacology", 2415 South Circle Drive, Ann Arbor, Michigan,
U. S. A. herausgegeben. Die gründenden Mitglieder dieses Institutes, die auch als Schriftleiter fungieren sind:
MALACOLOGIA est élaboré au Département de Mollusque de Musée de Zoologie du l'Université de Michigan
et publié à intervalles irréguliers par l'Institut de Malacologie, 2415 South Circle Drive, Ann Arbor, Michigan,
,
. S. .Les membres du Comité de Patronage de cel Institut constituent en même temps le Comité de Publication
de MALACOLOGIA. Ce sont:
MALACOLOGIA se publicará a intervalos irregulares por el Institute of Malacology, Inc., 2415 South Circle
Drive, Ann Arbor, Michigan, U. S. A. Los miembros adhérentes de este instituto, quienes también servirán como
editores, son:
D. W. TAYLOR, President
U. S. Geological Survey
:
Drive, Ann Arbor, Michigan, U. S. A.
.
-
R. CARRIKER, President-Elect
Systematics-Ecology Program
R.
2415
ROBERTSON,
The Academy
South Circle
Vice-President
of Natural
U. S. National Museum Marine Biological Laboratory Sciences of Philadelphia

Washington 25, D, C, U. S. A. Woods Hole, Massachusetts, U. S. A. Philadelphia 3, Pennsylvania,
U. S. A
A. G. SMITH, Vice-President E. G. BERRY, Vice-President N. F. SOHL, Secretary
Department of Invertebrate Zoology Laboratory of Parasitic Diseases U. S. Geological Survey
California Academy of Sciences National Listitute of Allergy U. S. National Museum
San Francisco 18, California, U. S. A. and Infectious Diseases Washington 25, D. C, U.S.A.
National Institutes of Health
Bethesda 14, Maryland, U. S. A.
J. B. BURCH, Treasurer
Museum of Zoology
The University of Michigan
Ann Arbor, Michigan, U. S. A.
Participating Members of the Institute of Malacology are:

Aktiv teilnehmende Mitglieder des Malakologischen Institutes sind:
Les membres actifs de l'Institut de Malacologie sont:
P.
Institute of Malacology
Third Annual Meetir^
Washington, D. C, August 1963
Notice of Third Annual Meeting of the Sponsor Members of the Institute of
the
Malacology hereby published as provided by Article III, Section 1 of the Constitution
is
of the Institute. Participating Members and Members-at-Large on record as of July 1,
1963 may attend the meeting if they so desire, but such attendance is not necessary
for proper handling of business matters. The meeting will be held sometime during
the International Congress of Zoology in Washington, D. C, U. S. A., August 20-27, 1963.
Notice of a specific time and place will be available at the Congress.
Malakologisches Institut
3. jährliche Sitzung
Washington D. , August 1963
Gemäss Artikel III, Abschnitt I, der Verfassung des Malakologischen Instituts wird
hiemit die 3. jährliche Zusammenkunft der gründenden Mitglieder des Instituts öffentlich
bekanntgegeben. Aktive Mitglieder, sowie alle bis zum 1. Juli 1963 ordnungsgemäss
eingetragenen ordentlichen Mitglieder, können, falls sie es wünschen, der Sitzung
beiwohnen. Anteilnahme ist jedoch für den Ablauf der Geschäfte nicht erforderlich.
Die Zusammenkunft findet in Washington D. C, währenddes Internationalen Zoologischen
Kongresses vom 20.-27. August 1963 statt, bei welchem dann Näheres Über Ort und
genauen Zeitpunkt in Erfahrung zu bringen ist.
Institut de Malacologie
Session Annuelle
Washington D. , Août 1963
Selon l'article de la section I de la constitution de l'Institut de Malacologie, nous

annonçons par présente publication la 3^"^^ réunion annuelle du Comité de Patronage
la
de l'Institut. Les membres actifs ainsi que les membres associés régulièrement
inscrits en date du l^r Juillet 1963 peuvent, s'ils le désirent, assister à la session,
mais leur présence n'est point nécessaire pour le déroulement des affaires. La
réunion aura lieu à Washington D. C, à l'occasion du Congrès International de Zoologie
allant du 20 au 27 Août 1963, pendant lequel des renseignements plus précis quant au
lieu et à l'heure de la réunion pourront être obtenus.
Instituto de Malacologia
Tercer Mitin Anual
Washington, D. , Agosto 1963
Convócase, de acuerdo con las provisiones del Artículo III, sección la de los Estatutos
del Instituto, al Tercer Mitin Anual de los Miembros Patrocinadores. Miembros regu-
lares y Adhérentes, registrados hasta el IQ de Julio, 1963, pueden concurrir al mitin
si asi lo desean, pero tal asistencia no es imprescindible para el manejo de cuestiones
administrativas y financieras. El mitin tendrá lugar en momento oportuno, durante
elCongreso Internacional de Zoología en Washington, D. C, Agosto 20-27, 1963. Fecha,
hora y lugar se notificará en ese Congreso,
,
, -- ,
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1961
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(161)
SOME ASSOCIATIONS OF MARINE MOLLUSICS AND ALGAE IN PUERTO RICO ^
Germaine L, Warmke and Luis R. Almodovar ^
ABSTRACT
During a study of marine moUusks associated with algae at La Parguera, Puerto Rico,
some 90 mollusk species were found living on 25 common species of algae. A total of
30,859 mollusks were obtained from 155 lots of algae in littoral zone ana coral reef
areas. Of those mollusks 99% were tiny gastropods, about 3/4% were pelecypods, and
about 1/4% Amphineura. Fifteen species of gastropods formed 94% of all mollusks col-
lected. Red algae appear to be the favored habitat, averaging 296 mollusks per lot.
Green algae ranked second, with 120 mollusks per lot. Brown algae yielded only 46
mollusks per lot collected.
Though most species of mollusks were collected on all groups of algae, in many cases
the animals showed a preference for one group. A few species showed a striking prefer-
ence for a specific alga, the best example being the association of Oxynoe antillarum with
the alga Caulerpa racemosa.
Numerous as these small mollusks proved to be on algae, the study brought to light
10 species not previously reported from Puerto Rico. These are as follows: Amphi-
thalamus vallei, Cyclostremiscus omatus, Marginellopsis serrei, Assiminea succinea,
CoTidylo cardia smithii, Cyclostremiscus pulchellus, Cylindrobulla beauii, Peristichia
agria,a.nd 2 unnamed species of Omalogyra.
INTRODUCTION —
nately have not been made in the Carib-
bean area.
Numerous small marine mollusks live While looking over various col-
on algae. Almost a century ago Bergh lections of reef and bay algae in Puerto
(1871) listed and described several species Rico, we found many marine mollusks, and
living onalgaeof theSargassumSea. Cole- this led to the present study of moUuscan-
man (1940)and Wieser (1952) supplied algal associations. The purpose of this
valuable information on the fauna (in- study was to identify and determine the
cluding mollusks) living on marine algae relative abundance of mollusks living on
in the Plymouth region of England. More common red, green, and brown algae at
recently Robertson (1960) pointed out that La Parguera off the Southwest coast of
algae on mangrove roots in the Bahama Puerto Rico (see map of collecting area).
Islands are thefavoredhabitatof many mi-
nute mollusks. To our knowledge, detailed METHOD OF COLLECTING
studies of association between mollusks
—
and algae particularly whether mollusks All algae were collected from clear
show preference for certain species of water, in depths of 1 to 10 feet, from bays
algae, or if they live on algae indiscrimi- and coral reef areas. The salinity in the
^This research was aided in part by National Science Foundation Grant 14020 awarded
to the junior author.
2 Curator of Mollusks and Algologist, respectively, Institute of Marine Biology, Univ-

ersity of Puerto Rico, Mayaguez, Puerto Rico.
(163)
164 WARMKE AND ALMODOVAR
-18°i
PUERTO RICO
Bahia Fosforescente
Bahia Montai va
Magueyes ^^';::=>
Magimo
<^^
Enri que ^^âta
Laurel (porral
Cristobal
îrooai /-) /-,
Media Luna *^^^Z!!^

Turrumote I
CARIBBEAN SEA
lin ni
05
FIG. 1. Map of collecting area, south coast of Puerto Rico.
collecting areas ranged between 34 and 36 mollusks that were alive at the time of col-
p. p.m.and the temperature from 27-32 C. lecting were recorded; obviously empty
To avoid contamination, each species of shells with hermit crabs are not included
alga was collected separately by the junior in the report. With the exception of four
author who used a face mask and snorkle. were made during the
lots, all collections
Only the part above the holdfast was col- months of January and February 1959 and
lected, thus eliminating any shells that October and November 1960.
might have been in the substrate at the base
of the plants. Each algal lot, which con- FINDINGS
sisted of approximately 1 pound wet weight,
was put into a plastic bucket and covered Distribution of all Mollusks on Algae
with 40% formalin. After 10 to 15 minutes
in the solution, the alga was agitated to In the present study 30,859 mollusks^
remove shells still clinging to the plants. were found living on 25 different species
The alga and liquid were removed, and the of algae.Of these, 30,529, or almost 99%,
residue was dried and examined under a belonged to the class Gastropoda. Only
binocular dissecting microscope. Only 258 Pelecypoda and 72 Amphineura were
SThese specimens will be divided and placed in the following Institutions: The Institute
of Marine Biology, Mayaguez, Puerto Rico; the Academy of Natural Sciences of Phila-
delphia; the Museum of Comparative Zoology at Harvard College; and the U.S. National
Museum in Washington, D.C.
ASSOCIATIONS OF MARINE MOLLUSKS AND ALGAE 165
collected (Table 1). lusks are, few are ever found by the
The red algae harbored many more average collector, and little is known about
mollusks than either the green or brown their habits. Of the ten most abundant
(Table 1). A total of 23,706 mollusks were gastropods collected from algae in the
collected from 80 lots of red algae, aver- present study, four have not previously
aging 296 specimens per lot. On the green been reported from Puerto Rico (see
algae, 5,997 mollusks were collected from Table 2).
50 lots, or an average of 120 specimens Amphithalamus vallei was by far the
per lot. The 25 lots of brown algae yielded most common mollusk found living on algae
a total of 1,156 mollusks, or 46 per lot. during this study. Almost one -half of the
total number of specimens collected were
Distribution of Gastropods on the Three this species. Surprisingly enough it had
Groups of Algae not previously been reported from Puerto
Rico. It was more than twice as common
The 15 most common species of as the next most abundant species, Bitt-
gastropods found living on algae, arranged ium varium. Robertson (1960) found A.
in decending order of abundance, are shown vallei to be themost abundant species in
in Table 2. These 15 species, with a total the red algae on mangrove roots in the
of 29,277 specimens, represent 94% of the Bahamas.
total number of mollusks collected during
this study. Distribution of Gastropods on Red Algae
Table 2 also shows the distribution
of these gastropods on the red, green, and Table 3 shows the distribution of 31
brown algae. Although most species were species of gastropods on 12 species of red
collected on all groups of algae, in many algae. The minute rissoid, Amphithalamus
cases the animals showed preference for vallei, was the most abundant gastropod
one group. The following species show collected on red algae. It averaged almost
preference for the red algae: Amphi- 156 specimens per lot on the 80 lots
thalamus vallei Aguayo and Jaume, Bittum studied. This is twice as common as
varium Pfeiffer, Columbella mercatoria Bittium varium, which averaged 68 shells
Linné, Assiminea succinea Pfeiffer, and per lot. Bittium shows a striking prefer-
Rissoella caribaea Rehder. It should be ence for Laurencia obtusa; 2,859, almost
noted that 79 to 92% of the specimens of one-half of the total specimens of this
each of these species were found on red species were found on that alga. Another
algae. species with a preference for a specific
The gastropods showing preference alga is Assiminea succinea. A total of
for the green algae were: Oxynoe antilla- 352 specimens of this species were col-
rum Mörch, Anachis pulchella Sowerby, lected, and of these, 283or 80%, were found
Cyclostremiscus omatus Olsson and on red algae. Most of these (269) were
McGinty, Caecum pul he Hum Stimp son, and found on one lot of Amphiroa fragilissimxi
Schismope cingulata O. G. Costa. Between collected in the bay area. All 41 specimens
58 and 98% of the specimens of each of of Cerithium variabile . Adams were
these species were collected on green found on the single red alga, Centroceras
algae. The only gastropod preferring clavulatum. However, this species is more
brown algae was Caecum (J\âeioceras) often found under rocks or in Thalassia
nitidum Stimpson, with 66% of its speci- beds.
mens on the brown algae. A total of 2,401 Rissoella caribaea
Most of the mollusks found living on were collected (Table 2); of these 1,905,
algae are minute gastropods. Ihe 11 or about 79%, were found on red algae,
most abundant species all are under 5 mm Laurencia papulosa was host to a little
in length as adults, and the majority are over one-half (1,054) of the ÄzssoeZZa found
2 mm or less. Abundant as these mol- on red algae.
In addition to the 31 species of gastro- 1,506, or almost one-fourth of the

pods listed in Table 3, there were numer- specimens, were Schismope cingulata.
ous others which appeared less commonly This minute (1 mm) snail averaged
or only as juvenile forms; these are in- 127 shells per lot on Cladophoropsis
cluded under the miscellaneous column at membranácea. Amphithalamus vallei, the
the end of the table. The miscellaneous second most abundant species on the green
gastropods, listed in alphabetic order, are algae, also shows a striking preference for
as follows: Acmaea (young), Alaba incerta C. mem,branacea. Four -fifths of all A.
Orbigny, Anachis (young), s ira ea (young), vallei from green algae (815 of 1,026) were
Atys riiseanaM.<5rch,Atys{young), Caecum on this alga.
coronellus Dall, Cerithium algicola C. B. Anachis pulchella Sowerby shows a
Adams, Crassispira juscescens Reeve, preference for the Halimeda Silga.e. A total
Crepidula (young), Emarginula (young), of 185 specimens of this species were col-
Epitonium (young), Euchelus guttarosea lected; of these 133, or almost 72%, (Table
Dall, Eulima sp., Fissurella (young), 2) were on the green algae. Table 4 shows
Haminoea (young), Hyalina sp., Marginella that 109 of the 133 specimens on green
láctea Kiener, Modulus modulus Linne, algae were found on Halimeda (96 on
Nitidella laevigata hinne, {young), Nitidella Halimeda opuntia and 13 on Halimeda
ocellata Gmelin, Odostomia.canaliculata . tridens). Most of the shells of this species
. Adams, Odostomia laevigata Orbigny, were young; of the 185 total, only 18 were
Oxynoe antillarum Mörch, Pedipes mir- adults. This could indicate either that
abilis MQhlfeld, Peris tichia agria Dall, Anachis pulchella spends only the early
Persicula pulcherrimxi Gaskoin, Planaxis stage of its life on algae, or that the col-
were made during
Adams, Pus¿a //
(young), Pseudostomatella coccinea A.
Dohrn, Pyrgocythara
sp., Pyramidella sp., Seila adamsi H. C.
lections
of the breeding season.
An
the early part
excellent example of a gastropod

Lea, Stomatella picta Orbigny, Tegula showing algal specificity is Oxynoe
fasciata Born, Tricolia adamsi Phillippi, antillarum. A total of 329 specimens
Tricolia bella M. Smith, Tricolia thalass- were collected; of these 324, or ap-
icola Robertson, Triphora sp.. Trivia proximately 98.5% (lable 2) were found
leucospkaera Schilder, (young), Trun- on the green algae. Further examination
catella sp., (young), Turridal (young), (Table shows that 322 specimens re-
4)
Triptychus niveus Mörch, Vitrinella covered on green algae were on Caulerpa
(young). (294 on Caulerpa racemosa and 28 on
Of the 23,518 gastropods found on red Caulerpa serhdarioides). The shells
algae, only 174, or less than 1% are listed ranged in length from less than 1mm for
in the miscellaneous column. Although the young specimens, up to 5nim for adults.
total number of specimens included in this This would indicate that O. antillarum
group was small, it represents some 44 spends most of its life on Caulerpa, pre-
different species (many incompletely ferring the species racemosa. The
identified). These 44, plus the 31 species yellow-green color of the live animal
included in Table 3, make a total of about serves as an excellent camouflage on
75 species of gastropods found living on Caulerpa. This is in accord with findings
red algae. of Gonor (1961), Keen and Smith (1961),
and others indicating that Sacoglossans
Distribution of Gastropods on the Green often are found in association with
Algae . Caulerpa.
In addition to the gastropod species
Table 4 shows the distribution of the listed in Table 4, there were numerous
31 most common gastropods on 10 species others which appeared less commonly or
of green algae. A total of 5,926 specimens only as juvenile forms; these are included
were found on the green algae; of these under the miscellaneous column. These,
, ,
listed in alphabetical order, are as follows: sp., Planaxis lineatus, Tricolia adam,si,
Acmaea (young), Anachis (young), Bullata Vitrinella (young).
ovuliformis Orbigny, Cerithiopsis sp., Only 30 specimens were included
Crassispira (young), Crepidula (young), under the miscellaneous heading, but these
Eulima sp., Fissurella (young), Hipponix add 15 more species to the 31 listed on
(young), Hyalina {young) Lobiger souverbii Table 5, making a total of some 46 species
Fischer, Marginella láctea Kiener, of gastropods found living on brown algae.
Modulus (young), Nitidella (young), Peri-
stichia agria, Persi aula pulcherrima Distribution of Pelecypods on Algae
Gaskoin, Pedipes mirabilis MOhlfeld,
Planaxis lineatus da Costa, Pusza (young), The distribution of pelecypods found
Vitrinella (young). living on species of red, green, and brown
Only 111, or less than 2% of the algae is presented in Table 6, 7, and 8.
5,926 gastropods found on green algae, are Only 258 specimens, or less than l%of the
listed in the miscellaneous column. Al- total mollusks collected were pelecypods.
though this group is small, it includes some These pelecypods are listed below in
20 different species (most of them too order of relative abundance (the number of
young to be identified). These 20, plus the specimens follows each species).
31 species included in table 4, make a
total of approximately 51 species of Brachidontes exustus Linné (182)
gastropods found living on green algae. Condylocardia smithii Dall (13)
Musculus lateralis Say (13)
Distribution of Gastropods on Brown Algae Chione cancellata Linné, (young) (7)
Area imbricata Bruguiére, (young) (4)
Table 5 shows the distribution of the Area zebra Swainson, (young) (2)
31 most common gastropods on three Barbatia cancellaria Lamarck,
species of brown algae. Caecum {Meio- (young) (2)
ceras) nitidum was the most abundant Diplodonta sp., (young) (2)
gastropod on brown algae. This small, Anadara notabilis Röding, (young) (1)
pale -brown Caecum shows a strong Codakia sp., (young) (1)
preference for Dictyota bartayresii. Gouldia cerina .. Adams (1)
Of the 254 specimens of this species found Lima sp., (young) (1)
on brown algae 245, or 95%,were recovered Miscellaneous young pelecypods (29)
from D. bartayresii. Total number of 258
Another gastropod showing specificity
is Marginellopsis serrei; all 63 specimens In all, nine pelecypod species were
of this species were found on Dictyota identified; three others could be identified
divaricata. However, on the red algae, to genus only. Brachidontes exustus was
where 70% of the Marginellopsis were the only reasonably common species; 182
found, it was less specific. Zuanza auberi- or 70% of all the pelecypods found on algae
ana, when found on brown algae, also pre- were this species. With the exception of
ferred D. divaricata. Condylocardia smithii Musculus lateralis,
Miscellaneous gastropods, which ap- and Gouldia cerina all other species of
peared less commonly or only as juvenile pelecypods found were young. Conceivably
forms are shown at the bottom of Table 5. some of the bivalves that are attached by
These, listed in alphabetical order, are a by ssus were not dislodged by the formalin
as follows: Acmaea {young), Alaba incerta, treatment.
Astraea (young), Cerithium algicola, Among the pelecypods, one species,
Crepidula (young), Epitonium (young), Concylocardia smithii proved to be a new
Fissurella (young), Haminoea (young), record for Puerto Rico.
Liotia (young), Marginella (young), Odost-
omia canaliculata C.B, Ada.nis, Parvi tur bo
Distribution of Amphineura The brown algae are not common on
reefs. The two genera studied were se-
A total of 72 chitons were found on lected because of the abundant surface area
algae. This is less than 1/4% of the total they provide. Dîciyoto forms a thick carpet
specimens of mollusks collected. The over sandy bottoms and mollusks live
distribution of the chitons on the species among its protective branches.
of algae is given in Tables 6, 7, and 8.
The chitons are all less than 3 mm
in length
and probably are juvenile forms. Most of ACKNOWLEDGMENTS
these appear to be young Ischnochiton Gratitude is expressed to Dr. Robert
papillosus C. B. Adams. Robertson, of the Academy of Natural
The following species of algae aver- Sciences of Philadelphia, for his great help
aged about 1 chiton per lot: the red alga in the identification ofmany of the more
Spyridia filamentosa (7 chitons on 5 lots of difficult moUuscan species and for re-
algae); the green zlgdiCaulerpa eras sifolia viewing the manuscript. For help in col-
(2 chitons on 2 lots), a.ndHalimeda opuntia lection of live algae, special thanks are
(16 chitons on 13 lots); the brown alga due to Mr. Victor Rosado of the Institute
Dictyota bartayresii (17 chitons on 19 lots). of Marine Biology.
Importance of Red, Green, and Brown Algal

Species as Mollusk Habitats .
LITERATURE CITED
Twenty-five species of algae are
listed in Tables 9 and 10 in order of rela- BERGH, RUDOLF, 1871, Beiträge zur
tive importance as habitats for mollusks. Kenntniss der Mollusken desSargasso-
Eight of the 12 species of red algae aver- meeres. Verh. K. K. Zool.-Bot. Ges.
aged over 100 mollusks per lot (Table 9). Wien, Vol. 21: 1-36.
The filamentous red alga, Coelothrix, COLMAN, JOHN, 1940, On the faunas in-
grows in tangled cushions over rocks and habiting intertidal seaweeds. J. Marine
dead corals. Its wiry texture and growth Biol. Assoc. Vol. 24: 129-183.
habit doubtless are of importance in giving GONOR, J. J., 1961, Observations on the
shelter to such large populations of mol- biology oí Lobiger serradifalci,a. shelled
lusks. Centroceras, Acanthophora, Laur- sacoglossan opisthobranch from the
encia, Amphiroa, Ceramium, and Hypnea Mediterranean. Vie et Milieu, 12(3):
also have compact growth habits which 381-403, 5 figs.
provide suitable habitats for mollusks. KEEN, MYRA and SMITH, ALLYN, 1961,
Of the green algae, three species West American species of the bivalved
averaged over 100 mollusks per lot (Table gastropod genus Berthelinia. Proc.
10). Cladophoropsis membranácea is a Calif. Acad. Sei., 30: (2): 47-66.
common shallow -water alga that forms an ROBERTSON, ROBERT, 1960, The mol-
extensive mat up to 6 inches in thickness. lusk fauna of Bahamian mangroves [ab-
This mat provides ample space for abun- stract] Amer. Malac. Union Ann. Repts.,
dant moUuscan populations. Halimeda 1959, 22-23.
opuntia and Caulerpa racemosa possess a WIESER, W., 1952, Investigations on the
series of branches that are entangled and microfauna of seaweeds inhabiting
also provide shelter and food for many rocky coasts. J. Marine Biol. Ass.,
mollusks. 31: 145-174.

TABLE 1, Distribution of MoUusks by Classes on Red, Green, and Brown Algae

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TABLE 5. Distribution oí Gastropods on 3 Species of Brown Algae^^

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TABLE 8. Distribution of Pelecypods and Amphineura on 3 Species of Brown Algae^^

TABLE 10. Importance of Green Algae as Habitats for Mollusks
Species of Alga^"^
environ 3/4% des pélécypodes et environ 1/4% des amphineures. Pour ce qui concerne
les gastéropodes, 15 espèces formaient les 94% des mollusques recueillis. L'habitat
favori semblait être fourni par les algues rouges, qui abritaient en moyenne 296 mol-
lusques par lot. Les algues vertes venaient en second lieu avec 120 mollusques par
lot tandis que les algues brunes n'hébergaient que 46 mollusques par lot recueilli.
Quoique la plupart de ces mollusques furent trouvés sur chacun de ces 3 groupes
d'algues, ces animaux montraient souvent une préférence pour l'un ou l'autre de ces
groupes. Certaines espèces montrèrent même une préférence frappante pour une
espèce particulière d'algue, dont le plus éclatant exemple est fourni par l'association
d'Oxynoe antillarum avec l'algue Caulerpa racemosa.
Dix de ces espèces de petits Mollusques n'étaient pas encore connus de Puerto Rico
in dépit de l'abondance qu'ils y montrent sur les algues. Ce sont: Amphithilnmus
vallei, Cyclostremiscus omatus, Margine Hops is serrei, Assiminea succinea,
Condylocardia smithii, Cyclostremiscus pulchellus, Cylindrobulla beauii, Peristichia
agria, ainsi que de 2 espèces non-determinées d'Omalogyra.
RESUMEN
ALGUNAS ASOCIACIONES DE MOLUSCOS MARINOS Y ALGAS DE PUERTO RICO

Durante un estudio de moluscos marinos que viven sobre algas en La Parguera,
Puerto Rico, un total de 30.859 individuos perteneciendo a alrededor de 90 especies
fueron encontrados en 155 lotes de 25 especies de algas comunes colectadas en las
zonas litoral y de arrecifes. De estos moluscos, 99% eran minúsculos gastrópodos,
alrededor de 3/4% pelecípodos y 1/4 anfineuros. Quince especies de gastrópodos
formaban el 94% de todos los moluscos colectados. Algas rojas eran el habitat favorito
con un término medio de 296 individuos por lote. Algas verdes venían en segundo
lugar con 120. Algas pardas dieron solo 46 moluscos por lote.
Aunque la mayoría de las especies colectadas estaban presentes en todos los grupos
de algas, en muchos casos mostraban preferencia por un grupo, y en unas pocas esta
preferencia era decisiva por una alga específica, siendo el mejor ejemplo la asociación
de Oxynoe antillarum con el alga Caulerpa racemosa.
Entre los numerosos diminutos moluscos que se encontraron en las algas, 10 especies
no habian sido indicadas previamente para Puerto Rico. Estas son: Ampkithalamus
vallei, Cyclostremiscus omatus. Margine llopsis serrei, Assiminea succinea, Condvlo-
cardia smithii, Cyclostremiscus pulchellus, Cylindrobulla beauti, Peristichia agria y
dos especies aún no denominadas de Omalogyra.
, 90 . , ,
,.
.,,
,
, -
-
.
. .
30,859
155
-
25
.. ,
296
,
94^
99^
%
.-
-
46
fo
,
,
:
Caulerpa racemosa.
Amphithalamus
Assiminea succinea,
Cylindrobulla
10
vallei,
beauii,
Cyclostremiscus omatus,
Condylocardia smithii,
Peristicia agria
,.
Cyclostremiscus
antillarum
Marginellopsis
pulchellus,
Omalogyra.
-
serrei,
TAXONOMY AND ZOOGEOGRAPHIC RELATIONSHIPS OF THE
SOUTH AMERICAN NAIADES (PELECYPODA: UNIONACEA AND MUTELACEA)!
J. J. Parodiz and A. A. Bonetto
Carnegie Museum, Pittsburgh, Pennsylvania, U. S. A.
and
Consejo Nacional de Investigaciones Cientificas
Santa Fe, Argentina
ABSTRACT
A natural system of classification is proposed for the South American fresh-water

pearly mussels which were formerly all grouped in the superfamily Unionacea. The
systems recognized since the end of the last century which were based mainly on con-
chological, and partly on anatomical characters, are here discussed in relation to recent
embryological and phylogenetic research, especially as regards the structure and de-
velopment of the different types of larvae. Researches made by the authors during the
last decade have confirmed the existence of the "lasidium" larva, discovered by Ihering
in 1891, but not observed since by other authors. This larva is typical of the South
American genera Anodontites, Mycetopoda, Monocondylaea and Leila. At the same time
the research of other workers on African species of Mutela has revealed the existence
of a larva which, if not entirely similar to the lasidium, is similar in its basic structural
features. Comparative studies of such structures and their development show a family
differentiation between Mutelidae of Africa and Mycetopodidae in South America. On the
other hand, the close relation between these two families, and their extraordinary embry-
ological divergence from those other fresh-water mussels characterized by the well
known "glochidium" larva, warrants the distinction of a new superfamily, MUTELACEA.
All other South American fresh-water mussels with larvae of the glochidium type remain
in UNIONACEA.
The Mutelacea are living today in the southern hemisphere, excepting Australasia,
Whether South American groups are derived from African groups, or vice versa, is not
known. The anatomical and embryological differences between Mutelidae and the more
advanced Mycetopodidae seem to indicate an ancient separation. Paleontological records
are rare: none exist for Africa or Australasia; In North America, fossil casts from the
Triassic of Pennsylvania were referred to by Pilsbry as a Mycetopoda -like mussel;
Pleiodon priscus described by Ihering from the Cretaceous of Brasil is not a mutelid,
as was assumed, but belongs to the genus Paxyodon (Hyriidae). Some references to
Anodontites-\ik.e fossils from the Cretaceous of Bahia, Brasil are very doubtful.
The South American Mutelacea, i.e., the Mycetopodidae, are divided into 3 subfamilies:
Mycetopodinae, Anodontitinae and Monocondylaeinae; another subfamily, Leiliinae, might
be accepted in view of more recent anatomical and embryological research. Other
groups, at the subfamily level, indicated in previous classifications, cannot be main-
tained, their characters being insufficient and the intergradations numerous.
The South American Unionacea belong to the family Hyriidae, which also occurs in
Australia but is absent in the rest of the world; the exclusively South American forms
belong in the subfamily Hyriinae and are divided into 3 tribes: Diplodontini, Castaliini
and Prisodontini. Especially the Diplodontini, largely formed by the genus Diplodon,
are more closely related to forms of Australia and New Zealand, Here again, attempts
to separate a number of subgenera have failed on account of the difficulty of defining
constancy of characters. From the embryological point of view, however, we may dis-
tinguish two entities of subgeneric value: Diplodon s.S., with parasitic glochidia, and
Rhipidodonta with non-parasitic glochidia, i.e., having direct development. There is
paleontological evidence of Hyriidae in the North American Triassic, the Paleocene of
Southern Argentina, and the Eocene of Chile, the latter fossils being very similar to the
species now living in the region as well as to related groups from Australia. All these
fossils belong to the genus Diplodon, of which other species are known from the strata,
at different levels of the middle and upper Tertiary, scattered over the continent of
South America.
The monotypic genus Bartlettia of the so called "fresh-water oysters", currently
included within the Etheriidae, very probably belongs to a polymorph species of
Mutelacea, Anodontites tenebricosus. Larval stages of Etheriidae are unknown, but
further investigation may prove that the family, if it should be maintained as such, inte-
grates with the Mutelacea.
Comparative tables are given of the different systems of classification since 1900 as
well as of the new system adopted here, from the superfamüy down to subgeneric level.
^Research supported by a research grant, NSF-15032, to the senior author from the National Science Foun-
dation, Washington, D. C, U. S, A.
(179)
180 PARODIZ AND BONETTO
The current system of classification of Monocondylaea and Mycetopoda. The

the Neotropical fresh-water mussels dates organization and development of the lasid-
from the end of the last century, with im- ium proved to be more complicated than
portant improvements made during the last could be inferred from Ihering's original
three or four decades (Table 3). Some ana- diagnosis. Franc (1949) said that he found
tomical characteristics favor the concept "glochidia" (I) in the gills of Mutela dubia
of a single family, Mutelidae, with many and M. rostrata, but more recently (1959
genera not only from South America, but and 1961) Fryer of the East Africa Fisher-
also from Central America, Africa and ies Research Organization, in Uganda,
Australasia. The anatomical similarities, gave a complete description of the larva
however, are not consistently present, and oi Mutela bourguignati, with important and
embryological and conchological charac- detailed observations of its development.
ters, as well as Zoogeographie al factors, Its basic structure is that of a lasidium,
are at variance with that concept of single- although it is differentiated from the
ness. From the biological point of view, embryos of the South American genera by
especially, the division of Mutelidae into marked morphological features.
Mutelinae and Hyriinae is fundamentally
more distinctive than a separation at COMPARISON OF THE LARVAE OF THE
merely the subfamily level. AFRICAN AND SOUTH AMERICAN
In 1891 Ihering described a larval form, "MUTELIDS"
from one species of Mutelinae, Anodontites
wymanni Lea (= A. patagónica Lamarck) The African Larva
which was entirely different from the
"glochidium" larva until then considered The embryo described by Fryer from
common to all Unionacea. That type of Mutela (Fig. 1) has the anterior end of the
larva, named "lasidium", has a body di- body divided into two short lobes. The
vided into three recognizable regions: the second portion is covered by a single uni-
anterior, ciliated and somewhat conic or valve shell, furnished at the end with two
bell-shaped; the median, rounded and cov- rows of 3-7 small hooks and a row of small
ered by an indivisible shell, and the pos- spinulae. It develops while attached ex-
terior, forming two short lobes with cirri ternally to the body of a fish, probably by
or hooks placed in rows. Two peculiar means of the hooks.
ribbon-shaped appendages of considerable The larval
shell is folded on the sides
length evolve from the anterior end. and fused at the median ventral line,
Ihering later added the following remarks forming an integral, not bivalved, piece
to the description: "I know this larva only (Fig. 2, LS ). This shell is uncalcified.
in Glabaris [= Anodontites], but in Aplodon As the larva grows, two tubular append-
[=Monocondylaea], the anatomy and the egg ages, called "haustoria" by Fryer, are
agree so well that the larva can scarcely produced anteriorly, penetrating into the
differ. It is advisable now, to follow further fish's tissues and, apparently, acting as
the distribution of this larva in America both trophic and fixing organs. After this
and Africa" (Ihering 1893: 59). the organism experiences a complete
Because of the remarkable differences metamorphosis conducive to the organ-
between the two types of larvae and the ization of the juvenile mussel, and finally
fact that subsequent investigators failed to the haustoria-base is cut, initiating the
find, or to recognize, the lasidium, the free living stage. The size of this larva
existence of this larva remained doubtful, is over 150 miera, almost twice as large
Its rediscovery was reported by Bonetto as the lasidium known in the South Ameri-
(1951) with a preliminary description of can species» Larval specimens of Mwíeía,
the lasidium of Anodontites trapesialis kindly sent by Fryer, in different stages
from the Parana River, and further in- of development, allowed a more complete
vestigations revealed its presence also in comparative study.
SOUTH AMERICAN NAIADES 181
100 miera
FIG. 1. Larva of Mutela bourguignati FIG. 2. Larva of Mutela bourguignati

(Ancey) showing the long tentacle (from (Ancey), oblique-dorsal view (from Fryer
Fryer). 1961, Fig. 3). CL ciliated lobes; LS
embryonic shell.
The South American Larva
but in fact they seem to project from a
Investigations on the lasidium of An- more ventral position, in the portion half
odontites trapesialis forbesianus {Lbsl) and covered by the valve; they then expand,
its parasitic phase in fishes, revealed a fused over the shell at the base of the an-
coincidence of development with that of terior ciliated lobes. Thus, a conic en-
Mutela, but without formation of the tubu- closure is formed, with the apex toward the
lar appendages or haustoria indicated by anterior lower side of the larva. The upper
Fryer. The larva is attached to the host margins of the ribbons remain free,
by a cyst-like structure. forming a V-shaped canal extending about
The lasidium of Anodontites (Figs. 3-4) ten times the length of the larval body, and
is smaller, measuring an average of 85 then dividing into two or three branches.
miera or 56.6% of the size of the larva in Although the complexity of this process
Mutela. establishes a remarkable difference with
The two ciliated lobes of the frontal Mutela, the position of the axes are es-
portion, already indicated by Ihering, have sentially similar.
an elongate -conic shape, but it is obser- These differences, however, are con-
vable that the frontal portion can be sepa- siderable reduced in Monocondylaea, in
rated into two circular pieces, lying side which the lateral expansions are less de-
by side, and projecting shortly from the veloped, and also in Mycetopoda whose
embryonic shell. This section coincides, filament has no lateral expansions at all,
in general, with that of Mutela. as in Mutela. In Leila the lasidium differs
The filamentous appendages are very from that of Anodontites by its larger size
different. Ihering said that they consist the ribbon-like appendages are narrower,
of two wide flat ribbons emerging laterally, become thinner distally and the cirri or
line of folding anterior lobes cilia
hooks
posterior
lobes embryonic shell organ of fixation
25 miera
Dorsal view.
FIG. 3. hdiSidium oi Anodontites trapezialis forbesianus {heâ).
not
FIG. 4. Same as Fig. 3, with division of the anterior lobes, and posterior lobes
visible (folded ventrally).
.
hooks of the posterior end are apparently, single row at each side; they develop from
wanting. small papillae and are separated during the
In the larval body, the outstanding first days of life. The labial palps, di-
characteristic in both African and South gestive tract and heart do not differ. The
Americah forms is the uncalcified uni- mantle in Anodontites is closed from the
valve shell (Fig. 5), whereby they differ beginning to form the siphons, which do
essentially from the glochidia. The pos- not occur in Mutela.
terior end of the body is usually folded The ontogenetic processes in the African
under the ventral side, and shows curved and the South American forms are of
cirri or hooks (6-7 inAnodontites trape- evident common ancestral relationship.
sialis) forming a circle around a pair of They probably separated from all other
lobes; additional spinulae like those in Naiades with glochidia very early time
at a
Mutela are absent. in their evolution. On the basis of their
marked differences a more natural system
COMPARATIVE DEVELOPMENT OF THE of classification can be established.
JUVENILE MUSSEL
TAXONOMIC CONSIDERATIONS
The larva, inside the single, non-bi- (Compare with Table 3)
valved enclosure follows divergent ways
in Mutela and Anodontites Without discussing the very early essays
In Anodontites, the extremities are of classification that were merely con-
folded toward the center of the ventral chological, and artificial in their results,
side, while in Mutela the growth is longi- it is necessary to return to Simpson's
tudinal. The period of development is 25 synopsis of 1896, for the first seriously
days in Mutela, but may be shorter or founded system.
longer in Anodontites, 19 to 28 days. As Simpson distinguished two large fami-
a consequence of the differences in their lies, Unionidae and Mutelidae, the first
parasitic adaptations, the young mussel in with schizodont hinge and glochidium lar-
Mutela has an elongated body, regularly va, and the second with taxodont hinge
curved below and somewhat truncated (theoretically) and lasidium larva, thus ac-
anteriorly; in Anodontites it is short and cepting Ihering's discovery, although it
high (Fig. 6) and the surface of the valves was yet unknown how many genera had
is formed by a series of planes, offering this larva. The Unionidae included forms
a polyhedral shape. Inboth cases the cuti- from all continents in different degrees
cle of the embryonic shell adheres to the of relationship, and Mutelidae confined to
valves of the juvenile mussel, and the small the Neotropical region and tropical Africa.
and cylindrical ligament is located on the Since then, from the intensive and valua-
middle of the hinge line, simulating a ble work of Ortmann, to the most recent
chondrophore. In the first stage the shell speculations on classification, such as
is composed only of conchiolin, but it is those of Modell, several systems have been
slowly filled in isolated spots with calcium proposed, but they only complement or
carbonate. modify that of Simpson. Ortmann grouped
The internal organization is similar in the neotropical and notogeic forms of
both Mutela and Anodontites. The foot has Unionidae (equivalent to the "Lamphor-
a rudimentary byssogenous gland with its hamphus" group of Simpson), with the
separate channel, but some elements of Mutelidae of Africa and South America,
fixation indicated by Fryer for Mutela are in a single family Mutelidae, based on
not found in Anodontites; also, in the foot anatomical details of certain relevance,
of the latter, there is a pair of very large and also because the marsupia for the incu-
otocysts. Both adductors are present. bation of the larvae were located in the
The branchiae, formed by 13 ctenidia in inner laminae of the gills. He separated
Mutela and 7 - 8 in Anodontites, are in a the Mutelidae into subfamilies Hyriinae
25 miera
FIG. 5. Larval shell oí Anodontites trapeziales, dorsal and lateral -ventral views, in
the first day of parasitism on the fish Jenyssia lineata.
posterior adductor ligament anterior adductor
labial palps
anal granules of
aperture calcium
branchiae
larval shell
branchial
aperture
margin of
mantle
juvenile shell
byssogenous
gland
foot
200 miera
FIG. 6. Juvenile mussel oi Anodontites trapeziales îth new shell, and larval shell
still attached.
and Mutelinae. The most important, and the most over-

The differences between the African, looked, of Ortmann's taxonomical cons-
South American and all other known siderations were his own reservations with
Naiades are so remarkable that in all regard to the stability of the system. He
probability they do not belong to directly said in fact (1921:454) that: "It possibly
related groups, but rather represent di- might be advisable, in future, to elevate
vergent ways in the conquest of continental the two South American subfamilies to the
waters. The two different types of larvae, rank of families...", and "the Mutelidaeoi
i.e., glochidium and lasidium, cannot be Simpson (1900) correspond to our Mutel-
considered to be derived one from the other inae" (p. 455 footnote). Pg. 567: "al-
or from any hypothetical direct ancestry. though closely allied to Spatha [=Mutela],
Ortmann's sound and critical observations the South American genera form a group
of the the anatomical and conchological by themselves, and the [only] similarity of
characters, add support to such a con- Mycetopoda to Spatha in the anal opening
clusion. It is necessary to upgrade the apparently indicates only parallelism of
taxonomic categories in the family group development, no genetic relationship", and
in order to adjust the system to our present also (p. 568) that "According to our present
knowledge. knowledge, the two subfamilies are un-
AdP
AdA
AdP
AdA
FIG. 7. Anatomy of South American and African naiad genera. Top, left: Anodontites
(Mutelacea, Mycetopodidae); top right: Castalina (Unionacea, Hyriidae); below: Spath-
opsis (Mutelacea, Mutelidae). A, anal opening; AdA, Adductor anterior; AdP, adductor
posterior; B, branchial opening; E, elevator or dorsal scar; F, foot; EB, external
branchia; IB, internal branchia; PI, palps; P, protractor; RA, retractor anterior;
RP, retractor posterior; U, union of mantle separating anal and branchial openings.
doubtedly allied; but they are very sharply THE SOUTH AMERICAN MUTELACEA
separated by anatomical as well as shell-
characters, and it is impossible to form The family Mutelidae is only known from
an appropriate idea of their genetic con- Africa. In the New World they are replaced
nection". by Mycetopodidae, characterized by a
Regarding the condition of primitiveness hinge that is edentulous or has very rudi-
in these groups, Ortmann remarks: "It mentary teeth; by a wide prismatic layer
is not very likely that the Mutelinae on the internal margin of the valves, the
reached South America coming from presence of a supra-anal aperture, the
Africa" (p. 455). "It is hard to say which connection of the inner laminae of the
group is more primitive, since of the two branchiae with the visceral sac, and the
differing characters, the one (anal opening absence of dorsal muscle scars. The dif-
is more primitive in the American forms, ferences between the lasidium and the las-
the other (inner lamina of inner gill) more idium-like larva of Mutelidae already have
primitive in the African Spatha" (p. 567). been indicated.
These observations would have been In South America the two superfamilies
sufficient to justify the separation, even if Unionacea and Mutelacea, occupy over-
the system was based only on anatomy and lapping areas, though there are but
not on theembryology of the larger groups. marginal zones, having only the one or
Ortmann used the study of the South A- the other; these zones perhaps correspond
merican larvae to diagnose species, some- to different origins and times of dis-
times genera, but not at the family level. persion. The Mutelacea (Mycetopodidae)
So it is that both lasidia and glochidia are are more restricted in their southward
included in his Mutelidae, although the distribution (Map 1), occupying zones
larvae from African species were then still north, east and peripheral to the Pampas-
unknown and the South American lasidia Chaco districts, and some tributaries that
continued unobserved after Ihering. cross these districts and empty into the
The numerous genera developing through Parana River. In the affluents of the left
the embryonic stage of glochidium, are side of the Parana River the Mutelacea
separated into several families, according have a greater development than the Union-
to anatomical peculiarities, such as acea (Hyriidae) (Compare with Map 2). In
Margaritiferidae, Unionidae, andHyriidae. the northwest they extend into Central
Consequently, the larval condition has a America and Mexico, where the Hyriidae
taxonomic value, not merely at the family are absent.
but at the superfamily level. We have Invasion of northern forms into the lower
seen that, in the current system, it has La Plata River system probably occurred
only a minor importance, generic or spe- in relatively recent times (the basin as it
cific. We believe that adult mussels de- is known today, was formed during the
veloping from totally different embryos Pleistocene), through connection of tribu-
should not be in the same superfamily. taries of the Upper Paraguay with those
In conclusion, the Superfamily UNIONA- of the Amazon, as is the case with Ano-
CEA should be restricted to those groups dontites ensiformis. Only a single species,
with glochidium larva, and those with Anodontites puelchana (d'Orb.) is rarely
lasidium elevated to a new Superfamily found in northern Patagonia. No species
MUTELACEA. are known west of the Andes, in Peru or
A synoptic comparison of the two fami- Chile, a zone which is populated by
lies which comprise the MUTELACEA, Hyriidae of the genus Diplodon.
i.e., Mycetopodidae and Mutelidae, is given
in Table I.
TABLE 1. Comparative Characters of Mutelacea

40
Distribution of
MYCETOPODIDAE
in
South America
20
MAP 1. Distribution of Mycetopodidae (Mutelacea) in South America.

Mycetopodidae are divided into three Leila, he created Glabarinae. Since

subfamilies: Glabaris according to the majority of
authors is a synonym of Anodontites such
Mycetopodinae separation should be deferred until it may
be based on better grounds. Recognition
Edentulous shell very elongate and thin, of mere groups of species in the Anodont-
gaping variably anteriorly. Prismatic ites complex, as proposed byOrtmannand
layer narrower. Anal aperture with ten- Haas, is more acceptable. Regarding the
dency to close above, and supra-anal not possible use of Leilinae, see appendix.
well defined. Foot extraordinarily long,
cylindrical, ending in a knob, mushroom- FOSSIL MUTELACEA
like, protuberant. The lasidiumhasalong
anterior filament and resembles that of Paleontological evidence in Mutelacea is
Mutela more than those of other subfami- very poor. A single and very doubtful
lies. specimen was referred by Ihering (1912)
Genera: Mycetopoda, Mycetopodella. to the Cretaceous of southern Brasil as
Pleiodon priscus (discussed later). Pils-
Anodontitinae bry (1921: 36) described M>'ce¿o/)odad¿ZMc-
uli from Triassic beds in Pennsylvania
Shell edentulous, regular in shape. which also contain Diplodon: "While the
Valves not, or scarcely gaping. Peri- generic reference of the fossil [M.
ostracum marked with creases and folds. diluculi] is not positive, the interior being
Foot regular. Supra-anal aperture dis- unknown, its characters, so far as they are
tinct. Prismatic layer wide. Lasidium legible, agree well with Myceio/)oda, which
with very wide ribbon-like filament di- appear to indicate this genus or one closely
vergent at the distal end. similar". Four other Anodontites -like
Genera: Anodontites, Leila (see ap- species were described by Hartt (1870)
pendix, p. 206). and White (1887) under the generic names
"Unio" and "Anodonta" for the Bahia
Monocondylaeinae Series of Brasil regarded as Upper Cre-
taceous or possibly Pal eocene; these
Shell small, thick, solid and gaping. shells are smaller than the living Anodon-
Hinge with one or two tuberculiform teeth. tites, the hinge area is unknown, and the
Periostracum with cloth-like sculpture. generic identification uncertain; White
Prismatic layer wider. Supra-anal aper- suggested that some of them, A. mawsoni
ture and foot regular. Lasidium of an and A. allporti, may be Iridina, which is
intermediate type between the other sub- stillless likely.
families. Frenguelli (1945) described, among
Genera: Monocondylaea, Fossula, other fresh-water bivalves, several
Haasica, Tamsiella. species of Paleonanodonta and Paleo -
mutela, from Permian-Triassic strata of
The family name Mycetopodidae has Patquia, Argentina. These genera are
absolute priority, dating from Gray, 1840, known from the Permian of South Africa
and it was also used by Adams and Adams and Russia, and do not seem to be directly
(1858) and by Carpenter (1861) («Mycetop- related with modern types of Naiades of
idae"). the family Mutelidae. These may be dif-
Anodontitinae and Monocondylaeinae ferent branches of fresh-water mussels
were established by Modell in 1942. He evolved from marine ancestors which did
restricted the subfamily Anodontitinae to not survive.
the Anodontites, of the group crispatus- Viewed in their distribution in the
tenebricosus-clessini, and for the other southern hemisphere, the Mutelacea agree
species of the trapesialis groups, plus with that zoogeographical pattern indica-
live of a gondwanic originor, what Pilsbry iidae is still unknown. However, it is

(1911) called "Eogeic" fauna. In that "a- more probable that Bartlettia belongs to
sylum", to apply the term given by Suess to the Mutelacea^; close to, or included in
his original concept of Gondwana, evolved, the Mycetopodidae. Our personal obser-
according to Pilsbry, several typical fami- vations as well as those made by Carcelles
lies of continental moUusks. However, the (1940), Pain and Woodward (1961) and
group from which these naiades arose is Yonge young shells
(1962), indicate that the
unknown; probably Mutelacea and Union- of Bartlettia are not distorted as the adult,
acea are not derived from a common stock but very similar to (sometimes undis-
but, even if they were, the groups separated tinguishable from) Anodontites tenebri-
at a very early time. cosus. Adults of A. tenebricosus de-
Ihering emphasized the importance of veloping in crevices of rocky substratum
Pleiodon [=Iridina] pris cus as an African (as shells living on the west bank of La
element in the Brazilian Cretaceous, but Plata River commonly do) are strongly
the generic reference was questionable. distorted and acquire the characteristic
The single fragment of the fossil valve shape and aspect of Bartlettia. Also, the
figured by Ihering (1912), agrees more prismatic zone in both species is very si-
closely in umbonal and hinge characters milar and their distributions in southern
with Paxyodon (type P. ponderosus Schu- South America are somewhat coincident.
macher, 1817 = Mya syrmatophora Gron- It is likely that Bartlettia stefanensis,
ovius, 1891, according to clarification recorded by Carcelles from the Paraquay
and subsequent designation of Olsson and basin, and A. tenebricosus belong to one
Wurtz, 1951). The hinge in Paxyodon and the same species. Acostaea is re-
seems to vary with development to a stricted to the Magdalena River in Colom-
pseudotaxodont condition, which appears bia, and the fact that this species and
equally in Paxyodon ponderosus and Pseudomuelleria are mono-myarians is
Pleiodon priscus. Furthermore, Paxyodon due perhaps to convergent evolution from
belongs to Hyriidae and itsglochidiumhas their entirely sedentary life.
been studied by Bonetto (1959). Thus the
assumed relationship of Pleiodon priscus THE UNIONACEA (HYRIIDAE) OF SOUTH
with African Mutelidae is unsound. The AMERICA AND AUSTRALIA
strata in which P. priscus was found proba-
bly are younger than the indicated Bauru In 1896, Simpson related all the Aus-
Formation of the Upper Cretaceous. tralian forms of naiades to the genus Dip-
lodon, assuming migration via Antarctica
NOTE ON THE GENUS BARTLETTIA from South America. Ortmann (1912) con-
cluded that the Australian naiades belonged
The genus Bartlettia Adams, 1866 (type to the subfamily Hyriinae, but did not
Etheria stefanensis Moricand), of which establish any direct relationship among the
Rochanaia Morretes, 1941, is a synonym, genera. Iredale (1933/34) on tne contrary
currently is placed within the family emphasized the differences, creating for
Etheriidae which includes: Etheria from the Australian forms the family Pro-
Africa, Pseudomuelleria from India and pehyridellidae with four subfamilies: Vel-
Acostaea from South America. esunioninae without umbonal sculpture,
Acostaea and Pseudomuelleria are Lortiellinae with ridged umbos, Cucumer-
monomyarians in the adult stage (only unioninae with plicate umbos and large
very juvenile individuals have two shells, and Propeehyridellidae but within
muscles); Etheria a.nd Bartlettia are di- the Mutelidae (still including in this family
myarians. The larval form of the Ether- the two types of embryos). They suggest
^Modell (1949) tentatively placed Etheriinae as a subfamily of Mutelidae, and Bartlett-

iinae between Anodontitinae and Mycetopodinae.
TABLE 2. Comparison of South American and Australian Hyriidae (Unionacea)

filament mantle of larva
larval tooth
basal expansion
of tooth
sensitive
hair brushes
striation of
shell margin
larval shell
marginated area of shell
100 miera
FIG. 8. Glochidium of Velesunio ambiguus (Philippi). Bogan River, Australia.

Lateral view.
distal extension
of filament
curved tooth
tooth-base
expansion
Granulations of inner
margin
FIG. 9. Same as Figure 8. Internal view.
that the Australian naiades were derived Museum collection (Fig. 16d), one can see
from a basic stock of northern hemisphere that all these glochidia from the Australian
ancestors which migrated to southwestern region are entirely similar to those from
Asia in the Triassic; but a year later South American species of Diplodon, in
(1959: 243) McMichael and Iredale agreed outline, shape, insertion and structure of
that "an equally good case can be made the curved hooks.
for southern distribution across a temper-
ate antarctic land mass". We understand
this as referring to the Unionacea
(Hyriidae) since no real Mutelacea are
known from Australia.
Modell (1942, 1949), segregated many
groups on the basis of the umbonal struc-
ture and hinge. In Mutelidae, which he con-
sidered the most primitive, he included
Velesunioninae and Lortiellinae, and he
placed Cucumerunioninae in Margaritifer-
idae and Hyriinae and Propehyridellinae
in Unionidae. Also he indicated the origin
of Unionidae andMargaritiferidaeasIndo-
Pacific, whence the Hyridellinae invaded
Australia, from where they moved to South
America. This interpretation is incon-
sistent with the fossil evidence of Hyriidae
Summarizing our own observations, the
differences and similarities of the Union-
FIG. 10. Larval shell of Velesunio am-
acea (Hyriidae) in South America and
biguus (Philippi).
Australia, are outlined in Table 2.
Classification of the Unionacea of the The internal organization of the glo-
Southern Hemisphere is more complicated chidia of many Australian species are not
at lower taxonomic levels, especially since well known, but according to Percivalthey
it seems to involve phylogenetic and zoo- lack the larval filament and the sensitive
geographical problems. However, our data cirri present in the majority of Diplodon.
are sufficient to establish the close re- Hiscock (1951) andBonetto (1952) indicated
lationship between the forms of South A- the presence of such a filament in Veles-
merica and Australia. Their affinités are unio ambiguus (Fig. 8-10).
closer than could be expected from di- An important variation in Velesunio is
vergence from common Eurasiatic an- the basal expansion of the larval tooth
cestors, even granting an extraordinary over the free margin of the embryonic
evolutionary stability combined with a shell, and the presence of fine striae or
high degree of parallelism. The dif- crenulations along the same margin; its
ferences are few and it is possible to out- internal organization is, according to
line a lineage of Diplodon-Hyridella, sup- Bonetto, coincidental in general with Di-
ported by recent researches in the glo- plodon, although the larval filament, short-
chidia. er and hollow, shows two distal expansions
From Percival's description (1931) of absent in Diplodon.
the glochidium of DzpZodon lutulentus Gould Except for these differences of detail,
(= Hyridella menziesi Gray, according to the glochidial phase in Diplodon and the
McMichael and Hiscock), that of D. Australian forms show greater similarity
menziesi hochstetteri (Dunker) by the than that to be expected between Diplodon
same authors, and thela.rva.oiD. menziesi and other genera of South American Hyri-
from one specimen in the Carnegie idae, such as Castalia, Castalina and
100 miera
FIG. 11. Glochidium of Diplodon delodontus delodontvs (Lamarck) Parana River,

Argentina. (For nomenclature of the organs see Fig. 8).
FIG. 12. Shell of the glochidium of Diplodon delodontus delodontus (Lamarck).

Map 2. Fossil Hyriidae. Triassic Pennsylvania and Texas; 2, Paleocene southern

1,
Argentina (Patagonia); Paleocene southern Brasil; 3, Eocene Chile; 4, Miocene
2^,
NE. Argentina; 5, "Upper Tertiary" (probably Pliocene), Argentina; 6, Pliocene Peru,
Ecuador; 7, Pleistocene Buenos Aires. With exception of 2 (faxyodon) all other fossils
belong to Diplodon.
Callonaia. The number of similarities other fossil group is found in the Paleocene
basis for separation
in the glochidia is the and Eocene of South America (southern
of the family Hyriidae in the southern Argentina and Chile). All these fossils
hemisphere from all the other Unionacea. are generally smaller than most of the
recent species (hence comparable to the
FOSSIL HYRIIDAE hylaeus group), except for Eocene Chilean
forms that differ very little from the
The oldest known Diplodon are repre- living Diplodon patagónicas, a form which
sented by several species from the more resembles Australian species?
Triassic of Pennsylvania and Texas. An- From younger and different Tertiary
comparative study of the types and other materials of these fossil species, is the
subject of a paper now in preparation by the present authors.
MAP. 3. Distribution of Hyriidae in South America.

It
300 miera
^SE^ss^
FIG. 13. Left: Glochidium of Z)¿/)Zodon/5, /)/5
x2 10; right: Filament of the gill of the fish
malabaricus (Characidae) (common name "tararira") of the Parana River, showing the cysts pro-
duced by glochidia of Diplodon charruanus.
FIG. 14. A. Juvenile mussel of Diplodon (Diplodon) charruanus xl66. B. Juvenile mussel of Di/Jiodon
(fihipidodonta) burroughianus(= variabilis) showing the wide opening of the valves, xl95.
-
levels, other fossil Diplodon are known expansion was from West to East, (earlier
from Colombia to southern Argentina, and forms distributed along the Andes) and
several genera were proposed by Mar- later especially from southwest to north-
shall: Prodiplodon, Eodiplodon, Ecuador east between northern Patagonia and the
ea. Also Marshall's Antediplodon has rivers of the Parana system or their
subsequently been used for all the oldest equivalents at the time. This type of dis-
species, despite the fact that no clear distribution is opposite to that oftheMyceto-
tinction between this and other named podidae, which was from North to South
genera, or even with Diplodon itself, has in the East, without reaching the extreme
been established. Umbonal sculpture in southwestern areas long habited by Di-
Antediplodon is of the same type as that plodon.
found in living species of the patagonicus The rivers running across the Pampas-
granosus group, as well as in other fossils Chaco districts, tributaries of the Parana,
of the late Tertiary. The hinge of Pro- are saltier than the tributaries from the
diplodon singewaldi Marshall is similar to East, a factor which seems to have re-
that of D. patagonicus . Some Triassic stricted the dispersalof theHyriidae more
species such as borealis and pennsylvan- than that of the Mycetopodidae. Another
icus do not seem to agree with the type factor may be temperature: Diplodon is
species of Antediplodon {Unio dumblei found in some cold bodies of water in
Simpson). The forms from the Paleocene which the Mutelacea cannot live, but
of Patagonia, as well as Diplodon gard- systematized data are yet too poor to
nerae Marshall from the Pebas Formation reach conclusions.
in Peru, and the same type of Aníed¿/)Zoííon, Regarding the relationship of the recent
resemble the group of hylaeus. The di- South American and Australian Hyriidae,
vision of the fossil species into genera as indicative of an "Antarctic Way" of dis-
as age-groups does not improve our tax- persion, fossil evidence is lacking.
onomic knowledge and, if a vertical classi-
fication or the maintenance of such names THE GROUPS OF RECENT SOUTH
eventually becomes necessary, it has to AMERICAN HYRIIDAE
be done on a more consistent basis.
Pilsbry accepted the generic identifi- The Unionacea of South America - and
cation of the fossil species under D¿/)/oíion of Australia for the most part - form a
sensu lato, primarily from the only conspi- well defined family, Hyriidae, with shells
cuous character that these fossils show: radially sculptured at the inner laminae
the radially sculptured umbos, not present of the internal branchiae that are in contact
in other living or fossil North American with the palps, and parasitic larvae with
"Unios''. This character was considered S-shaped teeth either ending in spinulae or
as primitive by Ihering, Marshall, Modell strongly pointed, but always without ad-
and Pilsbry himself. Ortmann, without ditional denticulations, and without supra-
giving to such character enough phylo- anal aperture; nonparasitic glochidia may
genetic significance, when diagnosing Di- occur, without teeth, but in any case all
plodon, stated, however, that: "the beak the glochidia are perfectly distinguishable
sculpture is the most important feature of from those of the other families of Union-
the group". By the presence of radial acea from the northern hemisphere. The
sculpture in widely separated Triassic prismatic layer is reduced to a fraction
species, Pilsbry inferred (1921: 31) that of millimeter, inconspicuous, or entirely
North America once possesed a large and absent.
varied Naiad fauna of South American The family name Diplodontidae Ihering
type. 1901 (or Diplodont-inae Morretes 1949) is
Comparison of the distribution of Terti- not valid, being preoccupied by Diplodont-
ary and living hyriid species in South idae Dall 1899, created for marine bi-
America (Maps 2 and 3) shows that the valves. Prisodontini Modell 1942 included
lines of growth
valve
foot
rim of mantle
mantle branchial
filaments
adductor
anus
FIG. 15. Juvenile mussel of Diplodon {JRhipidodonta) variabilis (Maton). Ventral view.
the genus Hyria {=Prisodon) which cannot Tribe Diplodontini

be separated as a subfamily by itself. The
name Hyriidae Swainson 1840 has priority, Shell regular in shape, not alate, always
but Diplodontini and Prisodontini can be with radial ribs on the umbo, but of vari-
used as tribal denominations. able growth and posterior ridge scarcely
The typical subfamily, Hyriinae, hasthe developed, except in a few more elongated
radial ribs on the umbo coalescent toward and more posteriorly acute forms, as in
the center with very few exceptions, Diplodon parallelipipedon (Lea) or D.
branchial diaphragm imperforated, anal parodizi Bonetto. Branchial opening not
aperture forming a simple groove without entirely closed at the front.
expansions, and the branchial aperture Glochidium subtriangular-scalene, with
somewhat closed at the front. The glo- the teeth S-shaped, curved and ending in a
chidian tooth is triangular and not divided pair of spinulae (Fig. 16, b, e,); larval
at the end. The glochidium is with or with- filament long and rolled; with 2-4 sensi-
out larval filament, and the margin of the tive cirri. Species of direct development
embryonic shells lacks crenulations. have no teeth or hooks in the embryonic
The South American Hyriidae can be shell, but one, or several, marked bands
divided into the follwing tribes: of growth (Figs. 11-13).
FIG 16 Glochidia oiHyriidae. a, b, c, e, South American; d, Australian, a, Callonaia; ,

Diplodon solid-
Paxyodon alatus; d, Hyridella menziesi; e, Diplodon rotundus. All approximately x395.
ulus; c,
FIG. 17. Glochidium of Casíaízna psammo/ca (d'Orbigny) (for references see Fig. 8) . Parana River at
Santa Fe.
Genus Diplodon. the slow, intergrading, variations. Di-

plodon rhiLacoicus (d'Orb.), for example,
Tribe Prisodontini with a parasitic glochidium has been often
confused from shell similarities with D.
Shell subrhomboidal, bi-alate or álate charruanus (d'Orb.) whose glochidium is
only behind, but always with greater non-parasitic; on the same account, D.
posterior expansion. Umbonal sculpture charruanus is more closely related to
radial, very strong, with conspicuous the groups of D. hylaeus (d'Orb.) or D.
coalescence of the vertical riblets; rarely variabilis (Maton) despite the shell dif-
the sculpture may be inconspicuous. Pos- ferences.
terior ridge well marked. Branchial aper- Species with parasitic glochidia belong
ture as in Diplodontini. to Diplodon sensu stricto. In the post-
Glochidium triangular (isosceliform), larval stage the juvenile mussel shows
with teeth less curved and shorter than in the hooks still attached and a long, ciliated
Diplodontini, ending in 2-3 needle-like foot (Fig. 14) which soon disappears (Fig.
points (the glochidium was studied in Paxy- 15). The non-parasitic species are in-
odon alatus (Sowerby) (Fig. 16, c) but the cluded in the subgenus Rhipidodonta (type
internal organization is not yet completely species Diplodon variabilis (= paranensis,
known). burroughianus bulloides), of which Cycl-
,
Genera: Prisodon, Paxyodon. omya, Bulloideus, Ecvuidorea and Schles-

chiella are synonyms. In order to avoid
Tribe Castaliini the mistake of placing Rhipidodonta as
"nomen oblitum" (introduced in Article 23,
Shell subquadrangular, solid, umbos section b of the International Commission
elevated and umbonal cavity deep. Beak of Zoological Nomenclature Code of 1961),
sculpture of variable development, some- notice must be taken that, subsequently to
times very obsolete. Branchial opening its establishment by MSrch in 1853, itwas
becoming perfectly closed at the front. used by Adams and Adams, 1858, Fischer,
Glochidium subtriangular, equilateral 1887, and Thiele, 1935.
or isosceliform, with short, straight, tri- Ecuadorea Marshall 1932 was intro-
angular teeth, wide at the base but not duced for fossil forms of the very variable
divided at the end; cirri grouped in form group of D. hylaeus, which is within the
of brushes; without larval filament (Fig. subgenus Rhipidodonta.
16, a; 17). Schleschiella Modell, 1950 is an as-
Genera: Castalia, Cnstalina, Castal- semblage of unrelated species; its type,
iella, Callonaia. Diplodon burroughianus (Lea), is a syno-
nym of D. variabilis (Maton), used by
In the tribe Diplodontini the species pre- Mörch as type species of Rhipidodonta.
sent extraordinary ecological and indi- This species has non-parasitic larvae of
vidual variations, often repeated or mixed direct development, but Modell also in-
among the numerous local populations or cluded in Schleschiella (as a subspecies of
demes, but without taxonomic value. Sub- burroughianus) the form rhuacoicus, which
generic divisions of Diplodon have been actually has a parasitic glochidium, as
based on transitory shell characters only. well as D. parallelipipedon.
The most reliable separation is based Although the numerous species of Di-
primarily on the parasitic or non-para- plodon can be separated into minor
sitic condition of the larvae; secondarily "species-groups" for practical purposes,
groups of species may be recognized by only the groups listed below can be
shell characters, although this sometimes diagnosed by some definite character-
presents serious difficulties on account of istics.
Key to the genus Diplodon (Hyriidae)
la. With parasitic glochidia, Diplodon height as the internal. Marsupia

s.s 2 central or with posterior gravitation
lb. Glochidia non-parasitic, Subg. Rhipid- Group of D.(p.) rhuacoicus
odonta 3
3a. Size and shape variable, generally
2a. Shell elongated,compressed laterally more rounded. Sculpture less promi-
central costulae with marked con- nent and moderately convergent.
vergence and tendency to cross, Hinge teeth very variable in develope-
forming thick folds or nodules. Ex- ment Group of D. (.) variabilis
ternal branchiae higher than the in- 3b. Small but very solid shells. Strong
ternal and marsupia placed anteriorly sculpture extended toward the middle
Group of D. (D.) chilensis of the shell or beyond. Several
2b. Variable in length, diameter and alti- central convergent, chevron-like
tude. Sculpture less convergent, not costulae. Hinge teeth thick and strong.
crossed. External branchiae of same Group of D. (.) hylaeus
TABLE 3. COMPARAI
TABLE 4. SYNOPSIS OF PRESENT CLASSIFICATION
MUTELIDAE Africa
MUTELACEA
ANODONTITINAE
MYCETOPODIDAE -{
MYCETOPODINAE
MONOCONDYLAEINAE
. LEILINAE^
South
America
iPrisodontini
Castaliini
Diplodontini
HYRHDAE
VELESUNIONINAE, etc.
Australasia
L HYRIDELLINAE, etc.
UNIONACEA <
LAMPSILINAE
UNIONIDAE ANODONTINAE North
UNIONINAE America,
Eurasia,
Africa
^ MARGARITEFERIDAE
^See Appendix, page 206.

PRESENT CLASSIFICATION OF SOUTH AMERICAN NAIADES

AND RELATED AFRICAN FORMS
Superfamily UNIONACEA
Families UNIONIDAE Fleming 1828 (with several subfamilies in the northern

hemisphere) andMARGARITEFERIDAE Haas 1940^ (MargaritanidaeOrtmann
1910) are not included in the Neotropical region (see Table 4).
Family HYRIIDAESwainson(Hyria-nae) 1840; Herrmannsenl847
= Hyridinidae Carpenter 1861; Diplodontidae Ihering 1901 non

Dali 1899; Hyriinae Ortmann 1911.
Type genus: Prisodon Schumacher 1817. = Hyria Lamarck 1819

non Stephens 1829, Robineau 1863, Insecta; Gronovius 1763
-Meuschen 1778 nomen nudum; Hyria Blainville 1821.
Subfamily HYRIINAE Swainson 1840 (restricted South America) = Hyria-dae Agassiz

1847; Prisodontinae Modell 1942, Morretes 1949.
Tribe Prisodontini
Genus Prisodon Schumacher 1817. = Naia Swainson 1840; Harmandia Rochebrune

1881.
Type: by subsequent designation of Olsson and Wurtz 1951: P. obliquus

Schumacher.
Subgenus Triplodon Spix 1827

Type: T. rugosum Spix (= Hyria corrúgala Lamarck).
Subgenus Triquetrana ? Simpson 1900.

Type: Unio stevensi Lea.
Genus Paxyodon Schumacher 1817.

Type: P. ponderosus Schumacher = Mya syrmatophora Gronovius 1781.
Tribe Castaliini
Genus Castalia Lamarck 1819. = Tetraplodon Spix 1827.

Type: Castalia ambigua Lamarck non Sowerby = inflata d'Orb. ?
Genus Castalina Ihering 1891.

Type: C. m.artensi Ihering.
^The «Official List of Family-Group Names" [of the International Commission on

Zoological Nomenclature] London 1958, p. 57, establishes: "Margaritiferidae Haas
1940, Field Mus. Publ. (Zool) 24: 119, as validated under the Plenary Powers (type
genus: Margaritifera Schumacher 1816)" [emend, of Margartifera]. The name
Margaritiferidae was used previously by Henderson 1929; in 1936, however, Henderson
used Margaritiferinae as a subfamily of Unionidae.
Genus Callonaia Simpson 1900.

Type: duprei Simpson.
Genus Castaliella Simpson 1900.

Type: C. sulcata (Recluz)
Tribe Diplodontini
Genus Diplodon Spix 1827. = Iridea Swainson 1840.

Type: Diplodon ellipticum Spix.
Subgenus Rhipidodonta Mörch 1853. = Cyclomya Simpson 1900. Bulloideus

Simpson 1900. Ecuadorea Marshall 1932. Schleschiella Modell 1950.
Type: Unto variabilis Maton = paranensis + burroughianus Lea.
Genus Diplodontites^^ Marshall 1922.

Type: D. cookei Marshall
Australian subfamiles HYRIDELLINAE and VELESUNIONINAE are known to have a

glochidium larva; in Lortiellinae, Cucumerunioninae and Rectidentidae the larva is
unknown.
Superfamily MUTELACEA
Family MUTELIDAE Gray 1847 (restricted to Africa).
= Muteladae Conrad 1853; "Platiris I" group Lea; Iridinidae Bourguignat

1886; Pliodontidae Rochebrune.
Type genus: Mutela Scopoli 1777. =Spatha Lea 1838; Calliscapha Swainson
1840; Mutelina Bourguignat 1855; Pseudomutela Simpson 1900.
Family MYCETOPODIDAE Gray 1840 (restricted sensu Conrad 1853).
= Mycetopidae Carpenter 1861.
Type genus: Mycetopoda d'Orbigny 1835.
Subfamily MYCETOPODINAE Adams and Adams 1858 (Mycetop-inae).
Genus Mycetopoda d'Orbigny 1835 (JVIycetopus 1847).

Type: M. silicuosa (Spix).
Genus (?) Mycetopodella Marshall 1927.

Type: M. /a Zcaia (Higgins).
lOThe inclusion of this very little known genus, Diplodontites , within the Hyriinae is
only tentative. It has a prismatic layer like a mutelid, and other characters approach
Diplodon, but its embryology is unknown.
Subfamily MONOCONDYLAEINAE Modell 1942. = Monocondylae-idae Morretes 1949.
Type genus: Monocondylaea d'Orbigny 1835. = A/)Zodon Spix (non Rafinesque

1818). Spixioconcha Pilsbry 1893.
Type: M. paraguayana d'Orbigny.
Genus Haasica Strans 1932. = Marshalliella Haas 1931 (non Kieffer 1913, nec-
Poppius 1914). Iheringiella Pilsbry 1893. Plagiodon Lea 1856.
Type: Plagiodon balzani Ihering.
Genus Fossula Lea 1870

Type: Monocondylaea fossiculifera d'Orbigny.
Genus Tamsiella Haas 1931

Type: Monocondylaea tamsiana Dunker.
Subfamily ANODONTITINAE Modell 1942. = Glabariinae Modell 1942.
Type genus: Anodontites Bruguière 1792. = Patularia Swainson 1840;

Glabaris Gray 1847; Styganodon Martens 1900; Ruganodontites Marshall
1931; Pachyanodon Martens 1900.
Type: A. críspala Bruguière 1^;
Subgenus Lamphroscapha Swainson 1840. = Virgula Simpson 1900.

Type: A. ensiformis (Spix).
? Subfamily LEILINAE Morretes 1949 (See Appendix below).
Type genus Leila Gray 1840. = Columba Lea 1833 (non Linnaeus 1758).
Type: Anodonta blainvilleana Lea,
APPENDIX series of 6 or 7 parallel dorsal scars which

are not present in other Mycetopodidae,
Newobservations made on Leila blain- but represented in Mutela by a single one.
villeans (Lea) revealed a clearer dis- The prismatic layer is practically absent.
tinction from Anodontites .Leila has a The lasidium of Leila is of a type
pair of well devloped contractile siphons closer to the larva of Mutela, of large size
formed by a separate fold of the mantle (three times larger than in Anodontites),
and not by fusion of the mantle edges, and with a long filament instead of a ribbon-
consequently a well marked palliai sinus. like organ of attachment, the ciliated lobes
The palps are low and elongated instead well separated and without cirri at the
of high and rounded a.s in Anodontites. The posterior end.
shell is more winged and gaping, with a All these characters seem to indicate
11 To this group, apparently, belongs Bartlettia stefanensis (Moricand). K, however,

the previously supposed differences are sustained by further research on the
Bartlettia-Acostaea group, then the name Bartlettiinae Modell 1942 should have
priority. (See note on page 190).
that the differences between Anodontites CARCELLES, A., lQAO,Bartlettia stefan-

and Leila are of an importance greater ensis (Moricand) en el Paraguay. Notas
than previously assumed. Modell (1942- Mus. La Plata, 5 (Zool. 40): 217-221,
49) included Leila with his Glabariinae pLl.
( = Anodontitinae in our scheme), but the ,
First record oiBartlet-
1942,
genus constitutes rather a subfamily by tia in Paraguay.
Nautilus, 55: 93-94.
itself, for which we have the name Leilinae CARPENTER, P. P., 1861, Lectures on
Morretes 1949; that author, however did Mollusca; or "shell-fish" and their
not indicate the reasons for the separation. allies. Prepared for the Smithsonian
Institution.
DOELLO JURADO, M., 1923, Nuevas
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S. Nat. Mus., 61(16): 1-9, vations on other species. Notulae
pl. 1-3. Naturae 239: 1-9.
, 1928, New fossil pearly fresh- ORTMANN, A. E., 1912, The anatomy of
water mussels from deposits on the the Hyridella australis {ha.ma.rck)
'iia.ja.d
Upper Amazon of Peru. Proc. U. S. (= Diplodon australis). Nautilus, 25:

Nat. Mus., 74(3): 1-7. 100-103.
, 1930, New fossil land and fresh- , 1921, Marsupium und Glo-
water moUusks from the Reynosa Form- chidium der südamerikanischen Mus-
ation of Texas. Proc. U. S. Nat. Mus., cheln aus der Unterfamilie der Hyriinae.
76(1): 1-6, pl. 1. Arch. f. MoU.-k., 53: 103-111.
1931,
,
Anodontites: a genus of ,1921, South American Naiades;
South and Central American and Mexi- ...Mem. Carnegie Mus., 8: 451-670,
can pearly fresh-water mussels. Proc. pl. 34-48.
U. S. Nat. Mus., 79(23): 1-16, pl. 1-2. PAIN, T. and WOODWARD, F. R., 1961,
, 1932, New fossil fresh-water A revision of freshwater mussels of the
family Etheriidae. J. Conchol., 25(1): 1900, Synopsis of the Naiades,

,
2-8. or pearly fresh-water mussels. Proc,

PERCIVAL, ., 1931, note on the life U. S. Nat. Mus., 22: 501-1044, pi. 18.
history of Diplodon lutulentus. Trans. , 1914, A descriptive catalogue
New Zealand Inst., 62: 86-91. of Naiades or pearly fresh-water
the
PILSBRY, H. A., 1911, Non-marine mussels. Detroit (Bryant Walker): i-
Mollusca of Patagonia. Rept. Princeton xi, 1-1540.
Univ. expeditions to Patagonia., 1896- SWAINSON, W., 1840, A treatise on mala-
1899, 3(5): 513-633, figs. 1-38, pis. 38- cology. London (Longman); i-vii, 1-419.
47 + 5. WHITE, A., 1897, Contribuçoes á
, 1921 (in WANNER, H. E.), Some paleontología do Brazil, 4: Molluscos
faunal remains from the Trias of York Cretáceos d'agua doce do grupo da
County, Pennsylvania. Proc. Acad. Nat. Bahia. Arq. Mus, Nac. Rio de Janeiro,
Sei. Philadelphia, 73: 30-36, pi. 2-3. 7: 231-245, pi. 26.
RICHARDS, H. G., 1944, Fossil mollusks WINDHAUSEN, A., 1931, Geología Argen-
from the Triassic of Pennsylvania, tina, ,
Peuser edit,, Buenos Aires.
Proc. Penn. Acad. Sei., 18: 69-72. WICHMANN, R., 1927, Sobre la facies
, 1948, Fossil mollusks from the lacustre senoniana de los estratos con
Triassic of Pennsylvania, Notulae Na- dinosaurios y su fauna. Bol. Acad. Nac.
turae, 206: 1-4, 1 pi. Cié., Córdoba, 30: 383-405, pl. 1-21.
SCHUMACHER, F., 1817, Essai d'un YONGE, M.,
1962, On Etheria el-
nouveau système des habitations des liptica Lam. and
the course of evolution,
vers testacés. Copenhague: 1-287, pi. including assumption of monomyar-
1-22. ianism, in the family Etheriidae (Bi-
SIMPSON, T., 1896, The classification valvia: Unionacea). Phil. Trans. Roy.
and geographical distribution of the Soc. London, Ser. ., 244(715):423-458.
pearly fresh-water mussels. Proc. U.
S. Nat. Mus., 18: 295-343, pi. 9.
ZUSAMMENFASSUNG
TAXONOMIE UND ZOOGEOGRAPHISCHE BEZIEHUNGEN DER SÜDAMERIKANISCHEN

NAIADEN (PELECYPODA: UNIONACEA UND MUTELACEA)
Fürdie südamerikanischen perlmuttrigen SUsswassermuscheln, welche früher alle

in die Superfamilie Unionacea eingereiht wurden, wird hier eine natürliche Klassifi-
kation vorgeschlagen. Die seit dem Ende des vorigen Jahrhunderts gebräuchlichen
Systeme, die nach hauptsächlich konchyliologischen und nur teilweise anatomischen
Richtlinien aufgestellt wurden, werden hier in Hinblick auf neuere embryologische und
phylogenetische Forschungen besprochen, insbesondre was die Struktur und Entwicklung
der verschiedenen Larventypen anbelangt. Untersuchungen der Autoren innerhalb der
letzten 10 Jahre haben die Existenz einer "Lasidium" -Larve bestätigt, die seit ihrer
erstmaligen Entdeckung durch Ihering im Jahre 1891 nie mehr beobachtet worden ist.
Diese Larve ist für die südamerikanischen Gattungen Anodontites Mycetopoda, Mono-
condylaea und Leila typisch. Gleichzeitig haben die Untersuchungen anderer Forscher
an afrikanischen Arten von Mutela gezeigt, dass diese eine Larve haben, die, wenn sie
auch nicht mit dem Lasidium identisch ist, ihm doch in den wesentlichen Strukturen sehr
gleicht. Ein vergleichendes Studium des Aufbaus und der Entwicklung dieser Larven
zeigt einen Rangunterschied von Familiengrad zwischen den Muteliden Afrikas und den
Mycetopodiden Südamerikas an. Die nahe Verwandschaft dieser beiden Familien und
ihre aussergewöhnlich auffällige Divergenz gegenüber denjenigen SUsswassermuscheln,
die durch die wohlbekannte "Glochidium" -Larve gekennzeichnet sind, ermöglichen
andrerseits die Aufstellung einer neuen Superfamilie MUTELACEA. Alle übrigen
SUsswassermuscheln mit Glochidien verbleiben in den UNIONACEA.
Die Mutelacea leben heute in der südlichen Hemisphäre, mit Ausnahme von Aus-
tralasien. Es ist unbekannt ob die südamerikanischen von den afrikanischen Formen
abgeleitet sind oder umgekehrt. Die anatomischen und embryologischen Unterschiede
zwischen den Mutelidae und den fortgeschritteneren Mycetopodidae scheinen auf eine
frühe Trennung hinzudeuten. Paläontologische Aufzeichnungen fehlen beinahe völlig.
In Nordamerika bezeichnete Pilsbry Fossilien aus den Triasformationen Pennsylvaniens
als Myceiopoda -ähnliche Muscheln; "Pleiodon priscus", von Diering aus der Kreide
Brasiliens beschrieben, ist nicht, wie angenommen, ein Mutelide, sondern ein Hyriide der
Gattung Pexyodon. Verschiedene Angaben über Anodontite s-^nliche Fossilien aus der
Kreide Bahias in Brasilien sind äusserst zweifelhaft.
Wir teilen die südamerikanischen Mutelacea, d.h. die Mycetopodidae, in 3 Unter-
familien ein: die Mycetopodinae, Anodontinae und Monocondylinae; eine weitere Unter-
familie, die Leilinae, kann man vielleicht ebenfalls auf Grund neuerer embryologischer
und anatomischer Erkenntnisse unterscheiden. Andere, in früheren Klassifikationen
aufgestellte Gruppen mit Rangstufe einer Unterfamilie lassen sich nicht aufrechter-
halten, da ihre Kennzeichen nicht genügend beständig und die Zwischenstufungen
zahlreich sind.
Die südamerikanischen Unionacea gehören der Familie Hyriidae an, welche auch in
Australien vorkommt, aber auf der übrigen Welt fehlt. Die ausschliesslich süda-
merikanischen Formen gehören der Unterfamilie Hyriinae an und werden in 3 Tribi
eingeteilt: Diplodontini, Castaliini und Prisodontini. Insbesondre die Diplodontini,
deren zahlreichste Vertreter der Gattung Dt/^Zocion angehören, sind näher mit den Formen
Australiens und Neuseelands verwandt. Auch hier blieben Versuche die Gattung in eine
Anzahl von Untergattungen zu zergliedern, infolge der Schwierigkeit beständige Merk-
male zu definieren, erfolglos. Vom embryologischen Standpunkt aus kann man aber 2
Einheiten von subgenerischem Wert unterscheiden: Diplodon s.S., mit parasitischen
Glochidien und Rhipidodonta mit nicht parasitischen Glochidien, d.h. solchen mit
direkter Entwicklung. Es gibt paläontologische Angaben über das Vorkommen von
Hyriiden im nordamerikanischen Trias, im südamerikanischen Paläozän und im Eozän
von Chile, wobei die letzteren Fossilien den heute in dieser Gegend lebenden Arten,
wie auch verwandten australischen Arten, sehr ähneln. Alle diese Fossilien gehören
zu der Gattung Diplodon, von welcher auch verschiedene andere Arten in jüngeren
Schichten aus dem mittleren und oberen Tertiär des südamerikanischen Kontinentes
vorkommen.
Die den sogenannten "Süsswasseraustern" augehörige monotypische Gattung SaríZeííí'a,
die man allgemeinerweise zu den Etheriiden rechnet, gehört walirscheinlich einer
polymorphen Art der Mutelacea, Anodontites tenebricosus an. Die larvären Stadien
der Etheriiden sind noch unbekannt. Höchstwahrscheinlich aber dürften weitere
Untersuchungen ergeben dass diese Familie, i'alls sie überhaupt als solche erhalten
bleibt, unter die Mutelacea einzureihen sein wird.
Tabellen werden hier gegeben, welche die verschiedenen seit 1900 gebräuchlichen
Klassifikationssysteme vergleichen und auch das hier angewandte System von den
Superfamilien bis zu den Untergattungen zeigen,
RESUME
TAXONOMIE ET RELATIONS ZOOGEOGRAPHIQUES DES NAIADES DE L'AMERIQUE

DU SUD (PELECYPODES: UNIONACEA ET MUTELACEA)
Un système naturel de classification est proposé pour les moules nacrées fluviátiles
qui étaient jusqu'à présent toutes rangées dans la superfamille Unionacea. Les systèmes
en usage depuis la fin du siècle dernier, basés principalement sur des caractères
conchyliogiques et partiellement sur des caractères anatomiques, sont ici discutés à
la lumière de recherches embryologiques et phylogénétiques récentes, spécialement en
ce qui concerne la structure et le développement des différents types larvaires.
Les recherches faites par les auteurs pendant les 10 dernières années ont confirmé
l'existence d'une larve, le lasidium', qui n'avait plus été observé depuis sa découverte
'
première par Ihering en 1891. Cette larve est typique pour les genres sudaméricains
Anodontites, Mycetopoda, Monocondylaea et Leila. Simultanément, des recherches
faites par d'autres auteurs sur certaines espèces de Mutela de l'Afrique y ont révélé
l'existence d'un type larvaire qui, s'il n'est pas exactement conforme au lasidium, en a
néanmoins les traits structuraux essentiels. Des études comparatives sur l'organisation
et le développement de ces larves permettent de formuler les différences entre les

mutélidés de l'Afrique et les mycétopodidês de l'Amérique du Sud. D'autre part, les
grandes affinités entre ces 2 familles ainsi que leur extraordinaire divergence embry-
ologique d'avec toutes les moules fluviátiles caractérisées par les larves si bien
connues du type "glochidium", permettent leur groupement dans une nouvelle super-
famille, les MUTELACEA. Toutes les autres moules sudaméricaines à glochidies
restent dans les UNIONACEA.
Les Mutelacea vivent exclusivement dans l'hémisphère austral, sauf en Australasie.
Nous ne sommes pas en mesure de juger si les groupes américains dérivent des
groupes africains, ou vice versa. Les différences anatomiques et embryologiques entre
les Mutelidae et les Mycetopodidae, plus évolués, paraissent indiquer une séparation
ancienne. Nous ne disposons que d'indices paléontologique's fort restreints: ils font
défaut pour l'Australie et l'Afrique; en Amérique du Nord Pilsbry attribue certains
fossiles du triassique de Pennsylvanie à un genre ressemblant à Mycetopoda tandis que
le "Pliodon priscus", décrit par Ihering du crétacé brésilien, n'est pas un Mutélidé
comme on le croyait, mais un Hyriidé du genre Paxyodon. Enfin, certaines références
à des fossiles du type de Anodontites, du Crétacé de Bahia au Brésil, sont fort douteu-
ses.
Les Mutelacea de l'Amérique du Sud, les Mycetopodidae, sont divisés en 3 sous-
familles: les Mycetopodinae, Anodontinae et Monocondylaeinae, Une sous-famille
additioneile, les Leilinae sera peut-être à introduire suivant les recherches récentes,
mais il n'est pas possible de maintenir d'autres groupes, indiqués dans les classifications
antérieures, au rang de sous-familles, car leurs distinctions anatomiques et conchyliolo-
giques sont insuffisamment tranchées.
Les Unionacea de l'Amérique du Sud appartiennent à la famille des Hyriidae, qui vit
aussi en Australie, mais est absente du reste du monde. Les formes exclusivement
sudaméricaines sont rangées dans la sous-famille Hyriinae et divisées en 3 tribus: les
Diplodontini, Castaliini et Prosodontini, Les Diplodontini en particulier, dont le groupe
le plus nombreux est formé par le genre Diplodon, sont les plus étroitement alliés aux
formes australiennes et néozélandaises. Tous les essais de distinguer divers sous-
genres ont échoué en raison de l'impossibilité d'en définir des caractères constants,
même approximativement.
Du point de vue embryologique, pourtant, nous pouvons distinguer 2 groupes de valeur
sous-générique: Diplodon s.S., à glochidies parasites et Rhipidodonta à glochidies
non-parasites, c'est à dire à développement direct. Nous disposons de renseigne-
ments paléontologiques sur les hyriidés: des fossiles ont été décrits du triassique
de l'Amérique du Nord, du paléocène de l'Argentine australe et de l'éocène du
Chili, ces derniers étant très proches des espèces qui vivent actuellement dans
la région, ainsi que des groupes apparentés de l'Australie, lous ces fossiles
appartiennent au genre Diplodon, dont on connaît nombre d'autres espèces de différentes
couches plus récentes des niveaux tertiaires moyens et supérieurs distribués de par
le continent sudaméricain.
Le genre monotypique Bartlettia, connu sous le nom de "huitres d'eau douce" et
couramment inclus dans les Etheridae, fait très probablement partie d'une espèce
polymorphe de Mutelacea, l'Anodontites tenebricosus Les stades larvaires des Ethér-
.
idés ne sont pas encore connus mais nous pensons que les recherches futures montre-
ront peut-être que cette famille, si elle est maintenue, se rangera parmi les Mutelacea.
Des tableaux comparant les différents systèmes de classification en usage depuis
1900 sont ici donnés, ainsi que celui ici adopté, allant du niveau de la superfamille à
celui du sous-genre.
RESUMEN
taxonomía y relaciones DE LAS NAIADES DE SUDAMERICA
El presente trabajo propone una clasificación natural de las almejas nacaríferas de
agua dulce sudamericanas que se agrupaban en la supe rf am ilia Unionacea. Los sistemas
conocidos desde fines del siglo pasado, basados principalmente en caracteres concho-
lógicos y en parte anatómicos, se discuten a la luz de recientes investigaciones embrio-
lógicas y filogenéticas, especialmente acerca de la estructura y desarrollo de los
diferentes tipos de larva que hasta ahora eran muy poco conocidos.
Investigaciones llevadas a cabo por los autores en los últimos diez años han con-
firmado la existencia de la larva "lasidio", que no había sido observada de nuevo desde
su descubrimiento por Iliering en 1891; esta larva es tipica de los géneros sudamericanos
Anodontites, Mycetopoda, Monocondylaea, Leila y afines, Al mismo tiempo, otros
estudios realizados en especies africanas de Mutela han revelado la existencia de un
tipo de larva que, si bien no es exactamente igual al lasidio, participa del mismo plan
de estructura. El estudio comparativo de la organización y desarrollo de estas larvas
permite la diferenciación de dos familias, Mutelidae y Mycetopodidae, en Africa y
Sudamérica respectivamente. Además, la estrecha relación entre esas dos familias
y el extradordinario contraste de su embriología frente a las otras almejas cuyo tipo
de larva es el bien conocido gloquidio, permiten agruparlas y distinguirlas en una
nueva superfamilia MUIELACEA, mientras que los restantes géneros y familias se
conservan en la superfamilia UNIONACEA.
Las Mutelacea actuales son exclusivas del hemisferio sur pero ausentes en Austra-
lasia. Un posible origen africano de los grupos sudamericanos, o viceversa, no ha
sido demostrado todavía. Las diferencias embriológicas y anatómicas entre las
Mutelidae y las más avanzadas Mycetopodidae parecen indicar una separación remota.
Las referencias a hallazgos fósiles de Mutelacea son raras y carecen de confirmación:
ninguna en África o Australasia; en Norte América se encontraron moldes, atribuidos
por Pilsbry a Mycetopoda. o un género similar, en el 1 riásico de Pennsylvania; "Plei-
odon priscos" descripto por Ihering del Cretáceo del Brasil, no es un mutélido como se
había creído sino que pertenece al género Paxyodon de los Hyriidae, Otras referencias
sobre almejas de tipo Anodontites del Cretáceo de Bahia, Brasil, son también muy
dudosas.
Las Mutelacea sudamericanas, Mycetopodidae, se dividen en tres subfamilias
fácilmente reconocibles: Mycetopodinae, Anodontitinae y Monocondylaeinae; otra sub-
familia, Leilinae, podria aceptarse basada en estudios más recientes. Ctros grupos
al nivel de subfamilia intentados por previas clasificaciones no pueden mantenerse,
por insuficiente caracterización y demasiada intergradación.
Las Unionacea sudamericanas pertenecen a la familia Hyriidae, viviente también
en Australasia pero ausente en otras partes del mundo. Aquellas que son exclusiva-
mente sudamericanas forman la subfamilia Hyriinae, dividida en tres tribus: Diplo-
dontini, Castaliini y Prisodontini; las Diplodontini, especialmente, están más relacion-
adas con las formas de Australia y Nueva Zelandia; el grupo más numeroso es el
género Diplodon, y, también aquí, previos intentos para distinguir subgéneros han
fallado por la dificultad en definir caracteres constantes. Desde el punto de vista
embriológico, sin embargo, se pueden distinguir dos grupos de valor subgenérico:
Diplodon s.S., con gloquidias parásitas, y Rhipidodonta^ con gloquidias no parásitas,
es decir con desarrollo directo.
Fósiles de Hyriidae, todos pertenecientes al género Diplodon^ han sido encontrados
en el T riásico de Norte América, (Pennsylvania y Texas), Paleoceno de Argentina
austral y Eoceno de Chile, estos últimos ya muy parecidos a las especies actuales de
la región así como a grupos actuales afines de Australia, Se conocen también otros
fósiles del mismo género de otros niveles terciarios en diferentes localidades sud-
americanas.
El género monotípico Bartlettia, de las llamadas "ostras de agua dulce", corriente-
mente incluido en la familia Etheriidae, muy probablemente pertenece a una especie
polimorfa de Mutelacea, Anodontites tenebricosus. Estadios larvales de estos y otros
Etheriidae son desconocidos, pero futuras investigaciones pueden comprobar que las
Etheriidae, si deben mantenerse como tales, pertenecen a las Mutelacea,
Se dan tablas comparativas de los diferentes sistemas de clasificación propuestos
desde 1900, así como se presenta el sistema ahora propuesto, desde el nivel de super-
familia hasta subgénero.
,,
,
Unionacea .
,
.. . .
, ----
,
.,
, . "",
, .
SOUTH AMERICAN NAIADES
Anodontites, Mycetopoda, Monocondylaea

- Leila.
213
20
1891
,
Mutela
.
- .
, Mutelidae Mycetopodidae -
"", -
,..."
, ; ,,
: - Mutelacea
MUTELACEA
-
-
UNIONACEA.
Mutelidae
:,, , , .
Mycetopoda; Pleiodon
', priscus,
Mycetopodinae,
,
-
,
, .-
.
-
Leiliinae
,
Paxiodon
-
-
-
-
(Hyriidae)
Mutelacea ,
Mycetopodidae , Anodotitinae
-
Mycetopodidae
.
Anodontites
Monocondylaeinae;
;
-
:,, Unionacea Hyriidae ,
. ,
,;:
Diplodontini
,.
, -
,-
-
--
.
Diplodotini , Castaliini
Diplodon
Prisodontini.
ss.,
Diplodon,
Hyriinae ,
.
Rhipidodonta,
.
Bartlettia,
Etheriidae
,
", - ,
Hyriidae
Diplodon,
,
",
-
. ,
Anodontites tenebricosus Mutelacea.
1900
CLINES IN THREE SPECIES OF LAMPSILIS
(PELECYPODA: UNIONIDAE)
Alan M. Cvancara
Museum of Paleontology, Ann Arbor, Michigan, U. S. A.
ABSTRACT
In relating the latitudinal position of three species of fresh-water mussels to shell

measurements, an attempt is made to demonstrate a geographic north-south cline.
Specimens used for this study range from northern Michigan southward to southern
Alabama. Three shell measurement ratios: height/total length, width/total length and
posterior length/total length, are plotted against the latitudinal position of specimens
for each of the three species: Lampsilis ventricosa (Barnes) (121 specimens), L. ovata
(Say) (79 specimens) and L. excavate (Lea) (60 specimens). Scatter plot diagrams indi-
cate a trend of gradual shifting of points to one side or other of the diagram depending
on the shell measurement ratio used. Diagrams of the shell ratios height/total length
and width/total length show an increase in the ratio number as lower latitudes are
approached. The posterior length/total length plot shows a decrease in the ratio number
with lower latitudes. Some support for a cline is given in that the ranges of the three
species overlap. The more northerly species, L. ventricosa and L. ovata, overlap in
their respective ranges, and the range of L. ovata merges with that of the southern
species, L. excavata. A statistical analysis of the data offers little to prove or disprove
a cline. The concept of a cline, as suggested, raises the implication of a change in the
taxonomy of the three species considered here. It is suggested that these nominal
species are probably subspecies. A definite judgment must depend on confirmative

studies based on soft part morphology and cross-fertility experiments.
INTRODUCTION other way, shells are more obese (swollen)

downstream in the larger rivers and less
A gradual change in some character over obese (compressed) in the headwaters.
large areas has been noted in many animal Ball (1922) statistically analyzed obesity
groups. This character gradient or cline and size of stream and arrived at the same
has even led some workers to formulate conclusions as Ortmann.
"rules", which, in a general way, can be This study is an analysis of a cline of
applied to express gradual geographic regional extent and is based on shell
variation (for summary of rules, see Mayr, measurements of river naiads which range
1942: 88-94). The purpose of this paper from northern Michigan southward to
is todemonstrate a regional cline in fresh- southern Alabama. The specimens used
water mussels (family Unionidae), which, are from an area bordered approximately
as far as the writer is aware, has not been on the east by the Applachian Mountains
done previously. and extending west to eastern Kansas.
Clines in moUusks, in general, are Three species of Lampsilis (subfamily
rather well-known (examples in Clench, Lampsilinae) were chosen for the study
1954: 122-125 and van der Schalle, 1948: because they are suspected of forming a
p. 26-30 and p. 57-60). However, clines closely related group: L. ventricosa
reported in naiads are few, and these may (Barnes), L. ovata (Say) and L. excavata
be termed ecoclines (Huxley, 1938, p. 219). (Lea) (Fig. 1). They are typically de-
Ortmann (1920: 269-312) found that in veloped, as follows:
certain mussels of the upper Ohio
drainage, obesity (length of shell divided Lampsilis ventricosa (Barnes); shell
by diameter or width) increases gradually subovate to subelliptical; beaks high and
in a downstream direction. Stated in an- sculptured with few coarse irregular
(215)
216 A. M. CVANCARA
50 mm.
_J
Lampstlts ventrt-
FIG. 1. Comparison of the left valves of three species of fresh-water mussels:
(UMMZ 86356), Wabash
cosa (Barnes), A (UMMZ 165196), Grand River, Michigan; L. ovata
(Say),
Indiana; L. excávala (Lea), (UMMZ 65656); Cahaba River, Alabama. All specimens are of approxi-
River,
mately the same age, each having 5+ annulae.
CLINES IN LAMPSILIS 217
ridges; posterior ridge low or wanting MATERIAL AND METHODS

(upper Mississippi drainage; in lati-
tudinal extent from about northern Mich- Shell measurements used in the present
igan to northern Illinois). study are (Fig. 2):
Lampsilis ovata (Say); like L.ventricosa Total length - greatest length measured
but with strong, well-developed pos- parallel to the hinge line.
terior ridge (Kentucky and Tennessee).
Posterior length - distance from pos-
Lampsilis excávala (Lea); with well- terior end of shell to point of beak,
developed posterior ridge but smaller parallel to hinge line and measured on
than L. ventricosa andL. ovata (Alabama interior of left valve.
River system).
FIG. 2. Left interior (left) and anterior (right) views of Lampsilis ventricosa (Barnes)
showing where shell measurements were taken.,
218 A. M. CVANCARA
Height - distance at right angles to hinge terior length/total length (Figs. 3, 4, 5).
line at highest part of umbo. Each point on the three grades represents
a single specimen. In some localities the
Width - distance across both valves, spread of several points on a given lati-
measured at same relative position as tude gives some idea of station variation.
height.
DISCUSSION
Another measurement, the distance can be seen that points on the scatter
It
between the outer edges of the adductor plot diagrams (Figs. 3, 4, 5) show a trend
muscle scars (DAS) measured parallel to of gradual shift to one side or the other
the hinge line, was also made on all speci- with a change in latitude, depending on the
mens. However, this measurement does measurement ratio used. The gradual shift
not appear significant, as is mentioned seems to indicate quite clearly a gradual
again in the Discussion. All measurements change in a measurable character, i.e., a
were made with vernier calipers. line. Additional support for the concept
Measurements from the shell interior is given in that the three species overlap
were obtained by fixing the shell to the in their geographic ranges. The range of
back of a framed plate glass by stiff rubber the northerly species, Lampsilis ventri-
bands. Desired distances were trans- cosa and L. ovata overlap, and the range of
ferred through the glass on paper with L. ovata merges with that of the southern
triangle and T-square and then measured species, L. excavata. The linal situation
with calipers. also seems to be reflected in certain physi-
Since all three species involved here are cal factors of the shell. As pointed out
sexually dimorphic, care was taken to earlier, L. ovata, as well as L. excavata,
measure specimens of the same sex, i.e., typically differ fromL. venin cosa in the
males. Male shells have a somewhat possession of a well-developed posterior
pointed posterior end, whereas the pos- ridge, which is low or completely lacking
terior end in females is bluntly truncated in the latter species. However, the de-
and the postero-ventral part is more in- velopment of this ridge in L. ventricosa
flated. To maintain relative size within and L. ovata is gradual and intermediate
and between all three species, only river in strength in certain areas where the two
forms were measured;lake forms are often species occur together. Another gra-
typically stunted. Also, in this study are dational character appears to be size. Al-
included only those specimens which have though no statistical proof has yet been
four annulae or more. All specimens compiled, there seems to be a gradual
measured are in the collections of the decrease in size within the three species
Mollusk Division, Museum of Zoology, from north to south.
University of Michigan. In a way, it seems rather surprising that
From the measurements of the 260 a gradual trend is indicated by shell
specimens {Lampsilis ventricosa, 121, L. measurements of specimens over an area
ovata, 79, L. excavate, 60), various shell of regional extent. One might expect local
ratios were computed: height/total length, environmental effects to "mask", more or
width/total length, posterior length/total less, any regional trend which might be
length, distance between outer edges of present. Some idea as to the effect of
adductor muscle scars (DAS)/total length, habitat on size, shape and other characters
and width/height (these data are on file in on mussel shells is given by Ball (1922:
the Mollusk Division, Museum of Zoology, 93-97, summary of observations by earlier
University of Michigan). These ratios were workers).
then plotted against the latitudinal position The shell ratio height/total length (Fig.
of each specimen. Of five ratio plots con- 3) seems to show the most uniform shift
structed, three are presented here: height/ of points, with an increase in the ratio
total length, width/total length and pos- number as lower latitudes are approached.
= -.576
• • •'
.A AJÂA
220 A. M. CVANCARA
• • • •
* I*
. ,:•. •••
34 -
This would indicate a decrease in total muscles be spaced farther apart for
to
length of shell relative to height with more effectivelyclosing the two valves, with
southerly formso The shell ratio width/ the reverse being true in shorter shells.
total length (Fig. 4) gives a similar in-
crease in ratio number with the approach STATISTICAL ANALYSIS OF DATA
of lower latitudes, indicating a gradual
decrease in total length relative to width. Regressions were computed for three
The points on this plot, however, appear shellmeasurement ratios on latitude, for
more widely scattered, indicating that each species separately, and also for all
width is a more variable measurement than three species taken together (Table 1).
height (Fig. 3). For the height/ total length and width/total
Part of this apparent variability may be length vdiiiosoi Lampsilis ventricosa there
due to using shells of rather variable age. is a significant correlation at the 95
Ball (1922; 118) pointed out that younger percent significance level, indicating a
shells are more obese (swollen) than older change of ratio number with latitude. For
ones in the species he considered. How- the posterior length/total length ratio of
ever, this does not seem to be the case L. venlricosa there is little, if any, corre-
with the three species considered here, at lation. Lampsilis ovala and L. excávala
least as can be determined from those show, from the correlation coefficients,
localities where several specimens were essentially no correlation, for each of the
measured. The width/total length plot ratios analyzed. Two interpretations can
also differs from the preceding one in that be given for the data of the latter two
the points representing Lampsilis excá- species: (1) there is actually no change in
vala are rather markedly "skewed" toward any of the ratio numbers with a change
the side of higher ratio numbers, and in latitude; (2) the latitudinal distribution
show a break from the general trend. The in each of the two species is too limited
reason for this difference, at present, is to indicate a change, although one may be
unknown. present.
Figure 5 shows a plot of the ratio pos- Regressions computed for the three
terior length which gives a
length/total species taken together produce relatively
general trend of lower ratio numbers with high correlation coefficients for all three
lower latitudes. These measurements re- shell measurement ratios, indicating a
flect a change in the position of the beak, rather significantly high correlation or
and they indicate that a more centrally change with latitude.
located beak occurs in the more southerly However, these data should be treated
specimens. This plot also appears to show with some reservation. A high correlation
that the points for Lampsilis excávala are coefficient could be obtained for three
slightly skewed, in this case toward lower populations widely separated latitudinally
ratio numbers. and in some specific measured character»
Two other shell measurement ratios That is, there could be plots of three
were analyzed, but they are not shown here clusters of points, separated along one
in graphic form; these were width/height axis by a difference in latitudinal range
and DAS/total length. The width/height and along the other axis by a difference in
plot was very similar to the width/total some measurable character. If these
length plot (Fig. 4) with points for L. ex- clusters were more or less offset, en
cávala skewed somewhat more markedly echelon, there would result a relatively
toward higher ratio numbers. A plot of high correlation coefficient and a re-
DAS/total length showed no apparent trend, gression line would pass through the three
a condition taken to mean that the position separated clusters of points. If there is
of the adductor muscle scars is not signi- a valid correlation for all species taken
ficant and is probably a mechanical factor. together, it would point out that sufficient
A long shell would require adductor latitudinal range is needed for the corre-
222 A. M. CVANCARA
TABLE 1. measurement ratios for Lampsilis ventricosa, L. ovata, and L.

Statistical data of three shell
excávala. Numbers written as exponents and subscripts to the correlation coefficient (r) in-
dicate the 95 percent confidence interval on r.
Species Correlation Arithmetic Standard

Number
and shell measurement ratios (X) coefficient Mean {x) Deviation
(r) (Sx)
Lampsilis ventricosa
-.179
Height/total length -.346 121 .6912 .03736
494
-.179
Width/total length -.379 .4453 .04731
522
-.092
Posterior length/total length +.088 .6732 .03384
+.262
Lampsilis ovata
•.173
Height/total length +.050 79 .7291 .03316
-.268
+.208
Width/total length .014 .4416 .01553
234
-.085
Posterior length/total length +.139 .6508 .02922
+.350
Lam,psilis excávala
.247
Height/total length +.007 60 .7533 .03501
+.260
+.208
266
+.190
Posterior length/total length .067 .6268 .02730
-.315
All 3 species of Lampsilis combined
-.488
Height/total length .576 260 .7170 .04380
-.652
-.466
-.635
+.359
Posterior length/total length +.458
+.549 .6557 .03609
lation to appear. It would then follow that are to be applied to this group. It has been
the significantcorrelation of Lampsilis a common experience to label northern
ventricosa for the height/total length and forms L. ventricosa; that is, those forms
width/total length ratios is merely additive found in the upper Mississippi River drain-
to the more extensive trend. It is con- age (in latitudes encompassing New York,
cluded that the statistical analysis offers northern Ohio, Michigan, Wisconsin, and
but little proof or disproof of the thesis northern Illinois). On the other hand,
presented here, specimens from Kentucky and Tennessee
would be considered "southern" and with
CONCLUSIONS their more centrally placed beaks and their
high and well-developed posterior ridges,
The attempt to analyze the clinal possi- these forms are identified as L. ovata.
bilities as they relate to three nominal But in such intermediate areas as southern
species of fresh-water mussels, Lamps- Ohio and Illinois intergrades are common
ilis ventricosa (Barnes), L. ovata (Say) and they have usually been named L. ovata
and L. excávala (Lea), stems from the ventricosa. As for L. excávala, it seems
practical problem one faces when names to have most of the characteristics of L.
ovata with respect to the more central may be helpful in future studies designed
position of the beaks and the prominent to understand better the interrelationships
high posterior ridge; it varies essentially in this complex.
in being consistently smaller in size than
L. ovata. In distribution L. excávala is ACKNOWLEDGEMENTS
part of the fauna of the Alabama River The writer expresses thanks to Dr.
system; its differences probably reflect Henry van der Schalle, MoUusk Division,
a change brought about since the time the Museum of Zoology, University of Mich-
Alabama and Tennessee drainages may igan, He suggested that this fresh-water
have been connected. mussel species complex might be suitable
Indication that a cline exists among for a clinal demonstration and placed the
Lampsilis ventricosa, L. ovata and L. collections of the MoUusk Division at the
excavata, based on shell measurement writer's disposal.
ratios, raises some doubt regarding the
taxonomic status of these three species. REFERENCES
One would naturally suggest that these
species could be represented by sub- BALL, G. H., 1922, Variation in Fresh-
specific taxa. If various possibilities
so, water mussels. Ecology, 3: 93-121»
come to mind; for example, the three CLENCH, W. J., 1954, The occurrence
groups could be treated as separate sub- of clines in molluscan populations (part
species of a single species complex. On of Symposium;
subspecies and clines,
the other hand, the L. ovata ventricosa 99-126, 133). Zool., 3: 122-125.
Syst.
complex is already recognized as quite HUXLEY, J. S., 1938, Clines: an auxiliary
intimately related so that it could be taxonomic principle. Nature, 142: 219-
separated subspecifically from the south- 220.
ern form, L. excavata. It is also possible M AYR, ERNST, 1942, Systematic s and
to use taxa of lesser rank, but this approach origin of species. Columbia Univ.
does not seem warranted here. It must Press, New York. 334 p.
be emphasized that the taxonomic charac- ORTMANN, A. E., 1920, Correlation of
ters used are only those of shell measure- shape and station in fresh-water
ment ratios and that only the shells of mussels (naiades). Proc» Amer. Philos.
these animals were examined. Much work Soc, 59: 268-312.
remains on the anatomy of the animals VAN DER SCHALIE, HENRY, 1948, The
and the ros s -fertility among these three land and fresh-water moUusks of Puerto
species has not been studied. The recog- Rico. Misc. Publ. Univ. Mich. Mus.
nition of the possibility that a cline exists Zool., No. 70. 134 p.
ZUSAMMENFASSUNG
GEFÄLLE IN DER MUSCHELGATTUNG LAMPSILIS (PELECYPODA: UNIONffiAE)
Es wurde versucht ein geographisches Nord-Süd Gefälle in der Süsswassermuschel

Lampsilis dadurch nachzuweisen, dass Schalenmessungen an 3 Arten dieser Gattung auf
deren Vorkommen in verschiedenen Breitegraden bezogen und graphisch dargestellt
wurden. Die 260 Exemplare auf denen diese Studie beruht stammen aus einem Gebiet,
dass sich vom nördlichen Michigan bis ins südliche Alabama erstreckt; es handelt sich
um L. ventricosa (Barnes) (121 Exemplare), L. ovata (Say) (79 Exemplare) und L.
excavata (Lea) (60 Exemplare). Die angewandten Verhältniszahlen betreffen folgenae
Masse der Schale: Höhe/Gesamtlänge, Breite/Gesamtlänge und Hintere Länge/Gesamt-
länge. Die eingetragenen zerstreuten Punkte neigen zur Verdichtung auf der einen oder
anderen Seite des Diagrammes, je nachdem um welche Messungen es sich handelt. Die
Werte der Verhältniszahlen von Höhe und Breite zur Gesamtlänge steigen, je südlicher
die Fundorte der Muscheln gelegen sind, während sie für diejenigen der hinteren Länge
224 A. M. CVANCARA
zur Gesamtlänge nach dem Süden hin fallen. Für das Bestehen eines Gefälles spricht,
dass die Verbreitungsgebiete der 3 Arten sich überschneiden; dies trifft für die beiden
nördlichen Arten L. ventricosa und L. ovata zu, und auch für die südliche Art L.
excávala, deren Verbreitungsgebiet in dasjenige von L. ovata übergeht. Ob ein solches
Gefälle nun besteht oder nicht, liess sich mittels der statistischen Analyse nicht
eindeutig beweisen. Dennoch liegt es nahe, dass diese 3 nominellen Arten nur Unter-
arten sind. Ein endgültiges Urteil wird jedoch nur auf Grund weiterer Studien über
die Anatomie der Weichteile und die wechselseitige Fruchtbarkeit gefällt werden
können.
RESUME
«CLINE» DE TROIS ESPECES DE LAMPSILIS (PELECYPODA: UNIONIDAE)
Nous avons essayé de mettre en évidence un "cline" géographique nord-sud pour 3

espèces de Lampsilis, en mettant en rapport les latitudes de leurs lieux de provenance
et les proportions de leurs coquilles. Ces rapports ont été représentés graphiquement.
Les 260 specimens utilisés dans cette étude appartiennent aux espèces L. ventricosa
(Barnes) (121 specimens), L. ovata (Say) (79 specimens) et L. excavata (Lea) (60 speci-
mens) et proviennent d'un territoire allant du Nord du Michigan jusqu'au sud de
l'Alabama. Les rapports hauteur/longueur totale, largeur/hauteur totale et longueur
postérieure/longueur totale furent mis en face des positions latitudinales. La dispersion
des points des diagrammes, montre une tendance graduelle vers un côté ou l'autre, selon
les mesures considérées: pour ceux qui concernent les rapports de la hauteur et de la
largeur à la longueur totale, l'on distingue une augmentation des valeurs en allant vers
le sud, tandis que pour les rapports de la longueur postérieure à la longueur totale,
les valeurs calculées diminuent en allant vers le sud. Le fait que les aires de réparti-
tion de ces 3 espèces se chevauchent donne prise à la suppesition de l'existence de ce
"cline": c'est le cas pour les 2 espèces septentrionales, L. ventricosa et L. ovata et
également pour l'espèce méridionale L. excavata dont l'aire se confond avec celle
de L. ovata . Quoique l'analyse statistique ne nous ait pas permis de décider défini-
tivement pour ou contre l'existence de ce "cline", il existe probablement et il se pour-
rait bien que ces 3 espèces ne soient que des sous-espèces. Un jugement définitif ne
sera possible qu'après confirmation par des études supplémentaires sur la morphologie
des parties molles et sur interfertilité de ces espèces.
RESUMEN
CLING EN TRES ESPECIES DE LAMPSILIS (PELECYPODA: UNIONIDAE)
Se utilizaron en este estudio 121 ejemplares áe Lamps His ventricosa (Barnes), 79 de

L. ovata (Say) y 60 de L. excavata (Lea). Esas tres especies se distribuyen desde el
norte de Michigan hasta el sur de Alabama. Se señalaron, en diagramas de correlación,
tres medidas proporcionales de la concha - las razones entre altura/longitud total,
diámetro/longitud total y longitud posterior/longitud total - en oposición a la situación
latitudinal de los ejemplares de cada especie. Se observó una tendencia a la dis-
locación gradual de los puntos para un u otro lado del diagrama, de conformidad con la
razón utilizada. Así, en la dirección de las bajas latitudes aumenta la frecuencia de
las razones altura/longitud total y diámetro/longitud total, y disminuye la frecuencia
de la razón longitud posterior/longitud total. Las especies de más al norte, L. ventri-
cosa y L. ovata, se sobreponen en sus respectivas distribuciones, y la distribución
de.L. ovata se continúa con aquella de la especie del sur, L. excavata. Estas relaciones
espaciales entre las tres especies favorecen la hipótesis de un clino geográfico de norte
a sur. Sin embargo, el análisis estadístico de los datos ofrece poco para probar o negar
esa hipótesis. El concepto de clino implicará en un cambio en la taxonomía de las
tres especies nominales consideradas. Se sugiere que ellas son probablemente sub-
especies. Un juicio definitivo depende de estudios confirmatorios basados en la morfo-
logía de las partes blandas y en experimentos sobre la fertilidad de los cruzamientos.
LAMPSILIS (BIVALVIA, UNIONmAE)
,. : .
-
), , ,
;
Lampsilis ventricosa (Barnes) - (121
L. excavata (Lea) - (60
),
). L. ovata
..,-
(Say) -
:-
(79
-
,. .
. -
-
- -
-
:,
L. ventricosa L. ovata L.
ovata L, excavata
. .
-
MUSSEL DISTRIBUTION IN RELATION TO FORMER STREAM CONFLUENCE
IN NORTHERN MICHIGAN, U. S. A.
by
Henry van der Schalie

Museum of Zoology, University of Michigan
ABSTRACT
It has been shown previously that certain Mississippi River naiades have invaded the
St. Lawrence drainage system in the region of the Fox River and Green Bay on the
western shore of Lake Michigan, Wisconsin, in post-glacial times when the Fox River
was still joined to the Wisconsin River, a tributary of the Mississippi. The fluviatile
species Alasmidonta marginata and Actinonaias carinata, for instance, now occur in
3 streams flowing into Green Bay, but not in the impounded waters of the Lake tself.
The present discontinuous distribution of these mussels was explained by a former en-
larged Fox River system with a dendritic river pattern that comprized these rivers
during a low water stage of Lake Michigan.
The southern species Alasmidonta marginata, Elliptio dilatatus ama Lasmigona costata
have been found in 3 northern Michigan rivers: the Millecoquin, in the eastern part of
the Upper Peninsula, and the Carp and Ocqueoc, in the northern part of the Lower Pen-
insula. Again we have a discontinuous pattern of distribution. Particularly A. marginata
is strictly fluviatile and the 2 other species also do not occur in Lake Michigan. Such
an extension of their range from the northwest can be accounted for best by geomorpho-
logical evidence. Glacial geologists have indicated that a former Mackinac River occu-
pied the bed of northern Lake Michigan during a post-glacial low water stage in
Chippewa -Stanley times, approximately 8500 years ago. The rivers concerned, now
tributaries of Lake Michigan and Lake Huron, must have been connected with the now
submerged Mackinac River system before the higher lake levels separated them.
An Atlantic species, Elliptio complanatus, has reached northern Michigan through
other post-glacial confluences, for the east. It has also been found in the Ocqueoc River,
although it is known to be absent from the Lower Peninsula, and in the Millecoquin of the
Upper peninsula.
The relations of the distribution of since glacial times, are also present in
fresh-water mussels to geomorphology the upper reaches of the more recent Great
have been discussed in several earlier Lakes - St. Lawrence hydrographical
accounts. Although this subject has been system, which drains to the east and which
somewhat controversial, facts to indicate has captured some of the headwaters of
that mussel distribution does serve as a the Mississippi system. The present di-
means for determining post-glacial vides lie in northeastern Wisconsin and
stream confluence have been demonstrated in northern Illinois and Indiana and the
in several regions. For the Great Lake whole of Michigan belongs to the St.
region of the Northern United States of Lawrence watershed.
America, which was formerly glaciated, Particulars on naiad distribution in this
the sequence of events that occurred in area and on its correlation with glacial
post-glacial history, as revealed by glacial history are found in the literature. Good-
geologists such as Leverett and Taylor rich and van der Schalie (1939: 40) have
(1915), Stanley (1938), Hough (1955, 1958) shown that certain Mississippi naiades
and others, can best explain certain entered rivers tributary to the Green Bay
features of present naiad distribution. of Lake Michigan, Wisconsin, at a low
Broadly speaking, it is apparent that water stage of the Lake during which there
some species typical for the Mississippi existed an enlarged Fox River System with
drainage, a system draining southwards a dendritic river pattern. Since these
(227)
228 H. VAN DER SCHAUE
species are, in part, strictly fluviatile, or for the successful completion of the life
at least do not inhabitLake Michigan, such histories that mussels must have with
a fromerly connected river system ap- their fish hosts.
pears to be an essential factor in ac- Knowledge of the mussel fauna in the
counting for the present occurrence of streams of the Upper Peninsula of Mich-
these species in streams that are now igan is all too incomplete. There are
discontinuous due to the present higher several reasons for a lack of such data. The
water level in Green Bay. Further perti- area is still a rather extensive wasteland,
nent data are given, for northeastern Wis- with very few roads providing access to
consin and northwestern Michigan, by its streams. Many of the rivers are not
Walker (1913), H. B. Baker (1922), Morri- favorable for mussels throughout their
son (1932) and van der Schalle (1939, 1945); entire course because the bottom often
for southwestern Michigan, by van der tends to be too sandy; mussels do not main-
Schalle (1941, 1945) and, for southeastern tain themselves on shoals with shifting
Michigan, by Walker (1913), Ortmann sands. It thus becomes very difficult to
(1924) and van der Schalle (1939). In determine which species live in a particu-
central New York, Clarke and Berg (1959) lar drainage system, since an accessible
have more recently correlated mussel stretch of a stream bottom may be un-
distribution with geologic history. productive while not far away in the same
With additional information now availa- stream mussels thrive on a firm bottom.
ble, it is possible to show that the influx With these conditions in mind, it is obvious
of the southern Mississippi mussel fauna that one cannot be certain that the infor-
into the Northern Peninsula of Michigan mation available is anything but fragment-
extended a considerable distance north- ary. However, the few reliable data
eastward beyond the Green Bay area than available, already do enable one to postu-
previously known, as far east as, and even late the conditions that must have existed
beyond the Mackinac Straits between Lakes during the late Pleistocene, when lakes
Michigan and Huron. The studies on the and rivers were connected in such a way
rivers of the Great Lakes, as published by as to permit the patterns of distributions
Stanley (1938) and Hough (1958) are here now observed among some of the mussels
shown to account for this influx. in northern Michigan. It should be stressed
In view of past misunderstanding as to that information of this kind ought to be
the value of mussel distribution as an aid obtained before the rivers are subjected
in determining stream confluence, it must to detrimental human influences that may
again be emphasized that fresh-water eradicate the original fauna.
mussels are an especially useful group While 8 species are involved in showing
for relating their patterns of distribution stream confluence in northern Michigan
to both the present and the past history and Wisconsin, the distribution patterns
of the streams in which they now live» of 2 river-inhabiting species, Actinona-
The controversial aspects of the subject ias carinata Bsivnes Sind Alas midonta mar-
have been discussed by Ortmann (1913) ginata Say, have already been stressed
and van der Schalle (1939, 1945). Passive by Goodrich and van der Schalle (1939:
means of distribution, other than fishes on 41-44, Map 2). They occur in north-
which the animals are normally parasites, eastern Wisconsin, in the Oconto, Peshtigo
do not serve to account for the patterns in and Menominee rivers (Fig. 1), but are
distribution now observed. Most attempts unable to maintain themselves in im-
to explain mussel dispersion by birds as pounded waters such as Green Bay and
passive agents are based on untenable bi- Lake Michigan itself. These species, now,
ological conditions and do not serve either occurring in 3 separate streams, clearly
to explain the definite inter-relation of have a discontinuous distribution pattern.
mussels with their environmental con- For a previous continuous river pattern
ditions or the intricate relations necessary that would explain this distribution, a
MUSSEL DISTRIBUTION AND STREAM CONFLUENCE 229
230 H. VAN DER SCHALIE
low -waterstage must have existed in the Carp and Ocqueoc Rivers in the Lower
Green Bay and Lake Michigan in former Peninsula (Fig. 1). Another, Atlantic
times. In fact, during the post-Algon- species, Elliptio complanatus (Dillwyn),
quin period, the streams in Wisconsin and was also found in 2 of these streams»
northwestern Michigan, now draining into Former confluences must have existed to
the lake, connected with the enlarged Fox account for the presence of such faunal
River, then flowing through the bed of elements in the northern Michigan
Green Bay, to form a continuous dendritic streams, which evidently once were part
pattern. These former confluences have of a Mackinac River drainage. It is with
been indicated in that previous account. the evidence of these confluences that we
The Mackinac River drainage Recent
. can now understand how the southern
studies by Hough (1955, 1958) using radio- mussels could be established farther north
carbon dating techniques confirm this low- and east than hitherto suspected. Each of
water stage in post -Algonquin and pre- the 3 drainages will be considered sepa-
Nipissing time; Hough places the water rately.
level as much as 350 feet belov/ the present The Millecoquin River . On August 23,
lake surface. More recently Hough (1963) 1941, Dr. Carl L. Hubbs collected three
clearly summarizes the glacial lake levels species of fresh-water mussels from that
as they occurred in succession in the Lake River» Among the eight specimens col-
Huron basin. His Fig. 7 shows the time lected were three Alasmidonta marginata
sequences of the several lake stages in Say, an location even farther to the north-
relation to sea level. He indicates that the east than the Wisconsin-Michigan border
Algonquin stage, with anageof 11,000B.P. and also far from its normal southern
(Before Present) and at an altitude of 605 range in the Lower Peninsula of Michigan.
feet above sea level, was followed by a Because this discovery was so unusual the
large drop in level (about 370 feet) in the Millecoquin River was re-visited on Oc-
Stanley stage to a low level of 190 feet tober 4 of the same year. During this
above sea level. This was followed by a survey the following species were col-
gradual rise so that about 4500 years later lected:
the Nipissing stage was attained (about
4500 B.P.) when the water level was again Unioninae:
605 feet above sea level. In earlier ac- Elliptio complanatus (Dillwyn)
counts the low level of lake water was *Elliptio dilatatus (Rafinesque)
simply referred to as the "Algonquin-
Nipissing" stage. These newer data pro- Anodontinae:
vide a useful tool for studies of stream con- Anodonta grandis Say
fluence. Anodontoides ferussacianus (Lea)
Hough's studies and those by Stanley *Lasmigona costata (Rafinesque)
(1938) also show that a submerged Macki- *Alasmidonta marginata Say
nac River channel (Fig. 1.) existed in
northern Lake Michigan. Three additional Lampsilinae:
river connections in that area can now be Lampsilis siliquoidea (Barnes)
indicated to account for the presence in Lampsilis ventricosa (Barnes)
recent time of 3 Mississippi naiades, the
river - inhabiting species, Alasmidonta The three species marked with asterisks
marginata Say, and 2 other species not in above list are characteristically
the
found in Lake Michigan, Elliptio dilatatus among the Mississippi fauna (van der
(Rafinesque) and Lasmigona costata Schalle, 1939: 39-45), and they indicate
(Rafinesque). The 3 rivers in which this former conditions not now present in the
southern, Mississippi River faunal group Great Lakes. While Elliptio dilatatus and
has been found are the Millecoquin River Lasmigona costata have not been reported
in the Upper Peninsula of Michigan and (Goodrich and van der Schalle, 1932) from
Lake Michigan, both species are known to found as far north as Houghton Lake,
be able to live in a lake environment (van Roscommon county, the source of the
der Schalle, 1938), particularly if the lake Muskegon river". This location is about
has a stream influence as brought about by 170 miles up Michigan's Lower Peninsula,
a strong inlet or outlet, or both. Since a region accessible to Mississippi mussels
Lake Michigan does not support a rich from the south, through the Des Plaines-
mussel fauna, both species actually have a lUinois and the Maumee routes, which
discontinuous pattern; they probably were formerly connected with the Missis-
reached the Millecoquin with Alasmidonta sippi drainage.
marginata during a period of stream con- The Ocqueoc River drainage . This
fluence. lower Peninsula stream also flows to the
The presence of Elliptio complanatus north and drains into Lake Huron at Ham-
(Dillwyn) in the Upper Peninsula clearly mond Bay (Fig. 1). The following mussels,
indicates an invasion through confluence collected in 1948 by Harold W. Harry and
from the east, since this species belongs me, represent the recent mussel fauna in
to the North Atlantic (St. Lawrence River) Ocqueoc River:
mussel assemblage (van der Schalle, 1950:
449-450). The species probably traveled Unioninae:
westward as glochidia attached to fish, *ElUptio com.planatus (Dillwyn)
from Quebec through the Trent andNipis-
sing outlets into Georgian Bay, from there Anodontinae:
to spread, according to Walker (1913:45), Anodonta grandis Say
"along the north shore of Georgian Bay Alasmidonta calecolus (Lea)
[of Lake Huron] into the St. Mary's [River], *Alasmidonta marginata Say
and from thence into the eastern Lake Su- Lasmigona compressa (Lea)
perior, without getting either into Lake *Lasmigona costata (Rafinesque)
Erie, Lake St. Clair, or the lower part of Strophitus rugosus (Swainson)
Lake Huron". With regard to the period
of this spread, Clarke and Berg (1959) Lampilinae:
indicated that the Trent Outlet stage was Lampsilis siliquoidea (Barnes)
more likely than the Nipissing outlet stage
(which is the present valley of the Ottawa The species marked with an asterisk are
River). of special interest in that they would not
Carp River . This stream in the upper be anticipated as a part of the fauna of the
part of the Lower Peninsula flows north- river in this area while the 5 other species
wards into northern Lake Michigan (Fig. are normally found in northern rivers such
1), and there is one authentic record in as the Ocqueoc. These species have used
Museum of Zoology,
the collections of the two separate avenues of invasion, the
University of Michigan, to establish that Elliptio complanatus reaching the Ocqueoc
Alasmidonta marginata has been found in from the east, while Alasmidonta margin-
this stream. Royal Brunson collected this ata and Lasmigona costata definitely came
species on July 20, 1945, at the mouth of from the northwest.
Carp River, Emmet County (UMMZ The mussels of the Ocqueoc drainage
197969). The predominantly southern dis- have long been known to represent an un-
tribution of Alasmidonta marginata in usual assemblage in which northern and
Michigan has been recognized for more southern elements are found intermingling
than half a centry and is thus indicated by with remarkable discontinuity among
Bryant Walker (1898: 6) when he stated several faunal elements. The disparity
"Margaritina marginata Say [now known as can now be explained best in terms of the
Alasmidonta marginata] is apparently of submerged Mackinac River channel,
general distribution through the southern backed by means of additional evidence
part of the state [Michigan] and has been adduced by Stanley (1938) and by Hough
(1955, 1958), which shows that there was stantiated by the hundreds of distribution
a low-water stage known as "Lake Stan- records available in the Museum of Zoo-
ley" in the Huron basin. Hough (1955: logy at the University of Michigan. Mat-
967) stated: «Detailed study of the present teson (1948: 13, Fig. 1) used those records
lake -bottom topography should reveal to show distribution in Michigan; his map
many more effects of low stages of the clearly indicates its isolated occurrence
lake. Captain Vernon Seaman of the rein the Ocqueoc region. Ortmann (1924:
search vessel Cisco reports (Personal 116) suggested that the few scattered out-
communication, 1951) that fishermenhave lying records of Elliptio complanatus in-
brought up in their nets, from a depth of 85 dicated that it might be expanding its
feet, off Hammond Bay, Lake Huron, whole range southward at the present time.
trees complete with their roots and bearing However, this extension has not been ob-
no man-made scars. These trees, perhaps served to have taken place.
from a flooded forest, were identified as The faunal list for the Ocqueoc River
Tamaracks". This information tends to does not include Elliptio dilatatus (Raf-
strengthen the belief that a level suf- inesque) although the Museum of Zoology
ficiently low must have existed to permit collections record it from some of the
the river confluence necessary for the other rivers in the northern part of the
introduction of the river species Alasmi- southern Peninsula such as: the Devil
donta marginata into the Ocqueoc drainage River and Thunder Bay River in Alpena
from the northwest; this low level could County; the Rifle River in Ogemaw County;
account for the interesting pattern ob- Sylvan and Crystal lakes in Leelanau
served in the distribution oiElliptio com- County; Platte River, Benzie County; and
planatus. The time when this low-water Betsie Creek, Grand Traverse County.
stage occurred has been established as It would be interesting to discern the
7850 ± 350, according to radiocarbon reasons for its absence in the Ocqueoc
dates assigned by Crane and Griffin (1960: and its presence in the Carp and Mille-
31) for vegetable matter from Grand coquin rivers.
Traverse Bay; the radiocarbon age of wood Alasmidonta marginata and Lasmigona
from two stumps in situ in Thompson's costata are both well established in the
Harbor in Lake Huron was 7250 ± 300 Ocqueoc drainage. While the former has
years (1961: 106). Farrand (unpublished not been recorded farther east along this
thesis, 1960) places the beginning of the northern route, there is evidence that Las-
Chippewa-Stanley period at 8500 years migona costata, clearly a Mississippi
ago which is some 2000 years earlier species, did travel far to the east with
than the date estimated by Hough (1958). records for the Erie Canal in New York
The occurrence of Elliptio complanatus (Clarke and Berg, 1959: 8-9) and the Rideau
in the Ocqueoc drainage has been known River at Ottawa, Ontario (LaRocque and
for many years. Why it lived nowhere Oughton, 1937: 152-53). This species
else in the southern peninsula has been seems to have migrated both through the
a matter open to conjecture. The records southern Greater Maumee system (in the
(Walker, 1898: 11; Matte son, 1948: 14) for basin of present Lake Erie) and the former
its occurrence in Macomb, Lenawee and Mackinac River, in the north, to establish
Monroe counties are dubious and uncon- itself far in the St. Lawrence drainage
firmed, since many years of collecting system,
have shown that this species is not a normal
inhabitant in the southern regions. Its pre- REFERENCES
valence in the Ocqueoc and its absence as
a regular element in the fauna of all BAKER, H. ., 1922, The Mollusca of
streams in the remainder of the lower Dickinson County, Michigan. Occ. Pap.
Michigan Peninsula have been well au- Mus. Zool., Univ. Mich., HI: 1-44,
thenticated and can be adequately sub- 1 map.
CLARKE, A. J., Jr. and BERG, C. O., water m\xsséi Elliptio complanatus {Dil-
1959, The Freshwater Mussels of Iwyn, 1817). Nautilus, 61: 127-132; 62:
Central New York. Cornell Univ., 13-17.
Memoir 367: 1-79. MORRISON, J. P. E., 1932, A report on
CRANE, H. R. and GRIFFIN, J. ., 1960, the Mollusca of the northeastern Wis-
University of Michigan radiocarbon consion lake district. Trans. Wise.
dates V. Amer. J. Sei. Radiocarb. Acad. Sei., Arts and Letters, 27:359-96.
Suppl., 2: 31-48. ORTMANN, A. E., 1913, The AUeghenian
1961,
,
University of Michigan divide, and its influence upon the fresh-
radiocarbon dates VI. Amer. J. Sei. water fauna. Proc. Amer» Philos. Soc.,
Radiocarb. Suppl., 3: 105-125. 52: 287-390, pi. 12-14.
FARRAND, William R., 1960, Former , 1924, Distributional features of
shorelines in western and northern Lake naiades in tributaries of Lake Erie.
Superior basin. Unpubl. Ph.D. Thesis^, Amer. Midi. Nat., 9: 101-117.
Univ. Mich., 1-lv, 226 p. STANLEY, George, 1938, The submerged
GOODRICH, Calvin and VAN DER valley through Mackinac Straits. J.
SCHALIE, Henry, 1932, II. The naiad Geol. 46: 966-74.
species of the Great Lakes. Occ. Pap. VAN DER SCHALIE, Henry, 1938, The
Mus. Zool., Mich., 238: 8-14.
Univ. naiad fauna of the Huron River in south-
, Aquatic mollusks of the
1939, eastern Michigan. Misc. Pap. Mus,
Upper Peninsula of Michigan. Misc. ZooL, Univ. of Mich., 40: 1-83.
Pub. Mus. Zool., Univ. Mich., 43: 1-45. , 1939, Distributional studies
HOUGH, Jack L., 1955, Lake Chippewa, of the naiades as related to geomor-
a low stage of Lake Michigan indicated phology. J. Geomorph., 2: 251-257.
by bottom sediments. Bull. Geol. Soc. , 1941, Zoogeography of naiades
Amer., 66: 957-968. in the Grand and Muskegon rivers of
1958,
,
Geology of the Great Michigan as related to glacial history.
Lakes. Univ. Illinois Press, Urbana, Pap. Mich. Acad. Sei., Arts and Letters,
i-xviii, 1-313. 26: 297-310.
, 1963, The Prehistoric Great 1945,
,
The value of mussel
Lakes of North America. Amer. Sei., distribution in tracing stream conflu-
51: 84-109. ence. Pap. Mich. Acad. Sei., Arts and
LaROCQUE, Aurèle and OUGHTON, J., Letters, 30: 355-373.
1937, A preliminary account of the VAN DER SCHALIE, Henry and Annette,
Unionidae of Ontario. Canad. J. Res. 1950, The mussels of the Mississippi
D, 15: 147-155. River. Amer. Midi. Nat., 44: 448-66.
LEVERETT, Frank and TAYLOR, F. ., WALKER, Bryant, 1898, The distribution
1915, The Pleistocene of Indiana and of the Unionidae in Michigan. Detroit:
Michigan and the history of the Great privately printed by the author, 23 p.
Lakes. U. S. Geol. Surv. Monogr., 53: PI. 1-3.
1-529, pi. 1-32. , 1913, The Unione fauna of the
MATTESON, MaxR., 1948, Thetaxonomic Great Lakes. Nautilus, 27: 18-23; 29-
and distributional history of the fresh- 34; 40-47; 56-59.
Ôbtainable through University Microfilms Inc., Ann Arbor, Mich.

ZUSAMMENFASSUNG
DER EINFLUSS EHEMALIGER FLUSSVERBINDUNGEN AUF DIE
VERBREITUNG EINIGER MUSCHELN NÖRDLICHEN MICHIGAN, V. S, A.
Ich habe schon früher darauf hingewiesen, dass gewisse Naiaden des Mississippi-
flusses das St. Lorenz Entwässerungsbecken nach der Eiszeit in der Nähe des Fox-
flusses und der Green-Bucht, am westlichen Ufer des Michigansees, in Wisconsin,
besiedelt haben, zu einer Zeit als der Foxfluss noch mit dem Wisconsinfluss, einem
Nebenfluss des Mississippi, in Verbindung stand. Z.B. kommen die Flussmuscheln
Actinonaias carinata und Alasmidonta marginata jetzt in 3 verschiedenen Neben-
flüssen der "Green Bay", nicht aber in den Stauwässern des Sees selbst vor. Die
gegenwärtige diskontinuierliche Verbreitung dieser Arten habe ich im Zusammenhang
damit erklärt, dass der Foxfluss während einer liefwasserperiode des Michigansees
ein ausgedehnteres, verzweigtes System bildete, dem diese Flüsse angenörten.
Die südlichen Muscheln Alasmidonta marginata, Elliptio dilatatus und Lasmigona
costata wurden nun auch in 3 Flüssen des nördlichen Micnigan gefunden: im Millecoquin,
der im östlichen Teil der Oberen Halbinsel gelegen ist, und in den Carp und Ocqueoc
Flüssen, die im nördlichen Teil der Unteren Halbinsel liegen, wobei zu beachten ist,
dass insbesondre A. marginata eine ausgesprochen fluviatile Art ist und die beiden
anderen Arten zumindest im See nicht angetroffen werden. Diese Funde zeigen, dass
es den Mississippiarten gelungen ist, vom Nordwesten her bedeutend weiter in das
St. Lorenz Abzugsbecken einzudringen als bisher angenommen wurde. Wieder sehen
wir ein zersplittertes Verbreitungsbild, das am besten durch die inzwischen bekannt
gewordene geologische Vorgeschichte der Gegend erklärt werden kann. Während einer
post-glazialen Tiefwasserperiode in der Chippewa-Stanley Zeit, vor ungefähr 8500
Jahren, durchströmte ein ehemaliger Mackinacfluss das Bett des nördlichen Michigan-
sees. Die erwähnten Flüsse, welche sich jetzt in die Michigan- und Huronseen ergiessen,
gehörten zweifelsohne dem Mackinacsystem an, bevor die höheren Wasserspiegel in den
Seen ihre Verbindung unterbrachen.
Andrerseits gelangte eine atlantische Art, Elliptio complanatus, vom Osten her
durch andere ehemalige Flussverbindungen ins nördliche Michigan. Diese Art wurde
ebenfalls im Ocqueocfluss gefunden, obwohl sie bekanntermassen sonst in der südlichen
Halbinsel fehlt, und auch im Millecoquin der nördlichen Halbinsel.
RESUME
L'INFLUENCE DE JONCTIONS FLUVIÁTILES DU PASSÉ, SUR LA DISTRIBUTION DE

CERTAINES NAIADES DANS LE NORD DU MICHIGAN, E. U. A,
Nous avons déjà montré que certaines espèces naiades fluviátiles du Mississippi ont
envahi lebassin du St. Laurent dans la région du fleuve Fox et de la "Green Bay" sur
la rive occidentale du lac Michigan, au Wisconsin, après la glaciation, pendant que le
Fox était encore uni au fleuve Wisconsin, un affluent du Mississippi. Par exemple,
on trouve à présent les espèces Actinonaias carinata et Alasmidonta marginata dans
3 rivières tributaires de la baie Green mais pas dans les eaux du lac même. La
présente discontinuité dans la distribution de ces bivalves a été expliquée par l'existence
d'un système Fox plus important autrefois qu'aujourd'hui et à disposition dendritique,
qui englobait les 3 rivières actuelles pendant une période d'eaux basses du Lac Michigan.
Les espèces méridionales Alasmidonta marginata, Elliptio dilatatus et Lasmigona
costata ont été trouvées dans 3 rivières du nord du Michigan: le Millecoquin, situé
dans l'est de la Péninsule Supérieure, et le Carp et l'Ocquéoc, situes dans le nord de
la Péninsule Inférieure. Nous avous donc là aussi une distribution discontinue. Notons
que l'espèce A. marginata en particulier est strictement fluviatile et que les 2 autres
espèces aussi font dé faut dans le lac Michigan. La grande extension qui n'est ainsi
produite à partir du nord-ovest trouve sa meilleure explication dans l'histoire géo-
morphologique de la région. Les géologues du glaciaire ont indiqué récemment divulgee.
Qu'une ancienne rivière Mackinac occupait le lit du lac Michigan du nord pendant une
période post-glaciaine d'eaux basses, au temps du Chippewa-Stanley, il-y-a environ
8500 années. Sans doute les rivières en question, maintenant tributaires des lacs
Michigan et Huron, faisaient-elles partie de l'ancien système Mackinac et étaient-elles
alors en communication, avant que les niveaux plus élevés des lacs les séparent.
D'autre part une espèce atlantique, Elliptio complanatus venant de l'est, a pénétré
,
jusqu'au nord du Michigan par d'autres confluences du passé. L'espèce, dont on

connaît l'absence de la Péninsule Inférieure en général, n'y a été trouvée que dans
l'Ocquéoc; et, dans la Péninsule Supérieure, elle a été trouvée dans le Millecoquin.
RESUMEN
DISTRIBUCIÓN DE ALMEJAS EN RELACIÓN CON ANTIGUAS CONFLUENCIAS

DE CORRIENTES EN EL NORTE DE MICHIGAN, U. S. A.
Ha
sido demostrado previamente que ciertas almejas del río Mississippi invadieron
el sistema de drenaje del San Lorenzo en la región del río Fox y Bahia Green en la
costa oeste del lago Michigan, Wisconsin, en tiempos post-glaciales, cuando el río
Fox todavía estaba unido al Wisconsin, tributario del Mississippi. Las especies
fluviales Alasmidonta marginata y Actinonaias carinata, por ejemplo, aparecen ahora
en tres cursos que desembocan en Bahia Green, pero no en las aguas embalsadas del
lago mismo. La presente distribución discontinua de estas almejas fué explicada por
un antiguo sistema más amplio y de configuración dendrítica del río Fox, que englobaba
esos ríos, durante un periodo de aguas bajas del lago Michigan.
Las especies meridionales Alasmidonta marginata, Elliptio dilatatus y Lasm,igona
costata se encontraron en 3 ríos del norte de Michigan: el Millecoquin en la parte
este de la Alta Península, y en los ríos Carp y Ocqueoc en la parte norte de la Baja
Península. Aquí, otra vez, tenemos un tipo de distribución discontinua. Particularmente
A. marginata es estrictamente fluvial y las otras dos especies tampoco se encuentran
en el lago Michigan, fal extensión de sus áreas desde el noroeste puede estimarse
mejor por evidencias geomorfológicas. Los glaciólogos han indicado que un antiguo
río Mackinac ocupaba el lecho del norte del lago Michigan durante una fase post-
glacial de aguas bajas en la época Chippewa-Stanley, hace aproximadamente 8.500 años.
Los ríos en cuestión, hoy tributarios de los lagos Michigan y Huron, deben haber estado
conectados con el sistema del río Mackinac, sumergido al presente, antes que el alto
nivel de las aguas lacustres los separaran.
Una especie del drenaje atlántico, Elliptio complanatus, alcanzó el norte de Michigan
desde el este, a través de otras confluencias post-glaciales. Ha sido encontrada
también en el río Ocqueoc, aunque se sabe que está ausente en la Baja Península, y
en el Millecoquin de la Alta Península.
, ...
, ,.
,, ,,, ,
,.
-
Alasmidonta
--
. marginata Actinonaias carinata,
. :, ,
.,,. - . -1,-
costata
., ,, , ,
.
- -
Alasmidonta
marginata
8500
marginata, Elliptio dilatatus Lasmigona
-
,.-
Elliptio
.
.
complanatus
-
-
FRESHWATER SNAILS OF THE SUBGENUS HINKLEYIA
(LYMNAEIDAE: STAGNICOLA) FROM THE WESTERN UNITED STATES^
D. W. Taylor^, H. J. Walter^, and J. B. Burch4
U. S. Geological Survey, Washington, D. C, U. S. A.
The Liberian Institute

The American Foundation For Tropical Medicine
Harbel, Liberia
Museum and Department of Zoology

University of Michigan
ABSTRACT
The subgenus Hinkleyia has heretofore included only Stagnicola caperata (Say), but is
herein increased by addition of S. montanensis (Baker) on various conchological and
anatomical criteria, and ofS. /)z7sbryz(Hemphill) on shell features only. New information
permits a revised diagnosis of Hinkleyia, which has several distinctive characters but
is worthy of only subgeneric rank.
Sixteen specimens of a preserved series of S. montanensis irom Idaho were dissected.
Anatomically they resemble S. caperata so closely that no distinguishing features could
be identified.
The two species can be distinguished by their shells. In shape, S. montanensis is
usually more narrowly elongate and S. caperata. more swollen, but these characters
overlap and surface texture and sculpture are the only consistently reliable criteria for
distinction. Relatively conspicuous raised spiral ridges of periostraciun are diagnostic
of S. caperata and a shiny surface without such sculpture but with spirally arranged
series of tiny crescents of S. montanensis.
The anatomical characters shared by these 2 species and typical American Stagnicola
and, as far as is known, exclusive to them, are: a well-developed vaginal sphincter
muscle; a large diverticulum appended to the proximal end of the upper prostate; a
relatively stout and comparatively short penis sheath; a relatively short, bilaterally
symmetrical penis having a muscular "knot" near its midlength; and a well-developed
series of clearly bicuspid lateral radular teeth.
In comparison with typical Stagnicola, Hinkleyia (i.e.,S. montanensis and S. caperata)
shows a somewhat different and less prominent development of the vaginal sphincter, a
notably stout vas deferens and a shorter penis that has a less prominent, somewhat more
distally placed "knot*. Of special note is the tapered portion of the penis that is distal
to the "knot", which is short and stout and much less distinctly demarcated from the
thick proximal portion than in Stagnicola proper. Both species also have an imusually
1 Contribution No. 1, Western American Freshwater Mollusks Program, Institute of Malacology.

9
Publication authorized by the Director, U. S. Geological Survey.
3work supported by grant E-952, National Institute of Allergy and Infectious Diseases, U. S. Public Health
Service. Present Address: Museum of Zoology, University of Michigan, Ann Arbor, Michigan, U. S. A.
4work supported by grant 2E-41, National Institute of Allergy and Infectious Diseases, U. S. Public Health
Service.
(237)
238 TAYLOR, WALTER AND BURCH
Short and thick penis sheath, a prostate of a particular conformation, and an uncommonly
robust relative development of the lower part of the duct of the spermatheca. Addition-
ally, in texture and pigmentation of their overall anatomy, they show similarities which,
further serve to set them apart from other sufficiently known lymnaeid species. Their
bodies are well pigmented and evidently their foot and tentacles are relatively slender
in life.
It was also found that S. montanensis had become sexually mature upon reaching a shell
length of 6 to 7 mm. It is conjectured that the unknown egg mass of this species will
prove to be similar to that of S. caperata, which among known masses is unique in the
relative thickness of the individual egg envelopes and the thin inconspicuous outer tunic
of the egg mass.
Anatomical evidence indicates that a species formerly assigned to Stagnicola under
the subgenus Nasonia is closely related to Fossaria and therefore not allied to S.
montanensis and S. caperata. A possible special affinity of S. arcit'ca with /^/
rather than the subgenus Stagnicola was considered but rejected on new anatomical data.
The haploid chromosome numbers of S. montanensis and S. caperata are 18; the
demonstrated diploid number of S. montanensisis36. These numbers are characteristic
of basommatophoran snails in general. Details of gametogenesis, including the mono-
centric nature of the chromosomes, seem to be the same as in other lymnaeids. During
spermatogenesis there are normally 6 spermatogonial divisions, followed by 2 meiotic
divisions. Therefore, spermatogonial cells occur in clusters of 2, 4, 8, 16 and 32,
primary spermatocytes in clusters of 64, secondary spermatocytes in clusters of 128
and spermatids and sperm in clusters of 256.
Of special cytological interest in the gametogenesis of both S. montanensis and S.
caperata, and not reported previously for other lymnaeid snails, are 5 to 7 large chroma-
tin bodies in each nucleus during early prophase of the first meiotic division of spermato-
genesis. These chromatin bodies are perhaps another character demonstrating a close
relationship between the two species.
S. montanensis has a wide distribution in the western United States in the eastern
Columbia River and northern Great Basin drainages, from western Montana and Utah
to southern Idaho and central Nevada. It has the unusual habitat for Lymnaeidae of
springs or clear mountain streams. S. caperata is found over most of northern North
America and includes the area of S. montanensis within its distribution. It is found in
such situations as irrigation ditches and muddy, seasonal waters where S. montanensis
does not occur. Descriptions of specific habitats and maps of geographic distribution
(living and fossil) of the two species provide data concerning non -morphological differ-
ences between the species. The observed geographic separation is due, in part, to
differences in écologie requirements, but is probably, in part, due to the geologic history
of the region. S. pilsbryi is known only from the original material from one locality in
western Utah, and no écologie data are available.
The common North American snail caperata. S. pilsbryi (Hemphill) is added

Stagnicola caperata (Say) has hitherto been to thisgroup on shell characters only.
the only species included in the subgenus Anatomical descriptions are by H. J.
Hinkleyia Baker, 1928. In this paper we Walter and chromosomal studies by J. B.
add S. montanensis (Baker) andS'. pilsbryi Burch. Taylor and Walter jointly are
(Hemphill) to the group. Both of these responsible for the discussion of classi-
species have been little known up to now, fication and for the revised description of
but extensive collections show thatS. mon- Stagnicola iflinkleyia) m,ontanensis{2âev),
tanensis is widespread in much of the Taylor is responsible for the discussion
western United States. In features of shell of variation, distribution and habitat, and
and anatomy it shows close ties with S. for notes on S. pilsbryi.
SUBGENUS HINKLEYIA (LYMNAEIDAE: STAGNICOLA) 239
CLASSIFICATION should be removed from Stagnicola.

Although Baker (1911, 1928) did not
The general systematic position of the discern certain of the distinguishing
Lymnaeidae as one of the higher Basom- characters of Stagnicola caperata, he
matophoran families has long been agreed nevertheless recognized it as an especially
upon. The North American species of the distinctive species, which he segregated
family have been most thoroughly studied by itself in the subgenus Hinkleyia. The
by F. C. Baker, whose two large works of anatomical characteristics of this species
1911 and 1928 established the modern have been described in detail (Walter,
groupings of Lymnaeidae. Some authors 1961) and are summarized elsewhere in
(such as Hubendick, 1951) include virtually this paper. From studyof several Ameri-
all living species in the single genus can species of Stagnicola s.s. and of S.
Lymnaea. Others, as Baker (1911, 1928) ißinkleyia) caperata and S. (H.) montan-
and Zilch (1959-60) recognize several ensis it is clear thatn^Ze>'a has several
genera. The fact that varying degrees of distinctive features, but is so closely simi-
relationship within this group can be lar to Stagnicola s.s. that a subgeneric
recognized, primarily on the basis of rank is appropriate.
genitalia and radula but to a certain extent
from shell sculpture, size of nuclear SYSTEMATIC DESCRIPTIONS AND
whorls, and other features, is a weighty FOSSIL AND RECENT DISTRIBUTION
objection to including all species in the
one genus Lymnaea, We believe that such Family Lymnaeidae
a classification obscures more than it Genus Stagnicola Jeffreys, 1830
reveals. Subgenus Hinkleyia Baker, 1928
The genus Stagnicola includes numerous
species of medium-sized to large Lymnae- Genitalia: Penis sheath and penis very
idae, all found north of the Equator and short and thick, terminal portion of penis
mostly north-temperate latitudes.
in the short and conical; vasdeferens very thick;
Most and two of the three
of the species, lower prostate flattened, weakly folded,
subgenera, are restricted to North Ameri- roughly D-shaped; vaginal sphincter well
ca. Baker (1928) divided Stagnicola into developed, somewhat eccentric, but not
the three subgenera Siog^mcoZa s.s.,Hinkl- ball-like.
eyia, and Nasonia. Although we are dealing Nuclear whorl relatively large,
Shell:
primarily with Hinkleyia, some discussion as and Nasonia. Shell
in Stagnicola s.s.
of the relationship of the other two sub- of small or medium size compared to other
genera is necessary for an appreciation of members of the family; spire longer than
the characters of Hinkleyia. aperture; columellar fold weak or absent.
The only species of the subgenus Nasonia Both axial and spiral sculpture well de-
that has been examined thoroughly is veloped.
Stagnicola (.) bulimoides 111{-
man). Despite the possession of bicuspid
Type (by original designation): Stagni-
cola caperata (Say).
lateral teeth and a relatively large nuclear Distribution: Most of North America,
whorl it is anatomically not a Stagnicola as far south as the central United States.
but much closer to Fossaria, another Hol- Middle Pliocene to Recent.
artic lymnaeid genus. This subspecies is S. caperata (Say) and S. montanensis
considered (Walter, 1961) to be a relative (Baker) are the only well-known species of
of Fossaria obrussa (Say), but in the the group. S. pilsbryi (Hemphill) is in-
present state of knowledge it remains un- cluded on the basis of shell characters
certain whether all of the group Nasonia alone.
Stagnicola (fíinkleyia) montanensis and regularly conical to convex and bullet-

(Baker) shaped. The surface of the shell is con-
spicuously shiny except when covered by
PL I, Figs. 1-17 dirt or algae. Over-all color is brownish
horn, without the dense red, purple, or
white markings seen in other Lymnaeidae.
Galba montanensis Baker, 1913, Nauti- The nuclear whorl is as in Stagnicola
lus 26: 115.
Lymnaea montanensis Baker: Walker, s.S. or Nasonia, larger than that of
1918, Mich. Univ. Mus. Zool. Mise. Publ. Fossaria. The apex of the shell is there-
6: 94. fore relatively blunt compared to a
Lymnaea äff. caperata Say: Taylor, Fossaria of the same size. The nuclear
1956,Wyoming Geol. Assoc. Eleventh
Ann. Field Conference, Jackson Hole,
whorl may have the same intensity of
1956, Guidebook: 123, Fig. 3. Love, 1956, color as the rest of the shell, or may have
Wyoming Geol. Assoc. Eleventh Ann. Field a deeper, reddish-brown color.
Conference, Jackson Hole, 1956, Guide-
book: 92. Love, 1956, Am. Assoc. Petrole- Larger shells have 5-6 whorls, evenly
um Geologists Bull., 40: 1910. convex or slightly flattened on the sides
and with a slight shoulder just below the
suture. The suture is slightly impressed,
Shell of Stagnicola montanensis
and below the periphery of the preceding
whorl. It is bordered by a white or light-
Diagnosis - A species oí Hink le yia with
evenly convex whorls, regularly tapering colored band created by thicker shell de-
spire, and glossy shell surface. Spiral
posited at the junction of the whorls. Often
a narrow line of darker (reddish-brown)
sculpture, when present, consists of ir-
regular series of short spirally arranged color borders the white band on its lower
arcs. Axial sculpture of fine to very fine edge.
growth lines. The body whorl is about 2/3 of total
Types - UMMZ 76196, a series of 35 shell length.The aperture is relatively
shells. Baker (1913) specified "Types in small (see measurements. Tables 1-5)
collection of L. E. Daniels," We restrict and regularly elongate -oval, except for
the type lot to this series which passed flattening on the columellar wall and an
from Daniels through Bryant Walker and arcuate excision by the parietal wall. The
retains the original label in Daniel's hand- outer lip is thin and simple, in profile
writing. The type locality is "Hayes Creek, slightly oblique. The columellar border
near Ward, Montana, in the Bitter Root of the aperture is straight or weakly
Mountains, altitude 3825 feet," L. E. curved. A columellar fold is often absent,
Daniels station 13, collected April 25, and when present is low and poorly defined.
1912. The town of Ward has not been The parietal lip is thin, appressed to the
traced, but the U. S. Geological Survey preceding whorl, and passes smoothly into
Hamilton quadrangle (1910) 1:125000 the columellar lip with no evident line of
shows a Hays Creek about 3 miles south demarcation at the mid-length of the aper-
of Ward Mountain. The creek flows east- ture. The columellar lip is broadly re-
ward from the Bitter Root Mountains, flected and leaves an inconspicuous but
entering the Bitter Root River in the NW definite umbilical perforation.
corner of sec. 1 1, T. 4 N., R. 21 W. Daniels Axial sculpture consists only of weak,
probably collected in about the S 1/2 sec. growth lines, slightly oblique to the axis.
3, T. 4 N., R. 21 W., judged by the ele- They may be slightly protractive at the
vation he gave. The locality is in the suture. Spiral sculpture varies from weak
Ravalli County, Montana. and nearly absent to moderately conspicu-
Description - The shell is narrowly ous at lOx magnification. It consists of
conical, about 6-13 mm
long in adults. spirally arranged seriesof tiny crescentic
The outline of the spire varies from flat ridges and depressions whose ends point
proximally. These series of crescents are conical S. caperata.

variably distinct, but generally weak, and Driggs, Teton County, Idaho (Table 2):
vary considerably in strength on a given A series from the locality described on p
shell and between different shells. They 270 is of larger average size than most
tend to be irreguarly wavy and vary also in series of shells from elsewhere. It in-
width. Shell and periostracum between cludes one of the two largest specimens
these spiral series of crescents do not have (14.6 mm long) seen. The shells are glo-
incised lines or raised periostracal ridges bose, often indistinguishable from Stag-
as in Stagnicola caperata. nicola caperata in form, and almost com-
Varices and color bands are not present pletely lack spiral sculpture. This lot
in every lot. When they do occur they of smooth, globose shells is more like the
are usually on all the larger specimens series from north of Driggs and from the
in a lot, but are weak and inconspicuous type locality than like shells from other
compared to these features as usually localities.
developed in S. caperata. A shell 10 mm Cottonwood Creek, Twin Falls County,
long may have 3 or 4 varices. These are Idaho (Table 3): All the shells from south-
diffuse, with ill-defined edges, and opaque central Idaho are closely similar to one
in the center. Usually these varices are another. They have evenly convex whorls
located just proximal to an interruption which are more elongate than in many other
of the shell growth marked by a slight lots, or which may show a slight shoulder
descent of the suture and minor change in toward the suture. Theoutlineof the spire
the angle of inclination of the growth lines. is usually slightly convex. Often the tip of
Color bands almost always are associated the spire is etched or eroded away. In
with varices. They are dark reddish- this series from Cottonwood Creek 13
brown with diffuse edges, inconspicuous, specimens were measured which had un-
and next to a varix are immediately distal. eroded apices and a length of more than
Shells found empty usually have a wash of 8 mm. Eleven other specimens over 8
deeper color just inside the aperture even mm long were not measured because of
when no varix is present. eroded spires.
Measurements, variation in proportions Pleistocene, locality 22328 (for data on
- The most conspicuous variation in Stag- this and other localities see Appendix p
nicola montanensis is in size and pro- 270), Bannock Co., Idaho (Table 4): Shells
portions. Ideally one would analyze this from localities 22328 and 22410 in Marsh
variation by biométrie analysis of selected Creek valley, Bannock County, Idaho, are
samples. Unfortunately the number of the smallest S. montanensis seen. Except
specimens available is inadequate for a for size they are like other samples, for
consistent treatment, in this way, of the surface sculpture and proportions (see
samples representing all the observed Table 5) are not unusual.
variation. In order to document the range The measurements of Stagnicola
of size and proportions in the species parts montanensis which have been given in
of the larger representative samples (3 Tables 1-4 are an adequate indication of
recent and 1 fossil) have been measured the variation in the larger shells of these
as discussed below and shown in Tables samples, which encompass the range of
1-5. variation of the species. The measure-
Cottonwood Creek, Franklin County, ments cannot all be compared on the same
Idaho (Table 1): This sample includes one basis, however, because of the small
of the two largest specimens (14.6 mm numbers of specimens in certain size-
long) seen. It is from the locality de- classes. The more nearly comparable
scribed on p 270. The shells here are measurements have been summarized in
fairly narrow, turriform in many cases. Table 5.
On shape alone, many specimens could be Comparisons of shells - The relatively
distinguished from the more broadly large nuclear whorl and shiny surface
PLATE I
Cottonwood Cr., Franklin Co.
A Driggs, Teton Co.
Cottonwood Cr,, Twin Falls
% Loc. 22328, Co.
Width oí SheU (mm)
FIG. 1. Shell of Stagnicola montanensis, FIG. 2. Shell measurements of four

x7.5, from type locality, Hays Creek, samples montanensis Diagonal lines
of S. .
Ravalli County, Montana. USNM 570589. represent linear regression of shell width
Length 9.7 mm, width 5.7 mm; 5+ whorls. on shell length. The regression line
equation for the four samples are as fol-
lows: Cottonwood Creek, Twin Falls Co.,
Idaho, W= .12 + .48 L; Driggs, Teton Co.,
Idaho, W= 1.5 + .37 L; Cottonwood Cr.,
Franklin Co., Idaho, W= 1.51 + .30 L;
Pleistocene, Bannock Co., Idaho, W= 1.08 +
.34 L.
PLATE I
FIGS. 1, 2. Twin Falls Co., Idaho. Head of tributary of South Fork Shoshone Creek,
SW 1/4 SW 1/4 sec. 27, T. 14 S., R. 18 E. Length 7.6 mm, 4 3/4 whorls. USNM 633882.
FIG. 3. Same locality. Length 7.6 mm, spire eroded. USNM 633883a.
FIGS. 4, 5. Same locality. Length 7.4 mm, spire eroded. USNM 633883b.
FIGS. 6, 7. Twin Falls Co., Idaho. Cottonwood Creek near center of south edge of sec.
3, T. 15 S., R. 18 E. Length 7.8 mm, 4 3/4 whorls. USNM 633885a.
FIGS. 8, 9. Same locality. Length 9.3 mm, 5 whorls. USNM 633885b.

FIGS. 10, 11. Lincoln Co., Wyo. Pasture at Bedford. Length 10.4 mm, 5 1/2 whorls.
USNM 536403a.
FIGS. 12, 13. Lincoln Co., Wyo. Grassland at Grover. Length 8.7 mm, 6 whorls.
USNM 536407c.
FIGS. 14, 15. Same locality. Length 10.8 mm, 6 1/4 whorls. USNM 536407a.
FIGS. 16, 17. Same locality. Length 10.3 mm, 6 1/2 whorls. USNM 536407b.
TABLE 1. Measurements (in mm) of shells^ of Stagnicola montanensis from Cotton-

wood Creek, Franklin County, Idaho
Shell
TABLE 2. Measurements (in mm) of shells^ of Stagnicola montanensis from Driggs

Teton County, Idaho
Shell
TABLE 4. Measurements (in mm) of Pleistocene shells'' of Stagnicola montanensis

from locality 22328, Bannock County, Idaho
Shell
TABLE 5. Summary of measurements (in mm) of Stagnicola montanensis shells^ from

four localities in Idaho
parative data which were obtained in an uncoordinated way, causing some struc-
earlier study of many species of Lymnae- tures to be "relaxed" and causing others
idae; most of these data have been to be contracted. For example, in a series
recorded, only in an unpublished disser- of specimens, according to the individual,
tation (Walter, 1961), although a brief a greatly retracted penis may occur in a
report on the results of that study appeared greatly extended penis sheath, the penis
elsewhere (Walter, 1959). Here, only a may be retracted while the sheath is ex-
limited amount of the more essential data tended, both penis and sheath may be
can be given. The anatomical terminology retracted, or both may be extended.
used in this paper is tentative and was The vagina, to give another example of
partly devised by the author; it corre- such "artifact-variation", may be found
sponds to that employed in the dissertation to be greatly contracted and narrow, or
mentioned, where the terms are elucidated else greatly inflated, in individuals of one
by many detailed illustrations in an account series; when contracted, the organ may be
ginata serrata
was observed about
().
of the morphology of S. (Stagnicola) emar-
Much that
the morphology of S.
very firm and quite strongly muscular,
and when inñated, its walls may be ex-
tremely thin and delicate and may seem to
montanensis, but is not included in this be only very weakly muscular. Such vari-
paper, as well as some of what is described ation, superimposed on the genetically
here is essentially no different from what determined, intrapopulation and inter-
is given about S. e. serrato in that account. population anatomical variation, that
Special problems inherent in this kind seems to be of considerable magnitude
of study affect the interpretation of obser- in lymnaeids generally, makes it difficult
vations made, and as such, they deserve to judge objectively whether a particular
explanation. The narcotization-killing organ of a particular individual or in a
process when most successfully applied to particular population may "really" differ
a collection of snails en masse, commonly morphologically from that of another indi-
results in considerable contraction of the vidual or from that in another population.
tentacles of all of the specimens, while In taxonomic work on such snails at the
causing the sole of the foot of most indi- species-level, one must then seek for
viduals to expand, and especially to minor consistencies among gross incon-
broaden, beyond the dimensions that it sistencies, and accordingly, statements
normally would show in life when function- about real differences pertaining to such
ing in locomotion. Concomitantly, indi- structures are likely to be inextricably
vidualistic reaction of specimens to the involved with subjective impressions and
narcotization-killing process results in can be subject to extensive qualification.
various modifications of the form of the Miscellaneous characters and pigmen-
body and of internal organs. The organs tation of S. montanensis - The head-foot
as a whole are plastic and are easily mass and the tentacles of S. montanensis
distorted by tensions and pressures. are apparently particularly slender in life.
Muscular or glandular activity may cause In the best-relaxed specimens the
the organs to be altered grossly in their tentacles are mostly rather slender and
shape and dimensions during their normal pointed, despite being contracted, while
functions. Therefore, the "normal" or the sole of the foot is about twice as long
"intrinsic" form of an organ cannot be as it is wide, which is fairly narrow for
readily defined, and must be considered in that of a lymnaeid.
relation to particular postures of that In S. montanensis there is a groove
organ. Such an organ may assume a wide which runs along the peripheral border of
variety of normal postures and may react the mantle collar and this groove divides
to a "relaxing" chemical in a variable way. that border into two shelf-like structures,
Even in a well-relaxed snail the muscular the one above the other. A homologous
organs often seem to have reacted in an development occurs in at least some other
Lymnaeidae. In S. montanensis these form a reticulate pattern. In the larger

"shelves" are readily noticeable on the individuals the ovals tend to be obliterated
whole, and in certain individuals the lower by broadening of the dark zones. From
one projects distinctly and even conspicu- the lung region the heavy pigmentation
ously beyond the other. The "shelves" in continues posteriorly onto the coiled, sac-
other Lymnaeidae are usually not nearly like part of the mantle (tunica propria)
so well developed. that encloses the earlier whorls, but it is
The renal organ of several specimens of confined to the posterior (spireward) half
S. montanensis was examined and it was of the outer face of each coil. These dark
observed to be quite slender for a areas are irregularly but sharply de-
lymnaeid. Running along the underside of limited from the more anterior parts of
that organ there were two well-developed the whorls, which are very pale. Within
and close-set parallel muscular bands, the dark areas, the pigment forms a
which constitute "the renal muscle" mottled dusky-gray to blackish pattern
(Hubendick, 1951). The 3 jaws were bright which in young specimens is clearly seen
brownish-orange. The dorsal jaw was to be made up of numerous tiny dark spots
noted to be unusually narrow and rather and dashes, while a greater concentration
asymmetrical; its free margin, which ex- of pigment in these areas forms a narrow
hibited no cusps, was broadly excavated black zone, interrupted by light oval spots,
in the center, while its dorsal margin of that runs along the posterior edge of each
attachment was broadly arched. whorl. The mantle collar anterior to the
The S. montanensis studied were more lung is gray owing to a subsurface suf-
darkly colored overall than are most fusion with dusky pigment, and on it there
lymnaeids, and they showed an unusually are some irregular black to dusky-gray
prominent pattern of pigmentation in the small blotches with diffuse edges. Much
palliai areas. The pigment mostly occur- of the internal structure of the animal
red as minute dusky flecks which were, in exhibits a pronounced, minutely granular
part, diffusely distributed and which were pigmentation, even in immature speci-
partly concentrated in particular parts of mens. The arteries and the digestive
the body. The larger specimens tended to system are quite dark. Longitudinal dusky
be darkest. The head-foot region was dark bands are conspicuous along the length of
gray to medium gray due to a general sub- the esophagus and the two diverticula of
surface suffusion with the dusky flecks. the osphradium are flecked with black.
Immediately over the pneumostome, on Hermaphroditic reproductive organs -
the far right side of the mantle collar, there The ovotestis of S. montanensis is largely
is a large, diffuse, sub-surface smudge of invested by, and tightly bound to, a complex
light gray to blackish color; this smudge of tissues consisting of the liver, con-
occupies the area of the "hypobranchial nective fibers, and elements of the vascu-
gland". Above this "gland", over the right lar system. The exact shape of the gonad
side of the lung and against the inside shell- was not determined, but it was seen to
surf ace, there runs an intensely black band consist of a number of large lobes, made
parallel to the free edge of the mantle up of compactly arranged follicles. The
collar. The remainder of the surfaces follicles do not project from the surface
which lie against the inside of the shell in of the lobes and are so intimately bound
the region of the lung and kidney are coated together that they could not be separated.
with a mottled film of black pigment. This The surface of the lobes showed yellow
film is interrupted by light oval areas with spots and irregular rusty-orange flecking
diffuse edges. These ovals are much arranged in a vaguely reticulate pattern
larger, relatively, in the smaller speci- against a general creamy -white back-
mens, particularly in those about 4 mm ground. Granules and yellow ova, applied
in shell length, in which the pigmentation to the internal walls of the follicles,
is largely restricted to narrow zones that account for the pattern and sperm account
for the background color. The remaining uterine coil is the bulkiest organ in the
part of the hermaphrodite system, the complex and is especially mucoid. The
hermaphroditic duct, arises near the an- free uterine arm is tubular and elongate;
terior end of the gonad. The duct is a its juncture with the succeeding portion
long tube which proceeds cephalad into the of the tract, the neck of the oothecal gland,
visceral haemocoel, in which it partly is hidden by two structures appended to
lies free. In all specimens it showed two the tract at that point. These structures
grossly different portions. The first are the oothecal gland pouch and the
portion is somewhat the longer, and is muciparous (nidamental) gland. The pouch
broad and bulky, although it is very narrow is flattened, slightly broadened, and
at its origin on the anterior part of the roundly diamond- shaped or sometimes
gonad. It is considerably contorted, es- short and angularly truncate; it is a blind
pecially near its origin; its walls are thin reverse extension of the tubular neck of
and it contains a compact mass of sperm, the oothecal gland and it is rather small
which causes its bulkiness and lends it a and difficult to demonstrate. The muci-
creamy white color. The bulk of the parous gland is bulky, bulbous and roundly
sperm mass projects into and fills out angular in outline: it attaches broadly to
abundant crowded seminal vesicles, which an opening in the base of the pouch, and
swell out laterally from opposite sides partly to the end of the free uterine arm
of the broad central duct to form two which attaches at the same opening and
uniseriate rows. The arrangement of the which is tightly sandwiched between the
vesicles indicates a bilateral symmetry gland and the pouch. Numerous coarse and
although the vesicles are rather irregular compactly arranged glandular internal
in size and arrangement near the beginning lamellae make up the bulk of the muci-
of the duct. The second, distal segment of parous gland.
the hermaphroditic duct is a much narrow- The main body of the oothecal gland
er, slender, and mostly smooth cylindrical (oviducal bulb, birnenförmiger Körper,
tube, that tapers somewhat toward its first accessory albuminiparous gland of
terminus. authors) is a swollen portion of the female
Female genitalia - The female and male tract in which internal glandular lamellae,
genital systems arise at the terminus of largely diagonal to the tract, are greatly
the hermaphroditic duct, and firstly form developed. The first part of the gland, or
a compact complex of glandular organs its neck, is short and stout in comparison
massed around that point. The albumen with that in some other lymnaeids, being
gland and uterine complex of the female about half the length and half the diameter
system are the most conspicuous elements of the main body. The latter is broadly
of this mass. These structures, when spindle-shaped or nearly cylindrical and
separated, are seen to be much the same is small compared to that of some Lymnae-
as in other lymnaeids. However, they are idae. In a specimen 8.9 mm
in shell-
more compactly arranged, and more length the organ contained 11 to perhaps
mucoid and more swollen than in some 13 large lamellae, and possibly some
species. The uterine complex consists of smaller ones. The ventral raphe of the
an apical uterine pouch, a uterine coil, and organ appears to be broad, but was not
a free uterine arm, which makes up suc- clearly discerned. The last large part of
cessive portions of the tract. The albumen the female system is the vaginal tract,
gland, which is large and roundly tri- which is succeeded by the short vagina
angular in outline, lies against this proper leading to the female pore on the
complex and empties into the apical pouch right side of the animal. The diameter of
by a short small duct. A small fertili- the vaginal tract for its whole length is
zation pocket was assumed to be present about 3/4 that of the oothecal gland pre-
at the base of the albumen gland but it was ceding it. It is rather flattened dorso-
not demonstrated unequivocally. The ventrally. Its distal half is thick-walled,
highly muscular, and hardly glandular, flattened tube of a light color; it abruptly
as a result of progressive differentiation and greatly increases in diameter immedi-
from the preceding gland. The vaginal ately beyond itsvery narrow point of origin
tract rapidly tapers in giving way to the at the terminus of the hermaphroditic duct.
vagina proper and there the duct of the As it swells it gives rise to a prominent
spermatheca is appended to it. The duct lateral evagination, the prostate pouch.
of the spermatheca, which is somewhat The pouch contrasts with the prostate in
shorter than the combined length of the being darkly pigmented, and in being very
vagina and oothecal gland, passes from its thin -walled; it is curved and much flattened
origin on the right to the left side of the and wrinkled and about twice as long as
animal and expands terminally to formthe it is broad, having nearly the width of the
spermatheca, which lies against the peri- adjoining prostate. Most of the pouch is
cardium. The spermatheca, at least in tightly sandwiched between the uterine
some specimens, exhibits pronounced, ir- complex and the albumen gland. The upper
regular, yellowish internal ridges and prostate is rather broad, thick-walled and
seems to be particularly variable in size twisted in its upper part; further down it is
and shape. It was sometimes as large as largely flattened against the underside of
the oothecal gland and sometimes much the oothecal gland andbecomes narrow and
smaller. In one specimen it was very thin toward its terminus. The lower
small and shaped like a short sausage, prostate is a swollen and highly glandular
but in nearly all cases it was observed to division of the male tract, set off by a
be more or less angular and usually it was constriction from the upper prostate,
found to be roundly quadrangular. The which it resembles in color and texture;
duct attaches far off-center on the sper- it is large to very large and may be almost
matheca, at an angle to the organ. It is the size of the whole female complex. This
rather thick in its upper part and is ex- organ is somewhat flattened and is some-
tremely thick and muscular toward its what longer than broad. Its right edge is
junction with the vagina, where it is usually very shallowly concave, while the opposite
nearly as broad as the vagina, although edge is strongly convex, so that it tends to
it was only about 1/3 as broad in one case. be bean-shaped or nearly D-shaped. The
Circular or transverse muscle elements upper side of the lower prostate that is
partly proceeding from the base of the partly appressed to the underside of the
spermathecal duct form a strong vaginal oothecal gland is quite flat or nearly so.
sphincter against and below the base of The wall of this flat side is invaginated
the duct. The sphincter is not abruptly longitudinally, so as to form a single
set off from the musculature of the pre- internal fold traversing its length. For
ceding parts and its upper border is the most part, this line of invagination
disposed diagonally to the vaginal axis. is obliterated by apparent fusion of the
It may bulge out, but not markedly, as a juxtaposed walls. In cross-section the
distinctly spheroidal structure, as is the fold forms a solid, roundly triangular
case in some lymnaeids. Passing through projection; it is rather weak although its
the sphincter, the tract becomes a short, walls are considerably thickened. As
fine tube attaching at the female pore. The compared to that found in some other
pore is a relatively small slit with two species, the wall of the lower prostate is
weakly defined lips, located high on the rather thin and the lumen is rather large.
neck of the animal, close to the insertion At the distal, lower end of the organ the
of the mantle collar. internal fold shows externally as a shallow
Male genitalia - An upper prostate, a groove which divides that end into two,
lower prostate, a vas deferens and a low, rounded protuberances. Arising near
penial complex, form the successive di- the end of the prostate but well on its
visions of the male tract. The upper underside, at the end of the groove, is the
prostate is a long, fleshy-glandular. tubular vas deferens. The vas represents
an abrupt and great reduction in the dia- it may be much swollen proximally and may
meter and glandularity of the tract; along show considerable distad attenuation. The
its length it is highly muscular, chiefly swelling of the preputium is caused by a
due to devlopment of circular muscles. large "velum" that fills its inner end. This
It tends to project from the prostate velum is a fleshy ring attached along the
parallel to the axis of that organ, but soon internal line of junction of the preputium
curves to the right to pass into the body and penis sheath. In some specimens it
wall near the female pore. It then pro- was found to be quite large but very
ceeds directly into the anterior part of asymmetrically developed, so that it pro-
the head-foot organ and then passes out of jected deeply into the preputium on one
the body wall at the base of the right side. The velum lies peripheral to another
tentacle by the male pore. It forms a long ring of tissue, the sarcobelum, which
free loop in the major anterior haemocoel surrounds and closes the outlet of the penis
of the body and then terminates at the sheath; it is small and ill-defined in S.
penial complex. The vas shows no con- montanensis. Internally, the preputium
striction at its originwhere it is about has a prominent glandular lining and two
1/4 the diameter of the lower prostate. longitudinal muscular pillars as long as
Although it tapers somewhat gradually the organ or nearly so. One pillar may
toward its terminus it is notably stout be split, forming an apparent smaller
overall, except where it is most narrowed third one. A few transverse to diagonal
near its attachment to the copulatory folds may pass from pillar to pillar.
organ. That part of the vas deferens in The penis is short and thick and is ex-
the body wall shows through the tissues, tremely so in its more contracted states.
and its stoutness is easily seen on ex- Grossly, it is divisible into two successive
ternal examination of the animal. parts, a basal column and a terminal
The penial complex begins as a tubular, "papilla". The latter is tapered and
abrupt broadening of the male tract, the elongate -conical and is well set off from
penis sheath. This sheath is succeeded the preceding part by its lesser diameter.
distally by a longer and more expanded It forms about 1/6 to 1/3 of the length of
tube, the preputium, which attaches to the the organ. The basal column is very
body wall around the male pore. Enclosed thick and firmly muscular, and it is
in the penis sheath is the penis which widened proximally. The penis shows
largely consists of muscular elements de- bilateral symmetry; it has a greater and
veloped around a direct continuation of a lesser curvature which may be taken
the vas deferens. The penial complex is respectively for dorsal and ventral sides.
of moderate to fairly small size. The There is a transverse swelling of the
sheath is very short and broad; its width lesser curvature that represents a muscu-
is 2/3 of its length, or less. Its length lar penial knot, and there is a median
was observed to be 1/3 to 1/2 that of the longitudinal depression on the opposite
preputium. The sheath is somewhat curvature that suggests the presence of a
tapered distally, and tends to be much- "dorsal" lacuna. The knot occurs at the
swollen proximally where it is 2/3 to 3/4 junction of the basal column and "papilla"
the diameter of the preputium. The of the penis, somewhat distal to the mid-
swelling of the end is due to unusual de- length of the organ. In a few specimens,
velopment of internal "proximal cham- where the organ was rather well extended
bers". The chambers are crowded around and not distorted, the knot was easily
the base of the penis and may extend down identified. However, there usually was
into the sheath well beyond the penial base; distortion resulting from compression of
they are very irregular in shape and have the penis by its sheath. Also, in the con-
glandular walls. The preputium may be tracted state the knot was often pressed
moderately to quite stout, and tends to be tightly backward, against the preceding
fusiform; in smaller specimens especially muscular base, so that it was separated
from the latter only by a closed, practically were faint or obsolete. In the highest
indistinguishable cleft. count obtained for a transverse row, with
Shell size and maturity - For the series vestigial or incipient teeth included, 20
of S. montanensis studied, it appeared that teeth were recorded for the left side of the
sexual maturity could be rather exactly radula and 23 were recorded for the right
related to shell-length. In several speci- side. The number of teeth in particular
mens about 4 mm in shell-length, the transverse rows could not be determined
smallest dissected, all of the genital exactly, and showed some irregular vari-
organs were present, but immature, and ation among rows. The transverse count
the major glandular structures of both given was made in the unworn part of the
male and female systems were especially radula, near the posterior end, where the
small. In these individuals, the penis was counts tended to be higher. The teeth
a distinct, separate structure which pro- closely resembled those figured by F. C.
jected well into the preputium beyond the Baker (1911, 1938) for many species of
sheath, which was very short. It was Stagnicola.
evident that the male system leads the Morphological comparison oiStagnicola
female system ontogenetic ally, and that montanensis with other species - In
the penial complex matures first, closely characters of its soft parts Stagnicola
followed by the lower prostate. The male montanensis clearly differs from all suf-
organs were practically adult in snails ficiently known lymnaeids except one, S.
having a shell 6 mm long, and both genital caperata. The author has acquired con-
systems were mature in size and ap- siderable anatomical evidence which con-
pearance in those having attained a shell- firms that S. caperata is allied to, but very
length of nearly 7 mm and over. distinct from, the species of typical Sto^^m-
Radula - A single radular membrane, cola (Walter, 1961), as contended by F.
from a specimen having a shell 8.1 mm Baker (1928), who partly relied on some
long, was examined. The radula was 1.3 of the more obvious anatomical characters
mm long and 0.7 mmwide; it bore an of S. caperata in erecting the subgenus
estimated 5,500 teeth, which were ar- Hinkleyia for the species. Hubendick's
ranged in 96 transverse rows and in about suggestion that S. caperata and Fossaria
60 longitudinal rows. All of the teeth of hum.ilis are synonymous deserves no con-
the median row, the centrals, were a- sideration. Anatomical characters which
nomalous; each was evidently fused with S. montanensis and S. caperata share with
the first lateral teeth adjoining it on either the American species of typical Sía^n¿ Za
side. The resulting compound structures are: a comparatively short, bilaterally
were extremely broad and irregularly symmetrical penis, with a knot near its
multicuspid. There were 5 and 6 normal midlength; a comparatively robust, com-
bicuspid laterals on the left and right sides paratively short penis sheath; a prostate
respectively, in addition to the anomalous pouch; and a well-developed vaginal
first laterals. In the marginal direction, sphincter. Among these snails there is
the laterals were successively narrower. a general similarity in the conformation
In each transverse row, two tricuspid of the whole genitalia and of the kidney,
intermediate teeth separated the series of and a characteristic common to them is
lateral and marginal teeth on each side. the possession of a series of bicuspid
There were many and smaller cusps on the lateral radular teeth. These forms, on
marginals except on the few that
first these characters, constitute a well-defined
were transitional to the intermediates in natural group among the American
each row and except for those that formed Lymnaeidae even though the status of the
the last several longitudinal rows along genus Stagnicola needs clarification, par-
the edge of the radula. The last two or ticularly in regard to certain European
three teeth in each transverse row were species (see Jackiewicz, 1959). However,
minute, simple, rod- shaped elements that the nominal subgenus Nasonia, of Stagni-
cola, erected for certain American relatively slender, as was noted for S.
species, must be rejected on new evidence. montanensis.

A series oi"Stagnicola ÍNasonia)"bulimo- The possibility of a close relationship
ides teckella from Texas was dissected of these two species to S. árctica, S.
and their anatomy proved to be particularly vahli, and S. diaphana, was raised on con-
similar to, and perhaps identical with, sideration of the report by Hubendick
that of Fossaria. A radula from this (1951) of an unusually short and stout
series was examined also, and its marginal penis in these three forms. However,
teeth were found to be tricuspid, with long the author (Walter) has examined S. árctica
daggerlike cusps: a type of dentition that from the same area (Moose Factory,
is grossly unlike that of Hinkleyia or Hudson's Bay, Ontario, Canada) from
Fossaria, and that is apparently unique in which Hubendick obtained his material of
Lymnaeidae. None of the lateral teeth the species, and its penis, like the
proved to be unequivocally bicuspid; the remainder of its anatomy, proved to be
margins of the two large cusps tended to like that of typical Stagnicola. S. vahli
be irregularly serrate, showing some and S. diaphana rema.in to be reinvestigated
fairly pronounced projections that might in this connection.
have been counted as additional cusps. The No anatomical differences between S.
teeth appeared to be thin and delicate in montanensis and S. caperata were surely
comparison with that observed by the identified, although the penial knot seems
author in the clearly bicuspid laterals of to be less developed in the former. Any
various species of Stagnicola. minor or subtle differences that exist must
The kinship of S. montanensis and S. have been masked by the observed con-
caperata is unquestionable. The charac- siderable variation of particular charac-
ters they share, in contradistinction to ters among individuals. This variation,
other known species of Stagnicola, are: in large part, may not represent ge-
1, a somewhat gradual transition between netically determined characters, but the
the vaginal sphincter and the musculature individual specimen's particular immedi-
preceding it; 2, remarkable stoutness of ate reaction to the killing process used on
the vas deferens; 3, extreme shortness and it, as discussed before. Consequently,
stoutness of the penis; and 4, the placement anatomical differences between S. montan-
of the knot of the penis somewhat distad ensis and S. caperata probably are to be
of the midlength of the organ. The two demonstrated adequately only by study of
species have other characters in common, considerably more material of both
each of which is parallelled in some other species. Differences might also be re-
lymnaeid, but which together set the two vealed by study of serial sections, by
species apart from others. These are: observations of the living animals, and by
great thickness of the lower spermathecal further examinations of radulae. The egg
duct, the shape of the lower prostate and mass of S. montanensis is not known, but
its fold (as described above for S. montan- it may be particularly similar to that
ensis), a short and thick penis sheath, and of S. caperata, which was found to be
an unusually short penial "blade" or unique among known lymnaeid egg masses.
papilla. Both species, in respect to the In the S. caperata mass, the egg-capsules
texture and form of the major glandular are nearly spherical and rather small,
organs of their female system also tend to while their individual sets of envelopes are
diverge from other Stagnicola, while more than twice as thick, relatively, as
tending toward Fossaria. Both show those of other Stagnicola. The egg-mass,
general similarities in their conformation instead of having the common type of thick,
texture, and pigmentation. Also, the foot firm and obviously differentiated outer
and tentacles of S. caperata are usually tunic, has a thin one that is hardly visible.
Cytological Study of the Reproductive Cells other Stagnicolas which have been in-
oí Stagnicola (Hinkleyia) montanensis vestigated (Table 6). This number is
characteristic of lymnaeids, indeed of
Previous cytological studies of Basom- aquatic pulmonate snails in general
matophoran snails include 25 species and (Burch, 1959; 1960b). The only Lymnae-
subspecies of the Lymnaeidae (Table 6). idae with chromosome numbers deviating
Ten of these species and subspecies, all from this basic number are Fossaria
with the haploid chromosome number 18, modicella rustica (n=19), and F. ollula
belong to the genus Stagnicola (9 to pervia. Radix auricularia, R. ovata, and
Stagnicola s.S., 1 to Hinkleyia). The R. pereger (n=17) (Table 6).
purpose of the present study was to de- In general, details of gametogenesis
termine if Stagnicola (flinkleyia) montan- (i.e., chromosome behavior and general
ensis had this same characteristic appearance of the cells) seem to be the
chromosome number and whether or not same as in other lymnaeids. This process
other cytological details of spermato- has been described in detail for Lymnaea
genesis were similar to those found in stagnalis by Perrot (1930) and for Stagni-
other lymnaeids. cola emarginata serrata by Burch, 1960b.
Materials and methods - Fifteen speci- In the male germ line, germinative cells
mens of Stagnicola (.) montanensis were lining the acini of the ovotestis go through
studied cytologically. Six were from Round a series of spermatogonial divisions in
Spring, Gooding Co., Idaho, and 9 from preparation for meiosis. The metaphase
Driggs, Teton Co., Idaho. The snails chromosomes of these cells appear highly
were fixed and preserved in Newcomer's contracted, and caryotype analysis was not
fluid (Newcomer, 1953). The procedure possible from this material. Nevertheless,
followed was to crack the shells (to allow the monocentric nature of the chromo-
rapid penetration of the fixative) and im- somes could still be seen in the "primary
mediately drop the snails into the fluid. constrictions", and side views of meta-
The ovotestes were later removed by phase -anaphase of the first meiotic di-
dissection and stainedby the acetic -orcein vision clearly confirm this. This is in
squash technique (La Cour, 1941). Obser- contrast to the chromosomes "en forme
vations were made with a Tiyoda micro- de bâtonnet ou de boule" described for
scope using a 90x (n. a. 1.25) oil immersion Lymnaea stagnalis by Perrot (1930) and
objective and 10-30x oculars. The the "dot-shaped" ones seen by Inaba and
chromosomes in Figures 3 and 4 were Tanaka (1953) in Fossaria ollula pervia
drawn with the aid of a camera lucida and Radix japónica, which shape the latter
and reproduced at a table -top magnifi- authors consider characteristic of aquatic
cation of 4650x. Photographs (Figs. 5, 6) pulmonate snails (see also Inaba, 1959).
were taken using a 20x ocular, oil im- Miotic chromosomes similar to those of
mersion objective, a Kodak Wratten 57 A Stagnicola montanensis were reported
(green) filter, and Kodak Micro-File film. for other basommatophoran snails by
Observations - This material was not Burch (1959, 1960a, b).
ideal for cytological studies because none Although Perrot (1930) speaks of prima-
of the snails examined were at the peak ry, secondary, and tertiary spermato-
of gametogenesis; they were either imma- gonial cells in Lymnaea stagnalis, there
ture or active gametogenesis had passed. are characteristically six spermatogonial
However, in all specimens there was divisions in Stagnicola montanensis. This
limited activity of the sex cells and two number is indicated by the appearance of
specimens from each locality had enough the sperm and spermatids attached to
dividing cells to determine the chromo- nurse cells in clusters of 256. Only rarely
some number of this species. are there less than six gonial divisions.
The chromosome number of Stagnicola Spermatogonia divide synchronously and
montanensis (n=18) is that found for all may be seen in clusters of 4, 8, 16, and
TABLE 6. Chromosome numbers in the Lymnaeidae^

SUBGENUS /JV^Lí;r/A (LYMNAEIDAE: STAGNICOLA) 257
in general as in other lymnaeids. In

polar view they may be seen as small
ovals or rods, but more often as "tetrads"
(Fig. 3). The chromosomes are often
difficult to count at this stage because some
tend to separate before others, and often
4SÍ this early separation begins before all the
chromosomes have been brought to lie
in one plane. The chromosomes can be
counted most readily and accurately during
late diakinesis and prometaphase of the
first meiotic division. All four speci-
FIGS. 3, 4. Chromosomes of Stagnicola mens from which counts could be made
montanensis 2770x. had 18 bivalents present during late
FIG. 3. Metaphase bivalents of the first prophase and metaphase of the first mei-
meiotic division of spermatogenesis (polar otic division of spermatogenesis. The
view). pairing behavior of the bivalents appeared
FIG. 4. Late prophase chromosomes of a to be normal, and during diakinesis the
gonial cell. paired chromosomes were held together
by one or more chiasmata. Thirty-six
32 cells. All cells of any particular chromosomes were present in gonial di-
cluster are always in exactly the same visions (Fig. 4).
stage of division. Primary spermatocytes The most striking feature in gameto -
are found in clusters of 64 and the genesis of this species compared to other
secondary spermatocytes in clusters of lymnaeids is the presence of five to seven
128. Similar conditions have been seen large chromatin bodies in early prophase
in other lymnaeids (Burch, 1960b). nuclei of the first meiotic division of
The shape, appearance, and behavior of spermatogenesis (Figs. 5 and 6). These
the meiotic metaphase chromosomes are bodies were present in all specimens ex-
FIGS. 5, 6. Cells of Stagnicola montanensis in spermatogenesis, 1370x.

FIG. 5. Early first meiotic prophase of spermatogenesis with six visible deep-staining
chromatin bodies.
FIG. 6. Cells at pachytene or early diplotene stage of spermatogenesis.
.
amined, from both localities. Sometimes men), 570929 (4);Junius Henderson, June
they appear vesicular. Similar structures 26, 1915; USNM 570590 (11).
have not been reported previously in IDAHO. Gooding Co.: SE 1/4 sec. 1,
lymnaeid snails, but they have been seen T. 3 S., R. 13 E. Unnamed tributary of
by Burch (unpublished) in Stagnicola Catchall Creek by house in southwest
caperafa. Perhaps they are similar in corner of Bowman Flat. D. W. Taylor,
nature to those seen by Husted and Burch July 26, 1959; USNM 633926 (14). SW 1/4
(1953) in the polygyrid land sna.il Allogona sec. 4, T. 3 S., R. 14 E. Round Spring
profunda. Although these chromatin (PI. 5, Fig. 2). D. W. Taylor, July 26,
bodies might suggest either the presence 1959; USNM 633927 (about 200).
of supernumerary chromosomes or that Twin Falls Co.: Center of sec. 28, T.
the snail is polyploid, obviously neither is 14 S., R. 18 E. South Fork of Shoshone
the case. The exact nature of these bodies Creek at road crossing 0.55 mile north
in relation to the chromosomes and the of junction of roads to Fawn Spring and
nuclear cycle remains to be determined. Cottonwood Creek. D. W. Taylor, Sept.
In any event they are perhaps another 2, 1957; USNM 633928 (1). NE 1/4 sec.
character linking Stagnicola caperata and 27, T. 14 S., R. 18 E. Seepy area beside
S. montanensis road to Fawn Spring 1.05 miles northeast
of junction with road to Shoshone Creek.
Distribution of Stagnicola montanensis D. W. Taylor, Sept. 1, 1955; USNM 633929
(3). SW l/4SWl/4sec.27,T. 14 S., R. 18
Recent occurrences of Stagnicola E. Head of tributary of South Fork of Sho-

montanensis - This species is widespread shone Creek beside road at junction of
in the eastern Columbia and northern Great road to Fawn Spring and road to Shoshone
Basin drainages in the western United Creek. D. W. Taylor, Sept. 1, 1955;
States (see Fig. 7). Most specimens were USNM 633882 (1, figured specimen), USNM
collected alive, but a few were recently 633883 (2, figured specimens), USNM
dead. 633884 (28). Near center of south edge
The locality information given for U. S. of sec. 3, T. 15 S., R. 18 E. Cottonwood
National Museum shells collected by Hall, Creek. D. W. Taylor, Sept. 1, 1955:
Hoffman, and Sinitsin is all that is on the USNM 633885 (2, figured specimens),
labels; in some cases it is not precise. USNM 633886 (60). Southeast corner of
The original data accompanying the speci- SW 1/4 sec. 4, T. 16 S., R. 18 E. SmaU
mens were destroyed several years ago. trickle of water in wash 0.1 mile north of
In the following list of localities, the quarter corner. D. W. Taylor, Sept. 2,
number of specimens is in parentheses 1955; USNM 633930 (71).
after the citation oí institution^ and cata- Franklin Co.: Leslie Wright's ranch,
logue numbers. Localities are listed 30 miles south of Soda Springs (according
from west to east and north to south. to local inquiry this locality is in theW1/2
MONTANA. Ravalli Co.: Hays Creek sec. 7, T. 12 S., R. 41 E). L. Sinitsin,
near Ward (probably about S 1/2, sec. 3, Sept. 12, 1929; USNM 530919 (12). NW
1/4
T. 4 N., R. 21 W.). L. E. Daniels, April sec. 33, T. 12 S., R. 40 E. Spring on
25, 1912; UMMZ 76196 (type series, 35), south side of Cottonwood Creek. D. W.
27799 (6), USNM 570589 (1 figured speci- Taylor, R. Bright, Sept. 10, 14, 1959;
10 CAS = California Academy of Science, San Francisco, California.

UCMNH = University of Colorado Museum of Natural History, Boulder, Colorado.
UMMZ = Museum of Zoology, University of Michigan, Ann Arbor, Michigan.
USGS = U. S. Geological Survey, Washington D.
USNM = U. S. National Museum, Washington D. C.
117 115'^ 113^ lll^'

119
EXPLANATION
Sfagnicola pilsbryi (Hemphill)
• Recent S. montanensis (Baker) 100 200
A Fossil 5 montanensis (Baker)

—
f-
J.
1^'
SCALE
.''
\ NX
FIG. 7. Map showing distribution of Stagnicola pilsbryi (Hemphill) and S. montanensis
(Baker) in the U. S. A. Precise locality data are given in the text.
USNM 633933 (67). Canyon, 22 miles east of Salt Lake City.

Bear Lake Co.: Pearl Cr., NW 1/4 sec. Half-dry spots by a ditch. L. Sinitsin and
30, T. 10 S., R. 43 E. D. W. Taylor, R. others, Sept. 26, 1929; USNM 530921 (9).
Bright, Aug. 29, 1961; USNM 633920 (2). Head of East Canyon, 25 miles east of
Unnamed tributary of Stauffer Creek, NW Salt Lake City. Swampy pasture, Mr.
1/4 SW 1/4 sec. 5, T. 12 S., R. 43 E. McPolim's ranch. L. Sinitsin and others,
D. W. Taylor, R. Bright, Aug. 29, 1961; Sept. 26, 1929; USNM 530922 (7). Head of
USNM 633921 (48). Emigration Canyon, East Canyon, 25 miles east of Salt Lake
W NE 1/4 sec. 23, T. 12 S., R. 42 E.
1/2 City. Swampy pasture along a ditch at
D. W. Taylor, R. Bright, Aug. 29, 1961; Mr. McPolim's ranch. L. and D. Sinitsin,
USNM 633922 (66). Sept. 26, 1929; USNM
531477 (6). Head of
Teton Co.: Center of sec. 35, T. 6 N., East Canyon, 27 miles east of Salt Lake
R. 45 E. Ditches beside state highway. City. Swampy pasture, Mr. Pace's ranch.
D. W. Taylor, J. D. Love, J. de la L. Sinitsin and others, Sept. 26, 1929;
Montagne, Aug. 28, 1959; USNM 633931 USNM 530923 (9). Creek at Park City.
(about 150). SE 1/4 NW 1/4 sec. 35, T. L. Sinitsin, Oct. 1, 1929; USNM 530912 (7).
5 N., R. 45 E. Ditch and small stream on Beaver Co.: Low's ranch, Beaver. Hall
west side of state highway at south edge and Hoffman, Sept. 24, 1929; USNM 530914
of Driggs. D. W. Taylor, J. D. Love, J. (6).
de la Montagne, Aug. 28, 1959; USNM NEVADA. Nye Co.: Springs 2 miles
633932 (about 125). Four miles west of southeast of MiUett P. O. May, 1931; CAS
Victor. Muddy stream, Henry Drake's 29060 (2).
11
ranch. L. Sinitsin, Sept. 15, 1929; USNM Fossil occurrences^ of Stagmcola
530998 (22). Four miles southwest of montanensis .
Victor. Half-dry ditch, Blanchard' s ranch. PLIOCENE. Teewinot Formation, Teton

L. Sinitsin, Sept. 15, 1929; USNM 531179 County, Wyo.: locality^ 19105. USNM
(10), 530997 (24). 563097 USGS (62).
(1),
WYOMING. Lincoln Co. Pasture at Bed-

: LATE PLEISTOCENE. American FaUs
ford. W. H. Krull, May 28, 1941; USNM Lake Beds, Power County, Idaho: locality
536403 (13). Grassland at Grover. W. 19169. UMMZ uncat. (8).
H. Krull, May 29, 1941; USNM 536407 Older alluvium, Power County, Idaho:
(about 100). Pasture at Grover. W. H. locality 20474. USGS (9).
Krull, May 30, 1941; USNM 536416 (78). Unknown Power County, Idaho:
unit.
Near Afton. W. H. KruU, Sept. 15, 1941; locality 20477. USGS (6).
USNM 536460 (16). Meadows and pastures Unnamed unit, Bannock County, Idaho:
in the region of Afton. W. H. Krull; locality 22328. USGS (about 400). Locality
USNM 522817 (45). 22410. USGS (9).
UTAH. Cache Co.: Between Logan and Deposits of Lake Thatcher, Caribou and
Smithfield. Muddy ditch across a swampy Franklin counties, Idaho: Locality ?21143,
pasture by the road. L. Sinitsin, Sept. 24, USGS (1). Locality 22396. USGS (25).
1929; USNM 531309 (1). Spring at north Locality 23066. USGS (1).
side NW 1/4 NW 1/4 sec. 23, T. 14 N., R. Unnamed unit, Bear Lake County, Idaho:
3 E., tributary to Beaver Creek. D. W. locality 21062. USGS (7).
Taylor, Aug. 27, 1961; USNM 633923 (59). Provo Formation, Franklin County,
Summit Co.: Small stream 2 miles Idaho: locality 22395. USGS (21).
northwest of Kimballs. Leslie Hubricht,
1938; UMMZ 176539 (4). Head of East
11
Precise geographic and geologic information on the localities is given in the Appendix,
P 270.
Stagnicola iflinkleyia) caperata (Say), 1829 USNM 570483.

Canyon Co.: Indian Creek near Caldwell.
PI. II, Fig. 2-12 H. M. Tucker; UCMNH 16354 in part.
Elmore Co.: NE 1/4 sec. 12, T. 5 S.,
R. 10 E., and NW 1/4 sec. 7, T. 5 S., R.
Galba caperata (Say): Baker, 1911,
Chicago Acad. Sei. Spec. Pub. 3: 225, PI. HE. Irrigation ditch oncommon section
28, Fig. 20-33; PI. 29, Fig. 1-3. line. D. W. Taylor, Aug. 8, 1959; USNM
Stagnicola caperata (Say): Baker, 1928, 633934. NW 1/4 sec. 8, T. 5 S., R. 11
Wisconsin Geol. Nat. Surv. Bull. 70(1): E. Irrigation over-flow. D. W. Taylor,
260, PI. 18, Fig. 43-47.
Aug. 8, 1959; USNM 633935.
Gooding Co.: Dry Creek, center E side
This species is accepted in the sense of sec. 15, T. 5 S., R. 14 E. D. W. Taylor,
F. C. Baker (1911,1928), andnodiscussion July 18, 1959; USNM 633936.
of variation or synonymy is necessary. Twin Falls Co.: Salmon Falls Creek,
Some descriptive and diagnostic detail on SW 1/4 NE 1/4 sec. 30, T. 8 S., R. 14 E.
the shell is given in the section on Stagni- D. W. Taylor, July 30, 1955; USNM 633937.
cola montanensis under "Comparisons Salmon River near Twin Falls (probably
of shells" p 241; anatomical details and same locality as preceding). L. Sinitsin,
a description of the egg masses can be Sept. 7, 1929; USNM 530983. Deep Creek,
found under "Morphological comparison NW 1/4 NWl/4 sec. 7, T. 11 S., R. 14 E.
of Stagnicola montanensis with other spe- D. W. Taylor, Aug. 28, 1955; USNM 633938.
cies* on p 253. Power Co.: Snake River, NE 1/4 sec.
Ecologie information for S. caperata 12, T. 9 S., R. 29 E. D. W. Taylor, Sept.
east of the Rocky Mountains is available 3, 1961; USNM 633924.
through the references cited above; the Clark Co.: North of Dubois on the road
habitats of S. caperata in the Snake River to Dillon, Montana. L. Sinitsin, Sept. 17,
and Great Basin drainages are discussed 1929; USNM 531442.
on p 267-270. Jefferson Co.: Ditch beside highway 15
The oldest known geologic occurrence of miles south of Dubois. L. Sinitsin, Sept.
S. caperata is middle Pliocene. Specific 16, 1929; USNM 531377.
locality data and the evidence for the age Bannock Co.: Bridge west of Pocatello
assignment are given by Taylor (1960). (probably Portneuf River). Junius
The geographic distribution of S. caper- Henderson and E. H. Nanney, June 20, 1927;
ata is from "Quebec and Massachusetts UCMNH 15192, USNM 570452. Wet pasture
west to California; Yukon Territory and near Lava Hot Springs. L. Sinitsin, Sept.
James Bay south to Maryland, Indiana, 11, 1929; USNM 531400.
Colorado and California" (Baker, 1928). Caribou Co.: Brook near reservoir 2
The following specific locality records are or 3 miles west of Soda Springs. Junius
for the eastern Columbia River drainage and B. R. Henderson, July 23, 1930;
and northern Great Basin only (compare UCMNH 17430, USNM 570453. Swenson's
with Fig. 8). They are intended to docu- ranch, 25 miles south of Soda Springs.
ment the relative abundance and distri- (From local inquiry this locality is near
bution of Stagnicola caperata and include the common corner of sees. 17, 18, 19,
both published and unpublished oc- 20, T. 11 S., R. 41 E). L. Sinitsin, Sept.
currences. 12, 1929; USNM 530916, 531387. Mee-
Recent occurrences of Stagnicola cham's ranch, 25 miles south of Soda
caperata. Springs. (From local inquiry this locality
MONTANA. Flathead Co.: no specific is in the NW 1/4 sec. 20, T. 11 S., R. 41
locality. L. E. Swanson; USNM 468507. E.). L. Sinitsin, Sept. 12, 1929; USNM
IDAHO. Washington Co.: Pool 2 mi. 530917, 531335, 531340.
SE of Weiser. Junius Henderson and E. Franklin Co.: P. Andersen's ranch,
H. Nanney, June 22, 1927; UCMNH 15205, 30 miles south of Soda Springs. (From
PLATE II
EXPLANATION
• Recent Slagnicola coperata (Soy)
* Fossil S coperata (Say)
FIG. 8. Map showing distribution of Stagnicola caperata (Say) in the eastern Columbia River and northern
Great Basin drainages. Precise locality data are given in the text.
PLATE U
FIG. 1. Stagnicola pilsbryi. "Fish Spring, Nevada". Probably Fish Springs, Juab Co., Utah. Length 7.9
mm, 61/4 whorls. Type. ANSP Ö2293.
FIGS. 2, 3. S. caperata. Logan, Cache Co., Utah. Length 8.0 mm, 5 1/4 whorls. USNM 522635c.
FIGS. 4, 5. S. caperata. Same locality. Length 9.0 mm, 5 3/4 whorls. USNM 522635a.
FIGS. 6, 7. S. caperata. Same locality. Length 8.0 mm, 5 1/4 whorls. USNM 522635b.
FIGS. 8, 9. S. caperata. Meecham's ranch, NW 1/4 sec. 20, T. 11 S., R. 41 E., Caribou Co., Idaho. Length
7.3 mm, 4 3/4 whorls. USNM 531335c.
FIG. 10. S. caperata. Same locality. Length 11.1 mm, 5 3/4 whorls. USNM 531335a.
FIGS. 11, 12. S. caperata. Same locality. Length 8.6 mm, 4 3/4 whorls. USNM 531335b.
local inquiry this locality is probably in 1934; USNM 633944.

the E 1/2 sec. 7, T. 12 S., R. 41 E.). Cache Co.: Swamp, Richmond. W. H.
L. Sinitsin, Sept. 12, 1929; USNM 530918, Krull, Mar. 31, 1941; USNM 536402. Near
531353. Leslie Wright's ranch, 30 miles Bear River, Benson Ward. W. H. Krull,
south of Soda Springs. (From local inquiry May 6, 1942; USNM 536465. Between
this locality is in the center of the 1/2W Logan and Smithfield. L, Sinitsin, Sept.
sec. 7, T. 12 R. 41 E.). L. Sinitsin,
S., 24, 1929; USNM 633945. Logan. W. H.
Sept. 12, 1929; USNM 530920, 531430. Krull, Feb. 20, 1939; USNM 522624.
NW 1/4 NW 1/4 sec. 20, T. 15 S., R. 39 Wennergren pasture 1, Logan. W.H. Krull,
E. Roadside ditch. D. W. Taylor, R. Mar. 1, 1940; USNM 522629. Wennergren
Bright, Sept. 11, 1959; USNM 633939. far pasture, Logan. W. H. Krull; Apr. 10,
Bear Lake Co.: Rivulet along auto road 1940; USNM 522632. Reese pasture,
17 miles northwest of Montpelier. Junius Logan. W. H. Krull, Apr. 10, 1940;
Henderson and F.E. Swisher, July 27, 1921 ;
USNM 522635.
UCMNH 11792, USNM 570451. Pasture, Rich Co.: Slough on west side of Bear
Liberty. W. H. Krull, June 6, 1941; USNM River valley about 10 miles west of Sage.
536427. Pasture, Ovid. W. H. Krull, June Junius Henderson; USNM 570464.
6, 1941; USNM 536434. Meadow, Paris. Weber Co.: Pond at head of "Confusin'
W.H. Krull, May 22, 1941; USNM 536365. Canyon", SW 1/4 sec. 21 T. 5 N., R. 1 W.
Pasture, Paris. W.H. Krull, May 22, 1941; 4700 ft. elev. J. H. Feth, Sept. 1954; USNM
USNM 536371. Roadside ditch 2 miles north 633947.
of Montpelier. Junius Henderson and F. E. Salt Lake Co.: Small slough on 4th
Swisher, July 27, 1921; UCMNH 11788. South, 11th West, near Jordan River, Salt
WYOMING. Teton Co.: Center sec. 20, Lake City. Treganza; USNM 199399. Salt
T. 40 N., R. 116 W. Flat Creek at road. Lake City. L. E. Swanson; USNM 424332,
D. W. Taylor, Sept. 2, 4, 1959; USNM 424333. Salt Lake City. F. A. Sampson;
633940. USNM 570466. Near Salt Lake City.
Sublette Co.: Dry hollow in floor of Dell Henry HemphiU; UMMZ 73688. Salt Lake
Creek valley about one mile above mouth City. UMMZ 142927. Southeast of Murray;
of creek, SW 1/4 sec. 21, T. 38 N., R. 113 L. A. Giddings, 1921; UCMNH 10571.
W. D. W. Taylor, Aug. 7, 1950; UMMZ Summit Co.: Creek at Park City. L.
178185. Wet moss and grass flat in floor Sinitsin, Oct. 1, 1929; USNM 633946.
of Dell Creek valley about one mile above Tooele Co.: No specific locality. A.
mouth of creek, SW 1/4 sec. 21, T. 38 N., G. Ruthven, 1930; UMMZ 49783.
R. 113 W. D. W. Taylor, Aug. 8, 1950; Utah Co.: Slough south of Lehi. Junius
UMMZ 178198. Small stream just north and B. R. Henderson, July 15, 1930;
of Faris ranch house, NW 1/4 sec. 28, UCMNH 17373, USNM 570463.
T. 38 N., R. 113 W. D.W. Taylor, Aug. Beaver Co.: Low's ranch, Beaver. Hall
9, 1950; USNM 633941. Dell Creek, center and Hoffman, Sept. 24, 1929; USNM 530913.
N 1/2 NE 1/4 sec. 29, T.38N., R. 113W., Beaver. L. E. Swanson, Sept. 22, 1934;
D.W. Taylor,Aug. 12, 1961; USNM 633925. USNM 539047. Beaver. L. E. Swanson,
Uinta Co.: Small creek where highway Oct. 8, 1934; USNM 539048, Beaver. L.
89 crosses Bear River. Honess, July 16, E. Swanson, Nov. 21, 1934; USNM 539049.
1953; USNM 633942. SmaU stream tribu- Small spring slough at south edge of
tary to Bear River on Jamison Ranch Beaver. Junius and B. R. Henderson,
(lower ranch), 7 miles S of Evanston on July 8, 1930; UCMNH 17334.
highway 89. Honess, July 16, 1953; USNM NEVADA. Elko Co.: Pools of Humboldt
633943. River vaUey. Helen Gaige, 1912; UMMZ
UTAH. Box Elder Co.: Adney farm, 67590. Humboldt River. Crystal
Corinne. W. H. KruU; USNM 513888. Thompson, 1912; UMMZ 5490, 5491, UC
Sloughs at Bear River Bay bird sanctuary, MNH 17121.
west of Brigham. S. S. Berry, Sept. 3, Nye Co.: Discharge of warm spring,
Monitor Valley. UMMZ 132552. Shell of Stagnicola pilsbryi

Fossil occurrences^^ of Stagnicola
caperata Diagnosis - A
species of Hinkleyia with
MIDDLE PLEISTOCENE. Unnamed slightly shouldered whorls whose greatest
formation, southwestern Idaho. Owyhee width lies above their middle, a narrow
Co.: localities! 2 20404, 20472. Elmore tapering spire, small aperture, and dull
Co.; localities 20105, 20107, 20109, 20112, shell surface. Spiral sculpture consists
20120. Twin Falls Co.: localities 19219, of weak, fine, irregular incised lines.
19222, 19225, 20410. Axial sculpture is of fine to moderately
Raft formation. Power Co., Idaho: coarse irregular growth lines.
Locality 20478. Type - Academy of Natural Sciences
LATE PLEISTOCENE. American Falls of Philadelphia 62293. «Fish Spring,
Lake Beds, Power Co., Idaho: locality Nevada" is the only locality data given by
21643. Hemphill. Baker(191 l)identified this more
American Falls Lake Beds or Michaud precisely as being in Nye County, in ap-
Gravel, Power Co., Idaho: locality 21057. proximately lat. 38o 45 ft. N., 30 ft.
Older alluvium or Aberdeen terrace W. Hemphill's label with the type says
deposits, Power Co., Idaho: locality 21636. "Fish Spring Nevada between Austin and
Unknown unit, Bingham Co., Idaho: Salt Lake", thus ruling out the locality
locality 20479. specified by Baker. Most probably Hemp-
Sunbeam Formation?, Power Co., Idaho: hill collected at Fish Springs, northern
locality 22332. Juab County, Utah (see map, Fig. 7).
Unnamed alluvium. Twin Falls Co., Description of shell - Shell turriform.
Idaho: locality 20413. Whorls 6 1/4, separated by a constricted
Deposits of Lake Thatcher, Caribou Co., suture. Nuclear whorl relatively large,
Idaho: localities ?23025, 23027, 23033, as in Stagnicola caperata. The greatest
23037, 23061, 23066. width of the last three whorls is above
Unnamed unit, Bear Lake Co., Idaho: their middle, so that they tend to become
locality 22429. shouldered and slightly flattened peripher-
Lake Bonneville deposit. Caribou Co., ally. Aperture relatively small, one -third
Idaho: locality 23069. of total shell length, slightly oblique.
Unnamed unit. Box Elder Co., Utah: Both inner and outer lips in the type are
locality 22359. broken, as shown in the drawing. Colum-
Saltair core, Salt Lake
Co., Utah: ella with only a slight trace of fold.
localities 21112, 21120, 21122, 21124. unbroken, the shell would not be as
Unknown unit, Humboldt Co., Nevada: umbilicate as illustrated, but with a
locality 19192. narrow perforation. Axial sculpture of
fine to moderately coarse, irregular
Stagnicola (flinkleyia) pilsbr'yi (Uem^phill), growth lines slightly retractive to the base.
1890 Spiral sculpture of weak, fine, irregular
incised lines; on the body whorl these are
PI II, Fig. 1 evident on the posterior part only.
Length of type 7.9 mm, width 3.3, length
LeptoUmnea, n, sp.?: Cooper, 1872, of aperture 2.6, width of aperture 1.7,
California Acad. Sei. Proc, ser. 1,4: 172. whorls 6 1/4.
Limnaea [LeptoUmnea) Pilsbryi Hemp- - Only three specimens of this
Remarks
hill, 1890, Nautilus 4: 25.
Galba pilsbryi (Hemphill): Baker, 1911, species are known, all from the original
Chicago Acad. Sei. Spec. Publ. 3: 254,
PI. 4, Fig. A,B.
12precise geographic and geologic information on the localities is given in the Appendix,
P 270.
lot collected by Henry Hemphill in 1868. flattened on its sides and occupying a
The two immature specimens are less than little more than half the length of the
half the length of the type. They agree growth very delicate, suture
shell; lines of
with the type in size of nuclear whorls, deep; aperture oval, longer than wide,
shape, and axial sculpture but lack spiral outer lip acute; inner lip subreflexed.
sculpture. The bleached periostracum and Length 3/8, breadth 1/8 of an inch.
the dirt inside the aperture shows that all Habitat: Fish Spring, Nevada.
three were collected as empty shells. I collected a few specimens of this
Hemphill's label notes "These are the best interesting shell in the month of June,
I can do for you. I have but two or three 1868, at this locality, after a long and
others the largest more globose than hard day's ride of 40 miles horseback.
these". These additional specimens have Another long ride next day of 50 miles to
not been traced. water, compelled an early start and thus
The relatively coarse growth lines, the opportunity to secure more specimens
small size, narrow shape, small aperture, was lost." (Hemphill, 1890, p 25-26).
and shouldered whorls of this species are
all characters much like those of Fossaria. DISTRIBUTION
It is surprising that Baker (1911),
not AND HABITATS
Hannibal (1912, p 144) and Hubendick (1951,
p 199) have grouped Stagnicola pilsbryi Stagnicola pilsbryi is known only living,
with species of Fossaria. Nevertheless but both S. caperata and S. montanensis
the relatively large nuclear whorl and are known from Pliocene deposits. The
spiral sculpture bar pilsbryi from earliest record of S. montanensis and of
Fossaria and ally it with Stagnicola S. caperata is middle Pliocene, but from
caperata (Say) andS. moniane^sis (Baker). the fact that they appear essentially
Stagnicola pilsbryi differs from S. modern at that time the geologic age of
montanensis in sculpture, rate of whorl the species is probably somewhat greater.
increase and form. The growth lines are Within the area of the maps (Figs. 7, 8)
coarser, so that the shell surface is dull there is no evidence of major changes of
rather than shiny. The spiral sculpture distribution. All of the fossil and recent
consists of incised lines, rather than occurrences of each species represent
series of tiny arcuate lines. The whorls the same pattern of distribution.
enlarge more slowly, so that the body The three recognized species of
whorl of S. pilsbryi is relatively smaller Hinkleyia have successively larger
than in S. montanensis and the shell as ranges, (see maps, Figs. 7, 8), so that the
a whole is smaller than shells of S. area of distribution of one is enclosed by
montanensis with the same number of that of another. Stagnicola pilsbryi is
whorls. The last whorls of S. pilsbryi known only from a single locality in
are shouldered, whereas in S. monianensfs western Utah. S. montanensis is found in
they are regularly convex. the northern Great Basin and southeastern
In making these comparisons only one Columbia River drainages. S. caperatais
adult specimen of S. pilsbryi is available, widespread in northern North America.
and hence no allowance for range of A list of localities and a description of
variation has been made. Several hundred habitats of the species of Stagnicola
'specimens of S. montanensis have been {Hinkleyia) have some value in themselves
examined, however, and their range of in recording what is known about the
variation in several characters does not species. The information available does
overlap the type of S. pilsbryi. not suffice to answer questions about
Original description - "Shell elongated, differentiation of the species oí Hinkleyia,^
narrow, somewhat solid, smooth, of a their isolating mechanisms or general
light horn-color; consisting of about six distribution, but it does suggest directions
roundly-shouldered whorls, the last of future study.
SVBGEÛS HINKLEYIA (LYMNAEIDAE: STAGNICOLA) 267
Habitats Scotland.
Biogeography - Certain details of the
Nothing is known of the habitat of distribution of Stagnicola montanensis and
Stagnicola pilsbryi. From the fact that S. caperata are simply and adequately
it narrowly localized species, where-
is a explained by their different habitats. In
as its close relatives are widespread, one the Snake River drainage in south-central
might infer it has some ecological special- Idaho, and in the Bear River drainage in
ization. southeastern Idaho, S. montanensis is an
Stagnicola caperata is found most often upland species and S. caperata is a low-
in seasonal bodies of water. It is charac- land form. This local distribution is
teristic of such habitats as irrigation evidently related to the differences in
ditches, sloughs and shallow ponds. Its water bodies of the mountains and low-
wide distribution is correlated with its lands.
tolerance for environments in which few Nevertheless not all of the differences
other snails live, and with the common in distribution between S. caperata and
and widespread occurrence of habitable S. montanensis can be explained ecolo-
situations. As aquatic snails go, it is gically. The restriction of S. montanensis
one not readily subject to geographic to the Great Basin and Columbia River
isolation. drainages is the most obvious case. Any-
Stagnicola montanensis is unique among one with field experience in the Green
North American Lymnaeidae in its habitat. River, and upper Mississippi River
It is a pure-water snail, living in the out- drainages as well, finds it hard to believe
flow of springs, or in clear mountain there is any sharp écologie difference
streams. It is never found in seasonal which would prevent species of the western
ponds, or stagnant or muddy water bodies, drainages from inhabiting the eastern.
but it is never in large clear waters such as In lesser details too the distributions of
lakes and rivers. Most Lymnaeidae of these two species seem to be influenced
clean water bodies live in large perennial by non-ecologic factors. Only S. montan-
rivers and lakes {Stagnicola (s.s.) emar- ensis has been found in the Teton River
ginata group, Bulimnea, Acella). Other and Salt River drainages. Only S. caperata
Lymnaeidae living in shallow marginal has been found living in the upper Snake
situations are usually found in ponds, River and Bear River drainages in
ditches, sloughs, or swamps without evident Wyoming, although S. montanensis is
regard for muddiness or the presence or known fossil from Jackson Hole.
lack of current. Their occurrence is more These apparently non-ecological factors
probably correlated with length of growing of distribution are probably biogeographic.
season and the vegetations. S. montanensis This is to say that they are the result of
is unique in combining a pure-water habitat the biologic and geologic history of the
with a small or even seasonal water body. region, and that they reflect past events
At several localities S.montonensis was instead of present differences between the
notably concentrated in the deeper parts of two species.
the shallow pools it lived in. Possibly this If these details of distribution are cor-
reflects a physiological specialization for rectly interpreted as biogeographic, then

breathing by means of a gas bubble in the there are significant differences in faunal
lung, or by cutaneous respiration, rather history between the Green River, and the
than directly inhaling air. Even if these Snake River and Bear River; and between
colonies were in fact breathing while sub- the upper Snake and Bear Rivers of western
merged, they need not be characteristic of Wyoming and the more western parts of
the species. Hunter (1953) found different these drainages. These differences may
states of respiratory behavior within be expected in other groups of aquatic
different populations oihoth Radix pereger animals. As knowledge of late Cenozic
and Physa fontinalis in Loch Lomond, geologic history in this region becomes
.
more detailed, then one may expect to find summer they dwindle to only a trickle, or
differences also in physical history of the a series of scarcely connected shallow
basins mentioned. pools, and may dry up entirely in the
Another implication of the historical uppermost parts of their courses. S.
control of such details of distribution is montanensis was common in the pools
that dispersal (active or passive) in these in these small water courses, staying in
species is very slow. Even though these the deeper water rather than crawling
snails live in swampy situations, springs about on mud barely in or out of the water,
and ponds, in shallow water bodies, they as Fossaria dalli or F. obrussa usually
are not effectively carried by birds. One do. At most places where S. montan-
can readily understand how snails or clams ensis was found alive it was the only
restricted to larger lakes and streams are aquatic moUusk; rarely Pisidium casert-
limited to dispersal through their habitat; anum was associated.
such seems to be the case also with One locality (PI. ,
Fig. 1) deserves
Stagnicola montanensis and S. caperata. specialmention because it is the place
Description of habitats in south-central from which the dissected series comes,
Idaho - Since 1955 Taylor has been making and at which the snails were found
a detailed survey of the living mollusks aestivating. The stream is a small,
of south-central Idaho in conjunction with seasonal rivulet tributary to the South
geologic studies. The sparse records of Fork of Shoshone Creek. It runs through
the two species of Hinkleyia in this area a grove of aspen and pine trees; beside
are therefore not due to inadequate the stream are sparse willows. The
collecting. bottom is formed of platy cobbles of latite,
The Snake River valley in the area the local bedrock, with mud and some
between Twin Falls and Glenns Ferry is gravel in the hollows. On September 1,
a nearly flat or gently rolling arid plain 1955, the stream had shrunk to a few
cut by the canyon of the Snake River. It pools (16 C.) in which 5. montanensis
is naturally vegetated by little more than was common. On September 2, 1957,
sagebrush and grass. Habitats suitable when the photograph was taken, no water
for aquatic mollusks are restricted to the remained in the stream bed. The snails
Snake River, its few perennial tributaries, were found adhering to the under sides of
and irrigation ditches and artificial reser- stones and logs in the low spots of the
voirs. Stagnicola caperata occurs only in bottom. They revived promptly when
the lowlands close to the Snake River. At placed in water, and were preserved for
all known localities it is in situations dissection. The only unusual feature
created or influenced by irrigation water. of the shells from this locality is their
Volcanic hills of basalt and latite rise corrosion (see PI. I, Fig. 3).
above the irrigated plains both north and Description of habitats in east-central
south of the Snake River. These are the Idaho - In the Teton River drainage in
Mount Bennett Hills north of the river, east-central Idaho, Stagnicola montan-
and the Rock Creek Hills in southeastern ensis is the only known species of
Twin Falls County. They are only slightly Hinkleyia. The two localities where Taylor
less arid than the lowland, and have aspen has collected it are roadside ditches.
trees or conifers only on north slopes or These situations were apparently more
in protected places. In both of these like those where S. caperata is found
upland areas the habitat of S. montanensis elsewhere than like other habitats of S.
is similar- small, seasonal streams or montanensis
seepages. These are the farthest up- In sec. 35, T. 6N., R. 45 E., roadside
stream locations of perennial or nearly ditches on either side of Idaho state high-
perennial water. During the spring run- way 33 just north of the bridge over the
off these streams are about 2 to 4 feet South Fork of Leigh Creek were dry at the
wide and several inches deep, but in time of collection. The lower spots were
-
PLATE
Habitats of Stagnicola montanensis
FIG. 1. montanensis
RcLbitât oí Stagnicola FIG. 2.Rabitâtoî Stagnicola montanensis
headwaters of
at the type locality in the in the canyon of Cottonwood Creek,
the South Fork of Shoshone Creek, Twin Franklin Co., Idaho. View westward, up-
Falls Co., Idaho. View eastward, upstream stream, in the upper area of seepage of
showing the seasonally dry stream bed. a spring on the south side of the canyon.
On Sept. 1, 1955, the stream had shrunk The smallest pools at the head of the
to a few poolswhich S. montanensis vâ.s
in seepage were inhabited by 5. montanensis,
common. On2, 1957, no water
Sept. Fossaria dalli (Baker), F. obrussa (Say),
remained. S. montanensis was found Gyraulus circumstriatus (Tryon), Pisi-
adhering to the undersides of stones and dium casertanum (Poli) and a small hydro
logs in the low spots of the bottom. No biid. Larger pools (foreground) had S.
other freshwater moUusks were found. montanensis, S. eZodes (Say), Physagyrina
Photograph by D. W. Taylor, Sept. 2, 1957. (Say), Pisidium casertanum. (Poli) and the
hydrobiid. Photograph by D. W. Taylor,
Sept. 14, 1959.
damp, but tall grasses grew all through area of seepage, hollows and crevices
the ditches and there was no trace of among the stones contained some water-
submergent aquatic plants. A few juvenile cress. Increasedvolume produced a small
S. montanensis were living, but numerous rivulet which formed and kept fresh quiet
adult shells were empty. The only associ- pools a foot or 2 across, 3 to 4 inches deep,
ated mollusks were the land snails Discus bordered by watercress, and with a soft
cronkhitei (Newcomb), Nesovitrea elec- mud bottom. Several such sources com-
trina (Gould), and empty shells of an bined in a pool several feet across, about 6
indeterminate succineid. inches deep, stagnant, vjithChara, Myrio-
Six miles south of this locality, on the phyllum, filamentous green algae, and
south side of Driggs, Stagnicola montan- watercress at the edge. This pool emptied
ensis was found again in a roadside ditch by a small stream into Cottonwood Creek.
on the west side of state highway 33. The The changes in volume and nature of the
ditch drained northward into a minor water bodies were found to be correlated
stream which was shown to be perennial by with changes in the moUusk population. In
patches of submergent aquatic vegetation the smallest water bodies at the spring
and by the continued moderate flow even in sources S. m,ontanensis Pisidium casert-
,
late summer. The ditch was mostly dry, anum (Poli), and a small hydrobiid were
but close to the stream the hollows were common. Gyraulus circumstriatus (Try-
wet and at the end of the ditch there was a on), Fossaria dalli (Baker) and F.obrussa
large pool 3 to 4 feet wide and 12 to 18 (Say) were also present but rare. The
inches deep (PI. IV, Fig. 1). S. montan- larger, fresh pools 1 to 2 feet in diameter
ensis was in slowly flowing water at the yielded S. montanensis, Pisidium caser-
culvert, in quiet water in the pool, and in tanum, and the hydrobiid as well as Sia^^m-
the dried-up hollows of the drainage ditch. cola (s.S.) elodes {Sa.y) a.nd Physa gyrina
Associated \\\1.^ ere Stagnicola {s.s.) (Say). The last two species were the only
elodes, (Say), Gyraulus circumstriatus mollusks in the stagnant pools and the
(Tryon), and Aplexa hypnorum (Linnaeus). lower parts of the discharge into cotton-
These snails are often found with S. ca/)er- wood Creek. The terrestrial snail
ata in the eastern United States, and they Oxylomzi was found also among grasses and
reinforce the impression gained from the watercress in the seepage area and along
habitat that here S. montanensis is in a the outflow of the large pool.
place where S. caperata might be expected Stagnicola montanensis occurred in
instead. pools varying in size from 1 or 2 feet to
Description of habitats in southeastern 2 or 3 inches in diameter, in water without
Idaho - In Franklin County, Idaho, D. W. perceptible current but nevertheless kept
Taylor and R.C. Bright found S. caperata fresh by seepage. It also was found in
and S. montanensis at one locality each. slow current, among algae in the rivulet
On thewestsideof the Bear River in north- toward the large pool. It was most abundant
ern Cache Valley, sec. 20, T.15S.,R. 39 E., in very small pools in the seepage area,
S. caperata was common in a roadside in watercress, on mud and leaves, always
ditch. The standing water was about 2 feet submerged.
wide and up to 6 inches deep. The sparse
vegetation gave the impression that the
APPENDIX
ditch had recently been scraped. S. caper-
ata was the only moUusk. Fossil Localities
At the south end of Gentile Valley S. mon-
tanensis was common in a spring on the Fossil localités in the following list
south side of Cottonwood Creek. The spring are arranged stratigraphically from older
issued from a number of small sources to younger, and geographically from north
next to the steep south wall of the canyon, to south and west to east. Locality numbers
and from among cobbles and boulders in the are those of the U. S. Geological Survey
canyon floor (PI, III, Fig. 2). At the upper Cenozoic series.
SUBGENUS // (LYMNAEIDAE: STAGNICOLA) 271
PLATE IV
Habitats of Stagnicola montanensis
Teton County, Idaho. A small

FIG. 1. Habitat of Stagnicola montanensis at the south side of Driggs,
the culvert opening in left center,
perennial stream flows westward under the highway, emerging through
Stagmcola elodes
and continues off left. The pool in the roadside ditch (foreground) was inhabitated by
(Tryon), and Aplexa hypnorum (Linnaeus) as well as S. montanensis. Photo-
(Say), Gyraulus circumstriatus
graph by D. W. Taylor, August 28, 1959.
Co., Idaho. View south down the bed
FIG. 2. Habitat of Stagnicola montanensis at Round Spring, Gooding
of Hot Creek just below the source of the small spring.
Elsewhere the stream bed is dry, but the spring
visible among the sedges growing
provides water along a distance of about 150 feet. Small pools of water are
montanensis was the only mollusk found. Photo-
in a bed of mud and cobbles and pebbles of platy latite. S.
graph by D. W. Taylor, July 26, 1959.

PLIOCENE 2785 ft -2800 ft elev. Scattered, scarce

19105. Teton Co., Wyoming. GrandTeton shells from light gray clay ft - 15 ft
National Park sheet (1954) 1:62,500. below l/2in basaltic glass sand at about
SE 1/4 sec. 25, T. 42 N., R. 116 W. 2800 ft elev. D.W. Taylor, H. E. Malde,
800 feet west, 800 feet north of south- 1956.
east corner. Exposure along irrigation 20112. Elmore Co., Idaho. Pasadena
ditch by shack. Upper claystone and VaUey quad. (1948) 1: 24,000. SE 1/4
pumicite member of Teewinot For- sec. 23, T. 6 S., R. 10 E. 600 ft W.,
mation of Love (1956b, p 90-91). D.W. 350 ft N. of SE corner. 2800 ft elev.
Taylor, J. D. Love, 1955, 1956. Clay below basaltic glass sand. H. E.
MIDDLE PLEISTOCENE Malde, 1956.
The following localitites are all in the 20120. Elmore Co., Idaho. Pasadena
Bruneau Formation, of middle Pleis- Valley quad. (1948) 1: 24,000. NW 1/4
tocene age (Malde and Powers, 1962). sec. 21, T. 6 S., R. 11 E. 2200 ft E.,
Localities 19219, 19222, and 20410 are 1750 ft S. of NW corner. 2900 ft elev.
in a brown soil horizon developed in Fossils from about 5 ft above base of
the top of the lake beds of this for- gray sand exposed on steep, west-
mation. facing slopes on E. side of Deer Gulch,
19219. Twin Falls Co., Idaho. Hagerman immediately north of tributary from
quad. (1950) 1:24,000. SW 1/4 sec. 16, east. D. W. Taylor, 1956.
T. 7 S., R. 13E.250ftE., 500 ft N., and 20404. Owyhee Co., Idaho. Oreana quad.
350 ft E. 700 ft N. of SW corner. 3180 ft (1950) 1: 24,000. SE 1/4 sec. 14, T.
elev. Brown soil. D. W. Taylor, 1955. 4 S., R. 1 W. 1425 ft N., 550 ft W. of
19222. Twin Falls Co., Idaho. Hagerman SE corner. 3025 ft elev. N. R.
quad. (1950) 1: 24,000. SW 1/4 sec. 17, Anderson, 1956.
T. 7 S., R. 13 E., 150 ft W., 1525 ft N. 20410. Twin Falls Co., Idaho. Hagerman
to 50 ft W., 1625 ft N. of SE corner. quad. (1950) 1: 24,000. NE 1/4 sec.
3180 ft elev. Brown soil. D.W.Taylor, 17 and NW 1/4 sec. 16, T. 7 S., R. 13
1955. E. 175 ft W., 1750 ft S. of NE corner
19225. Twin Falls Co., Idaho. Hagerman to 350 ft E., 1950 ft S. of NW corner.
quad. (1950) 1: 24,000. Center of 3180 ft elev. Brown soil. D.W. Taylor,
SW 1/4 sec. 16, T. 7 S., R. 13 E. 1955.
2875 ft-2940 ft elev. Sand on both sides 20472. Owyhee Co., Idaho. Castle Butte
of main gulch draining sec. 17. D. W. quad. (1948) 1: 24,000. SW 1/4 sec.
Taylor, 1955. 18, T. 4 S., R 2 E. 2350 ft E., 1500 ft
20105. Elmore Co., Idaho. Glenns Ferry N. of SW corner. 2530 ft elev. Sedi-
quad. (1948) 1: 24,000. NW 1/4 sec. 15, ments below basaltic glass sand. N.
T. 6 S., R. 10 E. 2300 ft E., 2050 ft S. R. Anderson, 1957.
of NW corner. 2800 ft elev. H. E. In the American Falls Reservoir area,
Malde, 1956. southeastern Idaho, the oldest Pleistocene
20107. Elmore Co., Idaho. Glenns Ferry formation is the Raft Formation. It is of
quad. (1948) 1: 24,000. NE 1/4 sec. 22, middle Pleistocene age, overlain by the
T. 6 S., R. 10 E. 3150 ft E. 550 ft S. of American Falls Lake beds whose base is
NW corner. 2790 ft elev. Shells from probably of Illinoian age (Love and Taylor,
light-gray to white-weathering clay 1962). Quite possibly the Raft Formation
about 10 ft below l/2in basaltic glass is equivalent to part of the Bruneau For-
sand. D. W. Taylor, 1956. mation.
20109. Elmore Co., Idaho. Pasadena
20478. Power Co., Idaho. N^1/4SW 1/4
VaUey quad. (1948)1:24,000. NE 1/4
sec. 22, T. 9 S., R. 28 E. Base of bluff
sec. 22, T. 6 S., R. 10 E. 800 ft S.,
at Snake River, in Wildlife Refuge about
1400 ft - 1100 ft W. of NE corner. 8 miles W. of Massacre Rocks on U.
SVBGENVS HINKLEYIA (LYMNAEIDAE: STAGNICOLA) 273
S. highway 30 N. Raft Formation. W. (1937) 1: 62,500. S 1/2 NE 1/4 sec.3,

J. Carr and D. E. Trimble, 1957. T. 6 S., R. 32 E. Exposure in cliff at
LATE PLEISTOCENE edge of American Falls Reservoir.
In the Glenns Ferry-Hagerman area, Sand and gravel in lower part of A-
southwestern Idaho, a few fossils have merican Falls Lake beds; or Michaud
been found in late alluvium. Locality gravel. M. L. Hopkins and others, 1957.
20413 is in a young alluvial unit not 21636. Power Co., Idaho. American Falls
formally named (H. E. Malde, written quad. (1936) 1: 62,500. NE1/4NW 1/4
communication, 1961). sec. 32, T. 7 S., R. 31 E. Exposure in
20413. Twin Falls Co., Idaho. Hagerman ditch for Michaud Flats pipeline. Older
quad. (1950) 1: 24,000. SW 1/4 sec. 16, alluvium or Aberdeen terrace deposits.
T. 7 S., R. 13E. 1650ft E., 975 ft N. of W. J. Carr, 1957.
SW corner. 2900 ft elev. Limey light 21643. Power Co., Idaho. American Falls
gray to white fine sand and silt. D. W. quad. (1936) 1: 62,500. NE 1/4 SW 1/4
Taylor, 1955. sec. 30, T. 7 S., R. 31 E. Bluff at edge
The following localities are in late of reservoir. Ten feet below base of
Pleistocene sediments in the vicinity of persistent white blocky clay. American
the American Falls Reservoir, south- FaUs Lake beds. W. J. Carr, 1958.
eastern Idaho. Overlying the Raft For- 22332. Power Co., Idaho. Michaud quad.
mation is a fluviatile gravel at the base of (1937) 1: 62,500. NW 1/4 NE 1/4 sec.
the American Falls Lake beds. The gravel I, T. 7 S., R. 32 E. 4445 ft elev. Road-
is probably of Illinoian age (Love and cut exposure. Possibly Sunbeam For-
Taylor, 1962). The upper part of the lake mation. W. J. Carr, 1960.
beds is no younger than early Wisconsin. The following localities are in an un-
The alluvial units are younger the Michaud
: named alluvial unit in Marsh Creek valley.
Gravel is early Wisconsin, deposited by This unit appears from reconnaissance by
overflow of Lake Bonneville, and the R. C. Bright and D. W. Taylor to be older
other sediments are of still later Wis- than the overflow of Lake Bonneville, hence
consin age. probably pre-Wisconsin.
19169. Power Co., Idaho. Michaud quad. 22328. Bannock Co., Idaho. Pocatello
(1937) 1: 62,500. SE 1/4 sec. 3, T. 6 sheet (1958) 1: 250,000. North side
S., R. 32 E. Shore of American Falls NE 1/4 sec. 31, T. 11 S., R. 37 E.
Reservoir. Basal gravel of American Road cut exposure on south side of
Falls Lake beds. M. L.Hopkins, 1954. Woodland road 2.6 miles west of
20474. Power Co., Idaho. American Falls Downey. R. Bright, D. W. Taylor,
quad. (1936) 1: 62,500. SE1/4NW 1/4 1959.
sec. 29, T. 7 S., R. 31 E. 4350 ft elev. 22410. Bannock Co., Idaho. Pocatello
Gravel pit exposures of older alluvium sheet (1958) 1: 250,000. SE 1/4 sec.
as mapped by W. J. Carr and D. E. II, T. 10 S., R. 36 E. Robin road west
Trimble (written communication, 1961). of Arimo. Big cut west of top of
D. W. Taylor and others, 1957. pediment and about 300 yards west of
20477. Power Co., Idaho. Rockland quad. big gravel pit. On east flank of first
(1937) 1: 62,500. SE 1/4 sec. 14, T. 9 big gully west of top of pediment. South
S., R. 30 E. 550 ft W., 200 ft N. of SE side of road in middle of cut. R. C.
corner. Rock Creek road. W. J. Carr Bright, 1959.
and D. E. Trimble, 1957. The follwing localities are in sediments
20479. Bingham Co., Idaho. American deposited in an early Wisconsin lake named
Falls quad. (1936) 1: 62,500. NW 1/4 Lake Thatcher by Bright (1960). Radio-
sec. 22, T. 5 S., R. 31 E. Two miles carbon dates (W-704, W
-855) of 32, 500 and
north of Aberdeen. W. J. Carr and 27,500 years were obtained from samples
D. E. Trimble, 1957. of fossil shells from these deposits (Rubin
21057. Power Co., Idaho. Michaud quad. and Alexander, 1960).
21143. Franklin Co., Idaho. Prestonquad. worked shore deposit of Lake Bonne-
in
(1918) 1: 125,000. NE 1/4 sec. 1, T. ville. Elev. about 5100 feet. R. C.
12 S., R. 40 E. Cut on west side of Bright, 1961.
Idaho state highway 34. Lake Thatcher The following localities are from sedi-
deposits. R. Bright, 1957. ments which may be related to a former
22396. Caribou Co., Idaho. Bancroft quad. high stand of Bear Lake, Idaho -Utah.
(1949) 1: 625,000. NE 1/4 SW 1/4 sec. 21062. Bear Lake Co., Idaho. Montpelier
29, T. 10 S., R. 40 E. Cut on west side quad. (1909) 1: 125,000. NW 1/4 sec.
of road below telephone pole and guy. 30, T. 12 S., R. 44 E. Fossiliferous
Sandy unit at base of cut, just a few sand from west side of pit. F. C.
feet above ditch. R. C. Bright, 1958. Armstrong, 1949.
23025. Caribou Co., Idaho. Prestonquad. 22429. Bear Lake Co., Idaho. Montpelier
(1913) 1: 125,000. NE 1/4 NE 1/4 quad. (1909) 1: 125,000. T. 11 S., R.
SW
1/4 sec. 9, T. 11 S., R. 41 E. Road 43 E. Two miles south of Nounan in
cut at sharp turn in road at base of road cut on west side of Road. R. C.
grade. Elev. about 5300 feet. Lake Bright, 1960.
Thatcher deposits. R. Bright, 1961. The following localities are in deposits
23027. Caribou Co., Idaho. Bancroftquad. of Lake Bonneville, Idaho-Utah. Locality
(1949) 1: 625,000. NE 1/4 NE 1/4 sec. 22359 is of rather recent age; those from
31, T. 10 S., R. 40 E. Just below the Saltair core range through much of the
gravel-capped knob about 1/2 mile west late Pleistocene according to the inter-
of Harris ranch house. Elev. about pretations by Eardley and Gvosdetsky
5340 feet. Lake Thatcher deposits. (1960).
R. Bright, 1961. 21112. Salt Lake Co., Utah, Core from
23033. Caribou Co., Idaho. Prestonquad. Saltair, depth 271 ft 4in - 271 ft 6in.
(1918) 1: 125,000. Center SW 1/4 SE A. J. Eardley, 1958.

1/4 sec. 26, T. 11 S., R. 40 E. Wawn 21120. Salt Lake Co., Utah. Core from
Hogan water well, depth 123-128 feet. Saltair, depth 428 ft 7in - 428 ft lOin.
Lake Thatcher deposits? R. C. Bright, A. J. Eardley, 1958.
1961. 21122. Salt Lake Co.. Utah. Core from
23037. Caribou Co., Idaho. Prestonquad. Saltair, depth 435 ft Hin - 436 ft lin.
(1918) 1: 125,000. Center N 1/2 sec. A. J. Eardley, 1958
12, T. 11 S., R. 40 E. About 1/4 mile 21124. Salt Lake Co., Utah. Core from
south of elev. 5376 on Idaho state high- Saltair, depth 463 ft 10 l/2in - 464 ft
way 34, in road cut on west side of road. l/2in. A. J. Eardley, 1958.
Elev. about 5345 feet. Lake Thatcher 22359. Box Elder Co., Utah. SW 1/4 sec.
deposits. R. Bright, 1961. 34, T. 9 N., R. 3 W. Depth 12in - 24in,
23061. Caribou Co., Idaho. Prestonquad. along Bear River. Post-Provo. J. H.
(1918) 1: 125,000. SW 1/4 SW 1/4 Feth, 1953.
SW 1/4 sec. 8, T. 11 S., R. 41 E. Road 22395. Franklin Co., Idaho. Prestonquad.
cut at north side of road. About 1/5 (1918) 1: 125,000. NE 1/4 NW 1/4
mile west of Bitten' s ranch on divide NE 1/4 sec. 28, T. 15 S., R. 39 E. About
separating East Fork Whisky Creek 1.3 miles duewestof center of Preston.
from West Fork Trout Creek. Elev. In road cut just south of highway 89 at
about 5270 feet. Lake Thatcher intersection of road to Refuge Club.
deposits. R. C. Bright, 1961. Prominently cross-bedded sand. Provo
23066. Caribou Co., Idaho. Prestonquad. sediments. 4702 ft ± elev. R. Bright,
(1918) 1: 125,000. NW 1/4 SW 1/4 1958. Shells from this locality were
SE 1/4 sec. 24, T. US., R. 40 E. About dated at 13,900 -400 radiocarbonyears
300 yards west of basalt cliff in sand by the U. S. Geological Survey labo-
pit on side of prominent knob. Some ratory (sample W-899).
probable Lake Thatcher material re- 23069. Caribou Co., Idaho. Prestonquad.
(1918) 1: 125,000. SW 1/4 SW 1/4 356-362 (p 360).

NE 1/4 sec. 24, T. lis., R. 40 E. About BRIGHT, R. C, 1960, Geology of the
50 yards north of Jay Turner's house Cleveland area, southeastern Idaho. M.
in tan sands. Elev. about 5100 feet. S. Thesis, Univ. Utah, Salt Lake City.
R. C. Bright, 1961. BURCH, J. ., 1959, Chromosomes of
The following locality is presumably of aquatic pulmonate snails (Basom-
late Pleistocene age: matophora). Amer. Malacol. Union Ann.
19192. Humboldt Co., Nevada. Antler Peak Rep., 1958: 9-10.
quad. (1940) 1: 625,000. Sec. 18, T. , 1960a, Chromosome morphology
33 N., R. 43 E. 5050' elev. at range of aquatic pulmonate snails (Mollusca:

front. Trout Creek valley at Marigold Gastropoda). Trans. Amer. Microsc.
Mine. Shells from red-brown clay from Soc, 79: 451-461.
drill hole in valley floor. R. J. , 1960b, Chromosome studies of
Roberts. aquatic pulmonate snails. The Nucleus,
3: 177-208.
ACKNOWLEDGMENTS
COOPER, J. G., 1872, On shells of the
For permission to study specimens west slope of North America. No. .
under their charge we are grateful to R. California Acad. Sei. Proc, ser. 1, 4:
T. Abbott, Academy of Natural Sciences, 171-175.
Philadelphia; G. D. Hanna and A. G.Smith, CRABB, E. D., 1927, The fertilization
California Academy of Sciences; H. A. process in the snail, Lymnaea stagnalis
Rehder, U. S. National Museum, and H. oppressa Say. Biol. Bull., 53: 67-97,
G. Rodeck, University of Colorado pi. 1-6.
Museum. We wish to acknowledge also EARDLEY, A. J. and VasylGVOSDETSKY,
the aid of Linda L. Bush, Department of 1960, Analysis of Pleistocene core
Biology, Western Michigan University for from Great Salt Lake, Utah. Bull. Geol.
preparing many of the cytological slides, Soc. Amer., 71: 1323-1344, 1 pi.
and to R. Bright, Department of HANNIBAL, Harold, 1912, A synopsis of
Geology, University of Minnesota, for the Recent and Tertiary freshwater
advice on the statistical analysis. Mollusca of the Californian Province,
Drawings of shells are by Mrs. Elinor based upon an ontogenetic classification.
Stromberg, U. S. Geological Survey, except Proc. Malacol. Soc. London, 10: 122-
for text Figure 1 which is by Miss Roberta 211, pi. 5-8.
Wigder, U. S. Geological Survey. HEMPHILL, Henry, 1890, New forms of
western limniades. Nautilus, 4: 25-27.
REFERENCES HUBENDICK, Bengt,1951, Recent Lymnae-
ARCHIE, V. E., 1942, The histology and idae; their variation, morphology, tax-
developmental history of the ovotestis onomy, nomenclature, and distribution.
of Lymnaea stagnalis. Ph.D. Thesis, Kungl. Svenska Vetensk.-akad. Handl.,
Univ. Wise, Madison. ser. 4, 3 (1): 1-223, pi. 1-5.
BAKER, F. The Lymnaeidae of
C, 1911, HUNTER, W. R., 1953, The condition of
North and Middle America, recent and the mantle cavity in two pulmonate snails
fossil. Chicago Acad. Sei., Spec. living in Loch Lomond. Proc. Royal
Publ. xvi + 539 p, 58 pi.
3. Soc. Edinburgh, sec. (Biol.), 65(2):
, 1913, A new Lymnaea from 143-165.

Montana. Nautilus, 26: 115-116. HUSTED, Ladley and P. R. BURCH, 1953,
, 1928, The fresh water Mollusca The chromosomes of the polygyrid snail
of Wisconsin. Part I. Gastropoda. Allogona profunda. Virginia J. Sei., 4:
Wise. Geol. Natur. Hist. Surv., Bull. 70 62-64.
(1): i-xx,1-507, pi. 1-28. INABA, Akihiko, 1950, Chromosome
BRETSCHNEIDER, L. H., 1948, Insemi- numbers in some pulmonate moUusks
nation in Limnaea stagnalis L. Proc. (in Japanese). Japanese J. Genetics,
Koinkl. Nederl. Akad. Wetensch., 51: 25: 58-59.
, 1959, Cytological studies in chromosomes dans les deux lignées

mollusks. .
A chromosome survey germinales du gastéropode hermaphro-
in the stylommatophoric Pulmonata. J. dite Limnea stagnalis {va.riété rhodani).
Sei., Hiroshima Univ., Ser. , Div. 1, Ibid., 41: 693-697.
18: 71-93, pl. 1-3. PERROT, Jean-Louis and Max PERROT,
INABA, Akihiko and Hidao TANAKA, 1953, 1938, Note sur les chromosomes de
Studies on the chromosome numbers oí cinq espèces de Limnées. C. R. Séan.
some freshwater gastropods. Ibid., 14: Soc. Phys. Hist. Nat. Genève (Suppl.
213-220. Arch. Sei. Phys. Nat.), 55(3,xi/17): 92-
LA COUR, L., 1941, Acetic -orcein: A 93.
new stain -fixative for chromosomes. RUBIN,Meyer and Corrinne ALEXANDER,
Stain Technology, 16: 169-174. 1960, .
S. Geological Survey radio-
LARAMBERGUE, Marc de,Etude
1929, carbon dates V. Amer. J. Sei., Radio-
cytologique de l' autofécondation chez carbon Suppl., 2: 129-185.
Limnaea auricularia L. C. R. Sean. TAYLOR, D. W., 1956, Pliocene mol-
Acad. 1027-1029.
Sei., Paris, 189(xii/2): lusks from Jackson Hole, Grand Valley,
LE CALVEZ, Jean and Philomène and Star Valley, Wyoming and Idaho.
CERTAIN, 1950, Données caryologiques Wyoming Geol. Assoc. Guidebook, 11:
sur quelques Pulmones basommato- 123-125.
phores. Ibid., 231(16): 794-795. 1960, Late Cenozoic moUuscan
,
LINVILLE, Henry R., 1900, Maturation faunas from the High Plains. U. S.
and fertilization in pulmonate gastro- Geol. Surv. Prof. Paper 337, 94 p, 4 pl.
pods. Bull. Mus. Compar. Zool., Harvard TRIMBLE, D. E., and CARR, W. J., 1961,
College, 35: 213-248. Late Quaternary history of the Snake
LOVE, J. D., 1956a, New geologic for- River in the American Falls region,
mation names in Jackson Hole, Teton Idaho: Geol. Soc. Amer. Bull., 72: 1739-
County, northwestern Wyoming. Bull. 1748, 1 pl.
Amer. Assoc. Petroleum Geologists, WALTER, Harold J., 1959. The mor-
40: 1899-1914. phology oi Lymnaea emarginata serrata
1956b, Cretaceous and Tertiary
,
Haldeman with remarks on the system-
stratigraphy of the Jackson Hole area, atic s of lymnaeids. Amer. Malaeol.
northwestern Wyoming. Wyoming Geol. Union Ann. Reps., 1958.
Assoc. Guidebook 11: 76-94. 1961, The morphology of Sia^m-
,
LOVE, and TAYLOR, D. W., 1962,

J. D., cola emarginata serrato (Haldeman) and
Faulted Pleistocene strata near Jack- its relation to lymnaeid systematic s.
son, northwestern Wyoming: U. S.Geol. Ph.D. Thesis, Univ. Mich., 274 p, pl.
Survey Prof. Paper 450-D: 136-139. 23.^2,
MALDE, H. E., and POWERS, H. A., WALKER, Bryant, synopsis of
1918, A
1962, Upper Cenozoic stratigraphy of the classification freshwater
of the
western Snake River plain, Idaho. Bull. Mollusca of North America, north of
Geol. Soc. Amer., 73: 1197-1219, 1 pl. Mexico, and a catalogue of the more
NEWCOMER, E. H., 1953, A new cytolo- recently described species, with notes.
gical and histological fixing fluid. Mich. Univ. Mus. Zool., Misc. Publ. 6:
Science, 118: 161. 1-213.
PERROT, Jean-Louis, 1930, Chromo- ZILCH, Adolf, 1959-60, Euthyneura,
somes hétérochromosomes chez les
et in Otto H. Schindewolf, Handbuch der
gastéropodes pulmones. Revue Suisse Paläozoologie, v. 6, Gastropoda, by
de Zoologie, 37: 397-434, Pl. 5-7. Wilhelm Wenz, pt. 2. Berlin (Born-
, 1934, A propos du nombre des traeger). xii + 834 p.
^Ôbtainable in microfilm form from Univ. Microfilms, Ann Arbor, Michigan, U.S.A.
SUBGENUS /]^/(:£ (:
ZUSAMMENFASSUNG
STAGNICOLA) 277
SÜSSWASSERSCHNECKEN DER UNTERGATTUNG HINKLEYIA (LYMNAEmAE:

STAGNICOLA) AUS DEN WESTLICHEN VEREINIGTEN STAATEN
Die Untergattung Hinkleyia, die bisher nur aus einer einzigen Art, Stngnicola caperata
(Say) bestand, wird hier um 2 Arten erweitert. Und zwar fügen wir auf Grund gewisser
konchyliologischer sowie anatomischer Merkmale S. montanensis (Baker) hinzu und,
auf Grund konchyliologischer Merkmale allein, S./)¿Zsbr>'¿ (Hemphill). Neue Erkenntnisse
gestatten nun eine revidierte Diagnose von Hinkleyia, welcher, obwohl sie einige kenn-
zeichnende Eigenheiten aufweist, doch nur subgenerischer Rang zusteht.
Sechzehn Exemplare einer konservierten Serie von S. montanensis wurden seziert.
Anatomisch waren sie S. caperata so ähnlich, dass keinerlei Unterscheidungsmerkmale
mit Sicherheit erkannt werden konnten.
Die beiden Arten kann man an ihren Schalen erkennen. S. montanensis hat gewöhnlich
eine schmälere und mehr langgestreckte Form als S. caperata, die eher geschwollen
ist, aber diese Kennzeichen überschneiden sich, so dass die einzigen immer verläss-
lichen Charaktere in der Oberfläche und Skulptur der Schale zu finden sind. Verhält-
nismässig auffällige spiralig verlaufende Periostrakumleistchen kennzeichnen S. caper-
ata, während eine glänzende, mit feinstem, spiral angeordnetem Sichelmuster verse-
hene Oberfläche, ohne Leistchen, für S. montanensis charakteristisch ist.
Die anatomischen Merkmale, die diese beiden Arten mit typischen amerikanischen
Stagnicola gemein haben und die, soweit bekannt, ausschliesslich bei diesen vorkommen,
sind: ein gut entwickelter vaginaler Sphinktermuskel; ein grosses Divertikulum am
proximalen Ende der oberen Prostata; eine verhältnismässig dicke und durze Pe-
nisscheide; ein relativ kurzer bilateral symmetrischer Penis mit einem muskulären
"Knoten" in seiner ungefähren Mitte; und eine gut entwickelte Serie von deutlich 2-
zackigen Seitenzähnen an der Radula,
Hinkleyia (d.h. S. montanensis und S. caperata) unterscheidet sich von typischen
Stagnicola durch einen etwas verschieden und weniger stark entwickelten vaginalen
Sphinkter, einen bemerkenswert dicken Vas deferens und einen kürzeren Penis, der
einen weniger prominenten, mehr distal gelegene "Knoten" hat. Bemerkenswert ist
ferner die sich verjüngende, distal vom Knoten gelegene Partie des Penis, die kurz und
dick ist und sich vom starken proximalen Teil viel weniger klar absetzt als dies bei
Stagnicola s.S. der Fall ist. Beide Arten haben auch eine ungewöhnlich kurze und starke
Penisscheide, eine Prostata von besonderem Bau und einen in seinem unteren Teil
ungewöhnlich robust entwickelten Spermathekdukt. Asserdem zeigen sie in der Be-
schaffenheit und Pigmentierung ihrer gesamten Anatomie Ähnlichkeiten, welche dazu
beitragen sie weiterhin von anderen genügend bekannten Lymnaeiden abzusondern. Auch
sind ihre Körper stark pigmentiert und, im lebenden Zustand, Fuss und Tentakeln
recht schlank.
Es wurde ferner festgestellt, dass S. montanensis bei einer Schalenlänge von 6-7 mm
geschlechtsreif wurde. Es wird angenommen, dass der noch unbekannte Laich dieser
Art dem eigenartigen von S. caperata gleichen dürfte, der sich durch die relative Dicke
der individuellen EihüUen und der unscheinbaren dünnen äusseren Tunika der Laichmasse
auszeichnet.
Weiters wurde ermittelt, dass eine Art, die als Untergattung Nasonia unter Stagnicola
eingereiht worden war, nahe mit Fossar¿a verwandt ist und daher nicht mit
Die Möglichkeit einer näheren Beziehung zwischen S. arciz'ca und ^;>>
/^.
statt Sia^g^-
nicola s.S. wurde in Betracht gezogen aber auf Grund von neuerem anatomischen
Beweismaterial abgelehnt.
Die haploiden Chromosomenzahlen von S. montanensis und S. caperata betragen 18;
die nachgewiesene diploide Chromosomenzahl von S. montanensis ist 36. Diese Zahlen
sind für Basommatophoren im allgemeinen kennzeichnend. In den Einzelheiten der
Gametogenese, einschliesslich der monozentrischen Natur der Chromosome scheint
Hinkleyia den anderen Lymnaeiden zu gleichen. Normalerweise verläuft die Spermato-
genese in 6 spermatogonischen Teilungen, auf welche 2 meiotische folgen. Es kommen
daher die spermatogonischen Zellen in Büscheln zu 2, 4, 8, 16 und 32 vor, die primären
Spermatozyten in Büscheln zu 64, die sekundären in solchen zu 128 und die Spermatiden
sowie der Sperm in Büscheln zu 256.
Von besonderem zytologischen Interesse in der Gametogenese von S. caperata und
S. montanensis ist die Anwesenheit von 5-7 grossen Chromatinkörpern in jedem Zellkern
während der frühen Prophase der ersten meiotischen Teilung der Spermatogenese, wie
sie für lymnaeide Schnecken noch nie beobachtet wurde. Diese Chromatinkörper stellen
vielleicht ein weiteres Kennzeichen für die nahe Verwandtschaft dieser beiden Arten
dar.
In den westlichen Vereinigten Staaten hat S. montanensis eine weite Verbreitxing im
östlichen Teil des Entwässerungsgebietes des Columbiailusses und dem nördlichen
des "Great Basin", einem Gebiet, das sich von Westmontana und Utah bis ins süd-
liche Idalio und Mittelnevada erstreckt. Man findet die Art in für Lymnaeiden ganz
ungewöhnlichen Wohnplätzen, nämlich in Quellen und klaren Bergbächen. Das grössere,
beinahe das gesamte nördliche Nordamerika umfassende Verbreitungsgebiet von S.
caperata schliesst dasjenige von S. montanensis ein. Man findet S. caperata in Bewäs-
serungsgräben und schlammigen, unbeständigen Gewässern in denen S. montanensis
nicht lebt. Durch Beschreibungen spezifischer Standorte und durch Landkarten welche
die geographische Verbreitung (lebend und fossil) anzeigen, werden Anhaltspunkte
für nicht morphologische Unterschiede zwischen den beiden Arten gegeben. Die be-
obachtete geographische Trennung der Arten ist teilweise auf ihre verschiedenartigen
ökologischen Bedürfnisse zurückzuführen, wahrscheinlich aber auch zum Teil auf die
geologische Vorgeschichte des Gebietes. S. pilsbryi istnur durch die Originalsammlung
und aus einem einzigen Fundort im westlichen Utah bekannt; ökologische Angaben fehlen.
RESUME
MOLLUSQUES FLUVIÁTILES DU SOUS-GENRE HINKLEYIA
(LYMNAEIDAE: STAGNICOLA) DES ETATS UNIS OCCIDENTAUX
Le sous-genre Hinkleyia, ne comprenant jusqu'à présent que l'espèce Stagnicola

caperata (Say), est augmenté ici de deux espèces: S. montanensis (Baker) et S. pilsbrvi
(Hemphill), la première d'après de critères conchyliologiques et anatomiques, et la
seconde uniquement d'après des caractères conchyliologiques. Ces nouvelles données
entraînent une révision de la diagnose de Hinkleyia qui, quoique possédant plusieurs
traits particuliers, ne mérite pourtant qu'un rang subgénérique.
Seize spécimens d'une série de S. montanensis provenant de l'Idaho furent disséqués.
Anatomiquement ils ressemblaient de si près à S. caperata, qu'aucun caractère dis-
tinctif ne put être trouvé.
Les 2 espèces se distinguent par leur coquille. Celle de S. montanensis est d'habitude
plus étroitement allongée que celle de S. caperata, plus gonflée; mais ces caractères
sont variables et les seuls critères sûrs et constants se trouvent dans la sculpture de
la surface. Des crêtes périostracales spirales relativement proéminentes déterminent
la diagnose de S. caperata tandis qu'une surface luisante, dépourvue de telles crêtes,
mais garnie de séries spirales de minuscules croissants, détermine celle de S. mon-
tanensis.
Les caractères anatomiques, particuliers, communs aux stagnicoles typiques améri-
caines et à ces 2 espèces sont: un sphincter vaginal bien développé; un grand diverticule
attenant au bout proximal de la prostate supérieure; une gaine péniale relativement
épaisse et courte; un pénis relativement court à Symmetrie bilatérale, ayant un "noeud"
musculaire à peu près à mi-longueur, et une série bien développée de dents radulaires
latérales clairement bicuspidées.
Comparés aux Stagnicola typiques, les Hinkleyia (tout-au moins S. montanensis et
S. caperata) montrent un développement un peu différent et moins proéminent du
sphincter vaginal; un vas deferens remarquable par son épaisseur et un pénis plus
court, ayant un "noeud" moins proéminent et de position plus distale. Notons aussi que
la portion effilée du pénis, distale par rapport au noeud, est courte, forte et moins
individualisée par rapport à la portion proximale que chez les Stagnicola proprement
dits. Les 2 espèces possèdent aussi une gaine péniale remarquablement courte et
forte, une prostate de structure particulière et un conduit spermatique plus robuste
que d'ordinaire dans sa partie inférieure. Dans leur pigmentation et leur anatomie
générale elles montrent des similitudes additionelles servant aussi à les séparer des
autres Lymnaeidés suffisamment connus. Leurs corps sont fortement pigmentés. A
l'état vivant, les pieds et tentacules sont relativement sveltes.
A une taille de 6 à 7 mm les S. montanensis avaient atteint la maturité sexuelle.
Nous supposons que la ponte encore inconnue de cette espèce ressemblera de près
à celle de S. caperata qui est très à-partau fait de l'épaisseur relative des enveloppes
SUBGENUS HINKLEYIA (LYMNAEffiAE: STAGNICOLA) 279
de chaque oeuf et du fait de la minceur de la tunique extérieure de la masse.

L'examen anatomique a aussi montré qu'une espèce rangée dans le sous-genre
Nasonia du genre Stagnicola est étroitement apparentée à Fossaria et, par conséquent,
pas à Hinkleyia. La possibilité d'une affinité de S. árctica avec Hinkleyia plutôt
qu'avec le sous-genre Stagnicola a été examinee, mais rejetée à la suite de nouvelles
recherches anatomiques.
Le nombre haploïde de chromosomes de S. montanensis et de S. caperata est de 18,
Le nombre diploïde de S. montanensis est de 36. Ces nombres sont caractéristiques
pour les mollusques basommatophores en général. Les détails de la gamétogénèse, y
compris la nature monocentrique des chromosomes, ne semblent pas différer de ceux
vus chez les autres lymnaeidés. Pendant la Spermatogenese il y a, normalement, 6
divisions spermatogoniales, suivies de 2 divisions meiotiques. Les cellules sperma-
togoniales se trouvent donc en bouquets de 2, 4, 8, 16 et 32; les spermatocytes primaires
en bouquets de 64; les spermatocytes secondaires en bouquets de 128 et les sperma-
tides et spermatozoïdes en bouquets de 256.
Fait intéressant spécial dans la gamétogénèse de S. montanensis et de S. caperata:
il y a 5 à 7 grands corps chromatiniens dans chaque noyeau cellulaire pendant les
premiers stades de la prophase de la première division meiotique de la Spermatogenese;
de pareils corps de chromatine n'ont été signalés chez aucun autre Lymnaeidé. Ils
constituent peut-être un autre trait d'affinité de ces 2 espèces.
S. montanensis est largement réparti dans l'ouest des Etats Unis d'Amérique, dans
le bassin hydrographique oriental du fleuve Columbia et le versant nord du "Great
Basin", un territoire allant du Montana occidental et de l'Utah jusqu'à l'Idaho méridional
et le centre du Nevada. L'espèce habite sources et ruisseaux de montagne clairs,
habitat inusité pour un lymnéide. L'aire de répartition de S. caperata, plus vaste,
couvre la majeure partie de l'Amérique du Nord septentrionale, incluant celle de S.
montanensis. S. caperata peuple des habitats tels que fossés d'irrigation et eaux
boueuses saisonnières où S. montanensis ne se trouve pas. Ces différences d'habitat,
et des cartes géographiques montrant leurs distributions actuelles et passées con-
firment la distinction des deux espèces que nous avait enseignée la morphologie. La
séparation géographique observée entre ces 2 espèces est partiellement liée à leurs
besoins écologiques différents, mais probablement aussi, en partie, à l'histoire gé-
ologique du territoire. S. pilsbryi n'est connu que par le matériel original, et d'une
seule localité dans l'Utah occidental, sans aucune donnée écologique.
RESUMEN
CARACOLES FLUVL\LES DEL SUBGÉNERO HINKLEYIA (LYMNAEmAE:

STAGNICOLA), DEL OESTE DE ESTADOS UNmOS
El subgénero Hinkleyia, formado por una sola especie, Stagnicola caperata (Say), se
amplía ahora por la adición de S. montanensis (Baker), en base a criterios anatómicos
y conquiliológicos en la primera, y exclusivamente conquiliológicos en la segunda. Los
nuevos datos obtenidos en este estudio permiten revisar la diagnosis de Hinkleyia, que
posee varias características distintivas pero insuficientes para conferirle categoría
genérica.
La disección de 16 ejemplares preservados de 5. montanensis de Idaho reveló
tal similitud con S. caperata que no ha sido posible distinguirlas anatómicamente.
Las dos especies pueden distinguirse por la concha. En general, S. montanensis es
más estrecha y alargada, y S. caperata más ancha, pero, como estes caracteres se
sobreponen, sólo quedan como caracteres diagnósticos seguros la textura y la escultura
superficiales. Así, S. caperata se caracteriza por costillas espirales del periostraco
relativamente conspicuas y S. montanensis por una superficie lustrosa sin esa escultura,
pero con series de minúsculos crecientes ordenadas en espiral.
Los caracteres anatómicos comunes a estas dos especies y a las típicas Stagnicola
americanas y, por lo que sabemos, exclusivos de ellas, son: esfínter vaginal bien
desarrollado; grande divertículo anexo a la extremidad proximal de la parte superior
de la próstata; vaina penial comparativamente corta y más bien robusta; pene corto,
bilateralmente simétrico y con un "nudo" muscular en la parte media; y una serie
bien desarrollada de dientes radulares laterales distintamente bicúspides.
En comparación con las Stagnicola típicas, Hinkleyia (es decir, S. montanensis y S.
caperata) tiene el esfínter vaginal menos desarrollado, el vas deferens notablemente
280
robusto
SVBGEliUS HINKLEYIA (: STAGNICOLA)
el pene más corto con un "nudo" menos prominente situado algún tanto más
distalmente. Especialmente notable es la porción atenuada del pene, distal al "nudo",
que es corta y robusta y mucho menos distintamente demarcada de la porción proximal
gruesa que en las Stagnicola típicas. Además, ambas especies se caracterizan por la
vaina penial más corta y gruesa que lo común, la conformación particular de la
próstata y la notable robustez de la parte inferior del ducto de la espermateca. Otros
caracteres que sirven para separarlas de otras especies de limneidos bien conocidas
son la fuerte pigmentación del cuerpo y la delgadez relativa del pie y de los tentáculos
en el animal vivo.
Se descubrió también que S. montanensis llega a la madurez sexual al alcanzar su
concha la longitud de 6 o 7 mm. Puede conjeturarse que la masa ovígera, desconocida
en esta especie, demuestre ser similar a la de S. caperata, la cual se singulariza por
el relativo espesor de la envoltura de los huevos individuales y por la delgada e incon-
spicua túnica externa de la masa ovígera.
La evidencia anatómica indica que una especie con anterioridad asignada a Stagnicola
bajo el subgénero Nasonia se relaciona estrechamente con Fossaria y por lo tanto no
es aliada a S. montanensis y S. caperata. Una posible afinidad especial de S. árctica
con Hinkleyia, 'más bien que con el subgénero Stagnicola, fué considerada pero rechazada
en base a los nuevos datos anatómicos.
El número haploide de cromosomas de S. montanensis y S. caperata es 18; el
número diploide de S. montanensis es 36. Estos números son característicos de los
basomatóforos en general. Los detalles de la gametogénesis, incluyendo la naturaleza
monocéntrica de los cromosomas, parecen ser los mismos que en otros limneidos.
Durante la espermatogénesis ocurren normalmente 6 divisiones espermatogónicas,
seguidas por 2 meióticas. Por lo tanto, los elementos germinales aparecen en racimos
de 2, 4, 8, 16 y 32 para las espermatogonias, 64 para los espermatocitos primarios,
128 para los espermatocitos secundarios, y 256 para las espermátidas y los esperma-
tozoides.
De especial interés citológico en la gametogénesis de S. montanensis y S. caperata,
de lo que aun no se tenia noticia en otros limneidos, es la ocurrencia de 5 a 7 grandes
cuerpos de cromatina en cada núcleo en el principio de la profase de la primera división
meiótica de la espermatogénesis. Estos cuerpos cromáticos son quizá otro carácter
demostrativo de una estrecha relación entre las dos especies.
S. montanensis tiene amplia distribución en el oeste de Estados Unidos, en la región
oriental del Río Columbia y septentrional de la Gran Cuenca, desde el oeste de Montana
y Utah hasta el sur de Idaho y Nevada central. Su habitat en claras corrientes de
montaña y manantiales es desusado para Lymnaeidae. S. caperata se encuentra en
casi toda Norteamérica septentrional e incluye el area de S. montanensis en su distri-
bución. Se encuentra en zanjas de irrigación y en aguas lodasas temporarias donde
S. montanensis está ausente. Descripciones de habitats específicos y mapas de
distribución geográfica (fósil y viviente) de las dos especies suministran datos sobre
las diferencias no morfológicas entre las especies. La separación geográfica observada
se debe, en parte, a las diferencias de exigencias ecológicas, pero probablemente
también a la historia geológica de la región. S. pilsbryi se conoce solamente por el
material original de una localidad en Utah occidental y no se dispone de datos ecoló-
gicos respecto a esa especie.
. . , . .
.
ñORFORk HINKLEYIA (LYMT:ÂEmAE: STAGNICOLA EA
. .
)
rata
S.
,
pilsbryi
(Say)
.
,
Hinkleyia
(Hemphill)
, ,
S.
-
.
Hinkleyia
montanensis,
S. caperata
,
..,-
S.
,
,
( Stagnicola
montanensis
cape-
-
, . ,
, , . . --
, - TAYLOR, WALTER AND BURCH 281
-.
-
S. montanensis -
S. caperata
;;
;
] S.
,: -
montanensis
S. caperata
. ,, , , -
Stagnicola ,
-
-
, , ,, . , , -
caperata
Hinkleyia (. -. S. montanensis
.., ,.
S. )
, ,, , S. m.ontanensis
.
. -
,
6 7
,.
S. caperata,
-
.
,
nicola
18;
Stagnicola
S. montanensis
S,
.
^,,
árctica
.
36
S.
,
Nasonia,
montanensis
Hinkleyia
S. m.ontanensis
, S. caperata.
S.
-
caperata
.-
-
KFossaria
Stag-
, .
. -
6
montanensis
5
-
,
,, S.
7
256.
caperata-,
64,
2
-
2, 4,
-
8, 16 32,
128
, S.
-
. ,.
, S. m.ontanensis
'
(.) . S. caperata
S.
,
,-
montanensis.
S.
.
m,ontanensis
S. pilsbryi
,
,
.-
- - -
.
THE DISTRESS SYNDROME IN TAPHIUS GLABRATUS (SAY) AS A REACTION
TO TOXIC CONCENTRATIONS OF INORGANIC IONS
Harold W. Harry^ and David V. Aldrich^
ABSTRACT
In very low concentrations of toxic materials this snail shows normal behavior, -
tending its body out of the shell, moving about, feeding and renewing its pulmonary air •
bubble at the surface of the water. In higher concentrations of toxicants the snailV
remains retracted in the shelL Between the concentration ranges of toxicants which
produce retraction and those allowing normal behavior is a range of concentrations
which induces a condition termed distress The distressed snail is extended but unable
.
to attach its foot, hence unable to move, feed or breathe atmosperic air. There is
deterioration of the tentacles and elimination of sand grains from the stomach, but
ciliary action and heart beat seem unaffected. Snails exposed for less than 24 hours
*usually recover.
Twenty-two ions were tested of which \^ produced distress. Ag, Cd and Cu ions
did so between 0.050 and 0.100 ppm; Mn and Co ions only at 20 to 150 ppm and 30 to
300 ppm respectively. Eight other ions were found to produce distress at concentrations
intermediate between these extremes (Zn, Al, Ni, Ba, CrO^, Pb, .). The non-
toxic range of concentrations of the remaining 9 ions is reported: Sn, Au, Sr, Li,
M0O4, 27, up to 10 ppm; F up to 100 ppm and W0O4 and SO3 up to 150 ppm. The
extent of the range of concentrations which produces distress is directly proportional
I
Ito the minimal concentration of an ion which will produce sustained retraction. The
distress syndrome may be identical with relaxation produced by menthol and other
I
¡organic compounds.
In the search for molluscicides to (Australorbis glabratus of authors)3 in

use against pulmonate snails which serve Puerto Rico, we observed another type of
as the intermediate hosts of trematodes, response to toxicants. This was termed
the sustained retraction of the animal in- the "distres s synd rome" (Harry et al.,
\to the shell has been used as a criterion 1957,1958). It was evoked by concen-
Ifor determining toxic concentrations trations of ions too low to produce re-
(Nolan et. al., 1953). It is generally nec- traction. These observations led to a
essary to transfer the snails to a non- systematic study of the production of this
toxic medium to determine whether theyi syndrome by very low concentrations of
subsequently show signs of life or decay. I a variety of ions.
During studies on the relation of water
quality to the ecology of /)/5glabratus
^Rice University, Houston, Texas, U. S. A.
2u. S. Fish and Wildlife Service, Ft. Crockett, Galveston, Texas, U. S. A.
^For further particulars on the no menc lato rial relationships

of the planorbids acting
as intermediate hosts of Schistosoma mansoni, the reader is referred to Barbosa
et al. (1961, Ann. Mag. Nat. Hist., Ser. 13, 4: 371-375), Wright (1962, Bull. Zool.
Nomencl., pt. 1, 19: 39-41) and Walter (1962, Malacologia, 1: 115-137). ED.
(283)
,
284 HARRY AND ALDRICH
MATERIALS AND METHODS mum quality of natural waters in which T. I
glabratus was found to occur in Puerto

Glas^âxêûsed experiments
in the Rico. (Harry et al., 1957).
was washed in dilute HCl^ and rinsed in
distilled water that had been passed RESULTS
through an ion exchange filter ("De-
eminizer", Aloe Co.). The latter was A normally behaving snail extends 1
done because the water from our metal from the aperture of the shell and attaches
still contained 0.050 to 0.070 parts per itself to the surface of the bowl with its foot.
millón of copper, an amount sufficient to It is usually moving about and quite capable
produce the distress syndrome. The of feeding and visiting the surface to ex-
filtered water contained l ess J han OjQlO change the air in the pulmonary cavity.
ppni_ copper, which was the limit of the The typically distressed snail lies on
imethod of analysis (Huff, 1948), and which jthe bottom of the bowl with the ephalopedal
¡was innocuous to the snails. The distilled, Imass extended. It is unable to attach the
^.demineralized water (DDW) was also used foot to the substrate althoughit frequently
to prepare all stock solutions, the test di- applies the anterior end momentarily, in an
lutions and the controls. Pyrex laboratory xapparent attempt to do so. After several
finger bowls (300 ml capacity) containing hours the tentacles usually become swollen
200 ml of the various dilutionsof the stock at their bases, and show extensive sloughing
solutions were used for the tests. Adult ôf cells distally. Movement of the foot
snails recently collected in the field were gradually becomes more feeble and less
/placed in DDW for half an hour before they frequent, but at later stages a series of
'were transferred to the test solutions. spasmodic contractions of the body stalk
Five snails of about 4-4 1/2 suture whorl sometimes seen. Although muscular
\size were placed in each bowl of the test action may later cease, ciliary activity on
/solutions and in the control bowl of DWW. ttie
external surface of the body and in the
(AH tests were made at room temperature ulmonary cavity seems unaffected, at
1(250 - 30° C). All ions studied (22) were least during the first 24 hours of exposure.
tested in the concentrations shown in Fig. The heart continues to beat while the snail
1, up to and including 10.0 ppm. In ad- is in distress, though at apparently a
dition, certain ions, as noted, were studied slower rate. There is a tendency for the
in those concentrations above 10.0 ppm distressed snail to eliminate most of the
listed in Fig. 1.The results were recorded jsand grains from the stomach by defe-
at the end of 24hours of exposure, and all Ication. Normal snails, once supplied with
experiments were repeated at least once. sand, retain the grains in the stomach for
All stock solutions were prepared weeks, if deprived of sand in their ex-^
from reagen t grade chemicals. The cat- ternal environment.
ions tested were prepared from the fol- Like the retracted snails, those in
lowing compounds: LÍSO4 H2O; 12(80^) distress are unable to feed, or crawl to thejj
•I8H2O; MnS04-H20; FeS04.7H20; FeClg surface to renew the air bubble in the (
•6H2O; NÍS04-6H20; C0SO4.7H2O; CUSO4 pulmonary cavity. There is no manifest

•5H2O; ZnS04-7H20; SrCl2-H20; AgS04 ;
difference in the distress syndrome as
3CdS04-8H20; SnCl2-2H20; BaCl2; AUCI3 (produced by one ion and that produced by
.HCL.3H2O; and Pb(N03)2. Solutions of I
another. Once distressed, snails may
the anions were made from: NaF; Na2S03; remain in this state for several days, if
24; 227;
.2H2O.
NaMo04; andNaW04 left in the medium which produced the phe-
nomenon. Snails which have been dis-
An artificial _stan dard reference tressed 24 hours or less will usually re-
waterJ^RW), used m some test, was based cover within a few hours if transferred to
on the formula of Freeman (1953). We a large volume of non-toxic medium, such
modified his formula to simulate the opti- as DDW. Attachment of the foot is fol-
DISTRESS SYNDROME IN TAPHIUS GLABRATUS 285
lowed by locomotion and the snail can again Fig. 1 shows results obtained
the
feed and crawl to the surface to "breathe". from the _13 more tox ic With the ex-
ions.
/The tentacles regenerate within a few ception of lead and divalent iron, the effect
'
weekSo of the various concentrations was well de-
In each of the 22 ions tested, the com- fined. In 5.0 and 10.0 ppm of lead, most
panion ion present was known to be non- of the snails were normal, but one or two
toxic at the concentrations used, from the showed sustained retraction. Divalent iron
|report of Deschiens (1954) and our own un- produced the same mixed reaction in
ipublished work. concentrations of 10.0 to 100.0 ppm, and
In the case of 9 ions, not even the the reaction was not uniform until the
higher concentrations tested produced concentration reached 150.0 ppm.
distress or retraction, as follows:
DISCUSSION AND CONCLUSIONS
still non -toxic The ions in Fig, 1 are arranged in

ions: at: the order of their increa sing toxicity. If
the concentrations beexpressed in mo-
tin larity, instead of parts per million, ap-
gold proximately the same series would result]
strontium as when the ions are ranked as in Fig. 1;
lithium
10.0 ppm
Cd, Ag, Cu, Zn, Ni, Pb, Ba, Al, Fe,4,
molybdate Mn, Co. It is notable that the three most
dichromate toxic ions, cadmium, silver, and copper,
produced distress at 0.05 to 0.1 ppm, or4|
fluoride 100.0 ppm to 6X 10"'^ M, under these experimental '
conditions.
tungstate From the data under consideration,
150.0 ppm
sulfite it appears that the ôwer the_concentration
parts
per
millón
300
150
100
80
40
30
20
10
5
1
0.100
0.050
0.005
DDW
jof an ion which produces retraction, the compounds also produce distress. Michel-
¡narrower the range of concentrations son (1957) has reported that minute quanti-
which produces distress, and the higher the •^ties or urethane or nicotine sulfate produce
jminimal concentration which produces re- similar reactions in this snail. Chemin
traction, the broader the range of concen- (1959) has reported several antibiotics \
trations which produces distress. Whether which seem to produce similar behavior,
this is generally true, or only coincidence and he noted that the toxic effect of some
in the series of ions tested, remains tobe of these was reduced by the presence of
demonstrated. Seven of the 13 ions of Fig. commercial potting soil (surely high in
1 seem tosubstantiate this hypothesis. organic compounds) or calcium, but that
Unfortunately, circumstances did not allow magnesium seemed to have no detoxifying
us to ascertain the minimal concentration effect.
\of barium, Chromate, manganese or cobalt The distress syndrome grossly re4
ions which will presumably produce resembles the response evoked by menthol,!
tractions. which is frequently used by malacologistsj
Lead may be an exception to the rule for "relaxing" fresh water snails before
that a range of concentrations which will fixing them for anatomical studies. Van
/produce distress occurs just below the der Schalle (1953) has reported the effect
•4 minimal concentration which will produce of nembutal in producing similar relax-
I '
! retraction. Perhaps the presence of or- ation.

ganic materials such as mucus or feces The actual physiological mechanism
was instrumental here in inhibiting the involved in the syndrome of distress is
3< toxic action of this ion. ûnknown, though it is apparently a neuro-
That extrinsic factors may limit the muscular phenomenon. Ciliary activity,
toxic action of copper is well known, and notoriously autonomous and difficult to
usually attributed to organicmaterial in the narrest by poison, seems unaffected by the
environment (Magalhaes Neto et al., 1953; ions we studied. Distress is a response,
Lacrange, 1952). Our experiments with which should receive much moreattentionl
zinc have shown that organic material may than it has in the past, not only for the'
not be the only factor which modifies the light it may shed on the general subject of
inability of the metal to produce distress. 4, •the physiology of fresh water organisms,
A layer of zinc powder covered the bottom but specifically for understanding thej
of the bowls that contained either 200 ml actions of moUuscicidesf
of DDW or of SRW-. In the latter the snails
showed normal activity, made trails LITERATURE CITED
through the powder, and zinc granules
could easily be demonstrated in the di- CHERNIN, E,, 1959, Notes on the effects
gestive tract and even in the liver lumen. of various antibiotics on Australorbis
In DDW, however, the snails soon showed glabratus. J. Parasit. 45: 268.
distress which persisted for the 24-hour DESCHIENS, R.1954, Incidence de la
period of the test. Zinc granules were not mineralization de l'eau sur les mol-
found in their digestive tracts. Presuma- lusques vecteurs des bilharzioses.
jbly the amount of organic material in each Conséquences pratiques. Bull. Soc.
/test lot was the same, and derived chiefly Path. Exot. 47: 915-929.
y from the mucus and waster products of the FREEMAN, L., 1953, A standardized
/ snails. method for determiningtoxicity of pure
Symptoms of the distress syndrome compounds to fish. Sewage and In-
have been noted only occasionally in the dustrial Wastes 25: 845-848.
Jitliterature (e.g. Nolan et al. 1953:719), HARRY, H. W., B. G. CUMBIE and J.
but have not previously been recognized MARTINEZ DE JESUS, 1957, Studies
>tas symptomatic with regards to toxicity. on the quality of fresh waters of Puerto
Certain concentrations of some organic Rico relative to the occurrence of
Australorbis glabratus. Amer. J. 1

Med. Hyg. 6: 313-322.
. influence
copper under
the molluscacide activity of
laboratory conditions.
HARRY, H. W. and D. V. ALDRICH, 1958, Pub. Avulsas Inst. Aggeu Magalhaes
Ihe ecology of Australorbis glabratus (Recife, Brazil), 2: 103-113.
in Puerto Rico. Bull. Wld. Hlth. Org. MICHELSON, E. ., 1957, Studies on the
18: 819-832. biological control of schistosome-
HUFF, L. , 1948, sensitive field test bearing snails. Parasitology (British).
for heavy metals in water. Econ. Ge- 47: 413-426.
ology, 43: 675-684. NOLAN, M. O., H. W. BOND and E. R.
LAGRANGE, E., 1952, A
propos de V MANN, 1953, Results of laboratory
action oligodynamique sur les mollus- screening tests of chemical compounds
ques pulmones en particulier et sur la for moUuscicidal activity. Amer. J.
faune aquatique en général. Arch. Inter- Trop. Med. Hyg. 2: 716-752.
nat. Pharmacodyn. Thér. 91: 185-193. VAN DER SCHALIE, H., 1953, Nembutal
MAGALHAES NETO, . M., A. M. DE as a relaxing agent for moUusks,
ALMEIDA, J. DE MORAES and O. Amer. Midland Naturalist, 50:511-512.
B. CALADO, 1953, Factors that
ZUSAMMENFASSUNG
NOT-SYNDROM IN TAPHIUS GLABRATUS (BASOMMATOPHORA: PLANORBmAE)
INFOLGE TOXISCHER KONZENTRATIONEN ANORGANISCHER IONEN,
In sehr niedrigen Konzentrationen wässriger Lösungen toxischer Substanzen verhält

sich diese Schnecke normal, d.h. sie streckt ihren Körper aus der Schale hervor,
kriecht umher, frisst und erneuert an der Wasseroberfläche den Luftgehalt ihrer Lunge.
In höheren Konzentrationen bleibt sie völlig in ihrer Schale zurückgezogen. Zwischen
den Grenzwerten derjenigen Konzentrationen die Retraktion verursachen und solchen
die ein normales Verhalten zulassen, liegen diejenigen welche einen Zustand hervorrufen
den wir als Not-Syndrom bezeichnen wollen. Eine Schnecke in Nöten streckt zwar
ihren Fuss aus der Schale hervor, ist jedoch unfähig damit zu fassen und, daher, weder
imstande sich fortzubewegen,noch sich zu ernähren oder atmosphärische Luft einzuatmen.
Es tritt weiters eine Degeneration der Tentakeln ein und die im Magen befindlichen
Sandkörner werden ausgestossen; die Zilienbewegungen sowie der Herzschlag scheinen
jedoch unbeeinflusst. Schnecken, die syndromauslösenden Konzentrationen für weniger
als 24 Stunden ausgesetzt waren, erholen sich meist.
Auf ihre Wirksamkeit hin wurden 22 Ionen untersucht, von denen 13 den Notzustand
hervorriefen. Bei Ag, Cd und Cu Ionen war dies schon bei Konzentrationen zwischen
0.050 und 0.100 1 eilen pro Million der Fall; bei Mn und Co hingegen erst zwischen 20
und 150 T./Mill. bzw. 30 und 300 l./Mill. Bei Konzentrationen die zwischen diesen
Extremen liegen verursachten folgende 8 Ionen das Syndrom: Zn,Âl, Ni, Ba, 4, Pb,
Fe2, Fe3. Die 9 restlichen Ionen waren in den untersuchten Konzentrationen nicht
wirksam, und zwar: Sn, Au, Sr, Li, M0O4, 27, bis 10 T./Mill.; F bis 100 T./Mill.
sowie Wo04und SO3 bis 150 T./Mill. Die Breite des Wirkungsbereiches jener Konzen-
trationen auf welch, bei einem gegebenen Ion, ein Notzustand erfolgt ist seiner zu
andauernder Retraktion führenden Mindestkonzentration direkt proportional. Das Not-
Syndrom ist möglicherweise identisch mit der von Menthol und anderen organischen
Verbindungen ausgelösten Entspannung.
RÉSUMÉ
LE SYNDROME DE DETRESSE CHEZ TAPHIUS GLABRATUS (BASOMMATOPHORA:
PLANORBIDAE) CAUSE PAR DES CONCENTRATIONS TOXIQUES
D'IONS INORGANIQUES.
Exposé à de très faibles concentrations de substances toxiques en solution aqueuse,

ce mollusque se comporte normalement, c'est à dire qu'il étend son corps hors de sa
coquille, rampe, se nourrit et renouvelle à la surface de l'eau l'air de sa cavité pulmo-
naire. En concentrations plus élevées il reste rétracté dans sa coquille. Entre les
limites des concentrations qui provoquent la rétraction et celles qui permettent encore
un comportement normal, se trouvent celles qui amènent un etat que nous appelons
détresse. Le pied d'un mollusque en détresse est étendu hors de sa coquille, mais il
est incapable d'adhérer, de sorte que l'animal ne peut se nourrir ni respirer l'air atmos-
phérique. Ses tentacules se détériorent, les grains de sable contenus dans son estomac
sont rejetés, mais ni l'action ciliaire ni les battements du coeur ne semblent affectés. Le
mollusque, d'habitude, se remet, s'il a été soumis moins de 24 heures à ces conditions.
Vingt-deux ions furent essayés, dont 13 déclenchèrent le syndrome. Les ions d'Ag,
Cd et Cu provoquèrent la détresse déjà entre les concentrations de 0.050 et 0.100
parties par million, tandis que ceux de Mn et Co ne la firent apparaître qu'entre 20 et
150 ppm et 30 et 300 ppm respectivement. Huit autres ions furent efficaces à des
concentrations intermédiaires entre ces extrêmes: Zn, Al, Ni, Ba, 4, Pb, Fe2, Feß,
Les 9 ions restants ne donnèrent aucune réaction dans les échelles de concentration
essayées, soit: Sn, Au, Sr, Li, M0O4, 27, jusqu'à 10 ppm; F jusqu'à 100 ppm ainsi
que W0O4 et SO3 jusqu'à 150 ppm. L'étendue de l'intervalle de concentrations provo-
quant le syndrome pour un ion donné est directement proportionelle à sa concentration
minimale donnant lieu à une rétraction durable. Le syndrome de détresse pourrait
bien être identique au relâchement causé par le menthol et par d'autres composés
organiques.
RESUMEN
SÍNDROME DEPRESIVO EN TAPHIUS GLABRATUS (SAY) (BASOMMATOPHORA:

PLANORBIDAE) COMO REACCIÓN A CONCENTRACIONES TOXICAS
DE IONES ORGÁNICOS
En concentraciones mínimas de materiales tóxicos este caracol muestra com-

portamiento normal, extendiendo su cuerpo fuera de la concha, moviéndose, alimen-
tándose y renovando su burbuja pulmonar en la superficie del agua. En mayores
concentraciones tóxicas, el caracol permanece retraído dentro de la concha. Entre
las condiciones que producen retracción y las que permiten comportamiento normal,
existe una serie de concentraciones que inducen una condición depresiva: el animal se
extende pero no es capaz de afirmar el pie y, por consiquiente, de reptar, alimentarse
o respirar aire atmosférico. Sobreviene deterioración de los tentáculos y eliminación
de granos de arena del estómago, pero la actividad ciliar y la pulsación cardíaca no
parecen afectadas. Los caracoles sometidos a esta condición por menos de 24 horas
generalmente se recuperan.
Se probaron 22 iones, de los cuales 13 produjeron estados de depresión. Los iones
Ag, Cd y Cu produjeron tal efecto entre 0.050 y 0.100 ppm; Mn y Co solamente entre
20 y 150 ppm, y 30 y 300 ppm, respectivamente. Ocho otros iones produjeron depresión
en concentraciones distribuidas entre esos extremos (Zn, Al, Ni, Ba, CrÜ4, Pb, Fe^^
). La gradación no tóxica de las concentraciones de los 9 iones restantes fué la
siguiente: Sn, Au, Sr, Li, M0O4 y 27, hasta 10 ppm; F, hasta 100 ppm; W0O4 y SO3,
hasta 150 ppm. La amplitud del rango de concentraciones que producen depresión
es directamente proporcional a la concentración mínima de un ion que produjera
retracción continua. El síndrome depresivo sería quizás idéntico al efecto relajante
producido por mentol y otros compuestos orgánicos.
) /75 GLABHATÎ/S ,(BASOMM ATO PHORA PLANOR-
,
(SAY)
,
,, ,,
, ,
. .
,,
,
. ,.
, -;, ..
-
-
-
.
,
, 24
. .
13
Ni,
.,
,
,
Ag, Cd,
mWo04,
,
9 :
Cu
Cr04, Pb, Fe2,
SO3
20
Sn,
Feß).
Au,
150
150
.
Sr,
.
Li,
, ,-
--
M0O4, 27,
0.050
10 ppm; F
0.100
(Zn,
100
-
Al,
-
I
THE EFFECTS OF ECHINOPARYPHIUM bARVAE ON THE STRUCTURE
OF AND GLYCOGEN DEPOSITION IN THE HEPATOPANCREAS OF
HELISOMA TRIVOLVIS AND GLYCOGENESIS IN THE PARASITE LARVAE 1
Thomas C. Cheng
Department of Biology, Lafayette College
Easton, Pennsylvania, U. S. A.
ABSTRACT
Histological examinations of the hepatopancreatic tissues of the planorbid snail

Helisoma trivolvisinvadedby the rediaeof the trematode Echinoparyphium sp. revealed
that the damage to host cells can be attributed primarily to ingestion and secondarily
to histolysis by redial digestive enzymes which are egested along with cellular debris.
Morphological evidence of the secondary lytic activity is furnished by lysed cells in
areas where there are heavy concentrations of cellular debris identical to that found in
the redial intestinal cecum. The presence of rediae in the hepatopancreas induces the
hypersecretion of yellowish globules by hepatic cells. Histochemical studies revealed
that less glycogen is removed from intact hepatopancreatic cells of H. trivolvis infected
with the rediae of Echinoparyphium sp. than in instances where the sporocysts of
Glypthelmins pennsylvaniensis are present. These rediae are believed to acquire
their carbohydrate requirements primarily through the direct ingestion of the host's
glycogen-containing hepatopancreatic cells rather than through the absorption of mono-
saccharides which have resulted from the breakdown of glycogen in intact cells, as do
the sporocysts.
The redial body wall and oral sucker are the primary sites of glycogen storage in
this larval stage. However, not all the PAS-positive material in the redial wall is
glycogen, since it is only partially digested with diastase. The sites of glycogen
deposition in developing cercariae include the sucker primordia, the primordia of the
pharynx, esophagus, intestinal ceca, and the two conspicuous lateral collecting tubules
of the excretory system. In addition, parenchymal cells also include glycogen, except
for a few large cells, probably developing cystogenous glands, which include PAS-
positive material that is only partially digested when treated with diastase. Compara-
tively little glycogen is present in cercarial tails where the heaviest concentration is
found along the "core" of each tail. This limited occurrence of glycogen might be
explained by the lack of a prolonged physiological need for carbohydrates in this tempo-
rary organ. It is believed that carbohydrate metabolism of stored glycogen serves as
the primary means for energy production in these intramoUuscan larvae since, although
fatty acids are present, the essentially anaerobic environment in the hepatopancreas
does not lend itself to energy production through lipid metabolism.
INTRODUCTION of the planorbid snail Helisoma trivolvis

(Say) infected with the rediaeof an echino-
The various types of known structural stome of the genus EchinoparyphiumDietz.
and physiological changes effected by Aspects of this study have been reported
trematode larvae on their moUuscan hosts in a preliminary abstract (Cheng, 1962).
have been reviewed in earlier papers The effects of trematode rediae on their
(Cheng and Snyder, 1962 a, b). This study molluscan hosts have been studied by
is concerned with the structural and Hurst (1927), F. G. Rees (1934), W. J.
chemical alterations in the hepatopancreas Rees (1936), Cheng and James (1960),
iThis research was supported by Grants E-3443, E-3443C1, and AI 3443-03 from the
Institute of Allergy and Infectious Diseases, National Institutes of Health, U. S. Public
Health Service. It is a contribution from the Laboratory of Parasitology and Inverte-
brate Zoology, Jenks Biological Laboratories, Lafayette College.
(291)
292 T. . CHENG
and pictured by Malek (1962). These collected from Green Pond, Bethlehem,
authors reported local histolysis, me- Northampton County, Pennsylvania. Twen-
chanical disruptions, decrease in hepato- ty per cent of these were found to be in-
pancreatic glycogen, and indirect damage fected with the rediae of an echinostome
to the digestive gland resulting from rediae trematode. The parasite could not be
located outside of that gland in the ad- identified definitely by its redial and
jacent gonads. This present study serves cercarial stages, although it was suspected
to confirm some of the earlier findings but that these represented stages in the life
more important, to present a new hypothe- cycle of a member of the genus Echino-
sis as to the mechanism involved in local paryphium. Therefore, experiments were
histolysis of hepatopancreatic cells. conducted to determine the life history of
Furthermore, presented below is a com- this trematode so as to obtain adults
parison and a possible explanation for the which could be identified more readily.
differences which exist in the amount of While dissecting the snails' internal
the host's hepatopancreatic glycogen which organs to determine the primary site of
is removed when rediae or sporocystsare infection by rediae, a number of encysted
present. metacercariae were found loosely attached
to the inner surface of the shells, to the
MATERIALS AND METHODS alimentary tract, and to the tunica propria
of the hepatopancreas. Such metacer-
During the summer of 1962, over 500 cariae,removed from their cyst walls,
specimens of Helisoma trivolvis were showed a striking resemblance to the
EXPLANATION OF PLATES
AL, acinar lumen of hepatopancreatic tubule; CF, cell fragments scattered in between
rediae; DCER, developing cercariae; FRIC, cell fragments in redial intestinal cecum;
HCG, globules secreted by hepatic cells; IHC, intact hepatopancreatic cells; LHC, dis-
lodged hepatopancreatic cells; PASCT, PAS+ material associated with lateral collecting
tubule of excretory system; PASG, PAS+ material in parenchymal gland cells; PASHC,
PAS+ material in hepatopancreatic cell; PASI, PAS+ material in intestinal cecal wall of
redia; PASIC, PAS+ material intermingledwithcellular debris in lumen of redial cecum;
PASP, PAS+ material associated with primordium of sucker; PASRW, PAS+ material
associated with redial wall; RED, redia; RW, redial wall;
young developing cercarla.
,
cercarial tail; YDCER,
PLATE I
FIG. 1. Photomicrograph of cross-section of redia of Echinoparyphium sp. in hepato-

pancreas ofHelisoma trivolvis showing cellular debris in intestinal cecum. Delafield's
hematoxylin. (40x obj.)
FIG. Photomicrograph of cross-section through hepatopancreas of infected

2. . tri-
volvis showing numerous rediae enclosing developing cercariae and a clump of cell
fragments. Mallory's triple. (40x obj.)
FIG. 3. Photomicrograph ofcross-section through hepatopancreas of infected H. tri-

volvis showing large number and dislodged
of light (yellowish) hepatic cell secretions
hepatopancreatic cells. Mallory's triple. (40x obj.)
FIG. 4. Photomicrograph of cross-sections of portions of several hepatopancreatic

tubules of uninfected .
trivolvis showing rich distribution of PAS+ material in cells.
PAS method. (40x obj.)
EFFECTS OF ECHINOPARYPHIUM ON HELISOMA 293
PLATE I
294 T. . CHENG
emitted cercariae in the number and ar- 10 infected snails were fixed in Zenker's
rangement of spines forming the collar. Fixative, embedded, and sectioned in the
In order to confirm that the metacer- same manner. Alternate slides of this
cariae found were of the same species as series were exposed to the periodic acid-
the cercariae, uninfected laboratory Schiff (PAS) reaction for glycogen, ac-
raised H. trivolvis and uninfected Physa cording to the procedure given earlier
gyrina (Say),collectedfrom another locale, (Cheng and Snyder, 1962a); the remaining
were placed in finger bowls together with were exposed to 0.5% diastase and the
individual infected H. trivolvis. Both PAS reaction to serve as controls. Three
species of snails thus exposed harbored of the uninfected snails were fixed in
encysted metacercariae within 24 hours, Zenker's, sectioned, and treated with the
which proved that the metacercariae had PAS reaction, with alternating slides
developed from the emitted cercariae. treated with diastase to serve as controls.
Encysted metacercariae were fed to The remaining 2 uninfected snails were
laboratory raised Japanese quails, Co- similarly fixed, sectioned, and stained
tumix coturmx japónica, and from their with one or the other of the two histo-
small intestine sexually mature adult logical stains given.
worms were recovered on the 8th day.
Examination of stained and mounted adult RESULTS
specimens verified that the trematode was
a member of the genus Echinoparyphium, Histological: The hepatopancreas of
although it could not be readily relegated Helisoma trivolvis is the primary site
to any of the known species. of infection by the rediae of Echino-
After successful identification of the paryphium sp. There is extensive damage
parasite, 20 infected and 5 uninfected of the hepatopancreatic tubules. Critical
snails were removed from their shells examinations revealed that most of the
and prepared for histological and histo- intact cells are severed and removed,
chemical studies. Ten of the infected since large areas of the hepatopancreas
snails were fixed in Carnoy's Fixative at all levels are devoid of visible cells;
(6:1:1), embedded, and sectioned at 8 resulting pockets are packed with rediae
microns, some in cross-section and others which enclose developing cercariae. Frag-
in longitudinal-section. Alternate slides ments of the hosts' hepatopancreatic cells
of thesewere stainedwithMallory's triple are present in the intestinal ceca of the
connective tissue stain and Delafield's rediae (Plate 1, Fig. 1). In addition, clumps
hematoxylin respectively. The remaining of similar cell fragments are found inter-
PLATE
FIG. 1. Photomicrograph of cross-section of a single hepatopancreatic tubule of
infected H. trivolvis showing less PAS+ material in cytoplasm than that observed in
Plate 1, Fig. 4. PAS method. (40x obj.)
FIG. 2. Photomicrograph of cross-section through a redia of Echinoparyphium sp.

showing PAS+ material associated with redial wall, developing cercariae, cecal wall,
and cecal contents. PAS method. (40x obj.)
FIG. 3. Photomicrograph of section through a redia showing PAS+ material associated
with area of presumptive sucker and the absence of PAS+ material in young differenti-
ating germ balls. PAS method. (40x obj.)
FIG. 4. Photomicrograph of cross-section through a redia showing PAS+ material

associated with lateral collecting tubules of excretory system of cercarla and lack of
PAS+ material in young developing cercarla. PAS method. (40x obj.)
PLATE II
PASP
PASCT %
YDCER
i
296 T. . CHENG
mingled with intact host cells outside of
the rediae in the hepatopancreas (Plate 1,
cells of uninfected /5
trivolvis are
characteristically rich in glycogen as
Fig. 2). These aggregates of cell frag- determined by the PAS reaction. The
ments are believed to come from the stored glycogen appears as finely granular
redial intestinal ceca because a con- PAS-positive material more or less evenly
siderable amount of dark, almost black, distributed in the cytoplasm, usually with
granules are intermingled among the a slightly greater concentration towards
cellular debris. These granules are also the luminal portion of each cell forming
present in the ceca and appear charac- the acinus (Plate I, Fig. 4).
teristically to be associated with ingested In . trivolvis infected with Echino-
cells which have been subjected to paryphium rediae, there is a slight de-
digestion. The blackish coloration of these crease in the amount of stored glycogen
granules is not due to differential staining, in the hepatopancreatic cells (Plate II,
since their appearance is identical in Fig. 1). The decrease is definitely not as
sections stained both by Mallory's and drastic as is the case in H. trivolvis
Delafield hematoxylin. Such aggregates, infected with sporocysts of Glypthelmins
when observed outside of the rediae, are pennsylvaniensis Cheng (see Cheng and
most plentiful in areas where large Snyder, 1962a).
numbers of rediae are found. Intact cells The redial wall is strongly PAS-
in the presence of cell fragments positive, as are the oral sucker, the
commonly show symptoms of lysis. intestinal cecal wall, and the cecal contents
A large number of irregularly rounded (Plate II, Fig. 2). The PAS-positive
yellowish globules are present in the prox- materials found within the redial wall,
imity of the few remaining intact cells however, are not all glycogen; in diastase-
(Plate 1, Fig. 3). Such globules, in lesser treated control sections, although there is
quantity, normally are found intracyto- a noticeable decrease in the amount of
plasmically in certain hepatopancreatic PAS-positive material in the redial wall,
cells of uninfected snails and undoubtedly some of it persists.
represent the liver cell globules of Bar- There is no PAS-positive material in
furth (1883). The yellowish coloration the bodies of germ balls and very little
of these globules is apparent in sections in young differentiating cercariae (Plate
stained both by Mallory's and Delafield II, Figs. 3, 4). Large deposits of PAS-
hematoxylin. positive material appear at the time the
Histochemical: The hepatopancreatic suckers begin differentiating. Atthattime,
PLATE
FIG. 1. Photomicrograph of section through redia showing PAS+ material associated
with area of presumptive sucker and with lateral collecting tubules of cercarial
excretory system. PAS method. (40x obj.)
FIG. 2. Photomicrograph of section through fully developed cercarla in brood chamber

of redia showing PAS+ material associated with parenchymal gland cells. PAS method.
(40x obj.)
FIG. 3. Photomicrograph of section through fully developed cercarla showing PAS+

material associated with dorsally located parenchymal gland cells. PAS method. (40x
obj.)
FIG. 4. Photomicrographof longitudinal section of fully developed cercarla showing

PAS+ material associated with parenchymal gland cells and slight amount of PAS+
material associated with cercarial tail. PAS method. (40x obj.)
PLATE III
RW
/
PAS6
3
298 T. . CHENG
the sucker primordia are extremely rich or S. palustris. Various authors (Tsuch-
in PAS-positive material (Plate II, Fig. 3; imochi, 1924; Mathias, 1926, 1927; Harper,
Plate III, Fig. 1). In addition, the primordia 1929; Suzuki, 1932) have demonstrated
of the pharynx, esophagus, intestinal ceca, that E. recurvatum (Linstow) can utilize
and the two conspicuous lateral collecting a gastropod as the second intermediate
tubules of the excretory system are also host. This pattern among Echinopary-
rich in PAS-positive material (Plate II, phium spp., which includes two molluscan
Fig 4; Plate III, Fig. 1). When observed intermediate hosts, is upheld by the
in cross-section, a few large parenchymal species under consideration.
cells lying along the dorsal body wall It is quite evident that the rediae of
include heavy concentrations of homo- Echinoparyphium do cause drastic damage
geneously PAS-positive material (Plate to the hepatopancreas of Helisoma tri-
III, Figs. 2, 3, 4). These undoubtedly volvis. The primary method of cell
represent gland cells, probably cysto- destruction and removal appears to be
genous glands. The remaining parenchy- through direct ingestion. This interpre-
mal cells also include finely granular tation is borne out by the presence of
PAS-positive materials, but these are not large quantities of cellular debris found
as heavily concentrated. in the intestinal ceca of the rediae and
Comparatively small amounts of PAS- the severe mechanical damage visible in
positive material are present in the cer- those areas of the digestive gland where
carial tails where the heaviest concen- the heaviest concentrations of rediae are
tration is seen along the "core" of each found. Direct ingestion of the molluscan
tail (Plate III, Fig. 4). host's hepatopancreatic cells have been
Practically all of the PAS-positive reported by Cheng and James (1960).
material found in cercariae is glycogen, Unlike Hurst (1927), this author does not
since such material was not observed in consider this type of damage secondary
in diastase -treated sections, except in the to histolytic damage although histolysis
large gland cells situated in the parenchy- is evident.
ma. Although the intensity of PAS-positive Hurst (1927) and F. G. Rees (1934)
staining in these cells is reduced con- attributed histolysis in mollusks infected
siderably when treated with diastase, these with rediae to the parasites' excretory
cells were still definitely PAS-positive products. Although there is strong evi-
even after 45 minutes of digestion. dence for this in instances where the
trematode larvae are sporocysts (Cheng
DISCUSSION AND CONCLUSIONS and Snyder, 1962a), it does not appear to
be the case when rediae are present, for
The life history pattern among Echino- symptoms of histolysis were not observed
paryphium spp. is well known. Reich in any cells not in contact with, or in the
(1927) reported that the cercariae of immediate proximity of, rediae. This
Echinoparyphium aconiatum Dietz, es- author proposes that local histolysis is
caping from Stagnicola palustris (Müller) affected by some lytic substance, most
(=Lymnaea palustris), enter and encyst probably digestive enzymes, which are
in the same and other species of fresh- egested through the mouth rather than
water gastropods. McCoy (1927) and excreted by rediae. This hypothesis is
Najarían (1954) reported that the meta- based on two major observations. (1)
cercariae of E. flexum (Linton) were Lysis of hepatopancreatic cells of H.
found in Helisoma trivolvis and Lymnaea trivolvis seen in this present study is
lacustris (Leach) after the cercariae had limited to areas where aggregates of
escaped from Physa integra (Haldeman) cellular debris identical to that found in
the intestinal ceca of rediae are present. is the case when sporocysts are present.
There can be no other source for such It known at this time whether the
is not
aggregates except from the redial ceca, glycogen present in the redial oral sucker
hence these must represent egested non- and wall is derived from the ingested
digestible material. It is suggested that, glycogen or whether it is carried over
along with the egestion of debris, cecal from an earlier stage of development. It
enzymes are passed out and it is these is apparent, however, that the ingested
which effect local cytolysis. (2) Cheng and glycogen is the carbohydrate source from
Snyder (1962c) and Cheng (1963a) have which the glycogen found in developing
demonstrated that there is a high concen- cercariae is derived. It does not appear
tration of acid and alkaline phosphatase possible that the glycogen molecule,
activity in the intestinal ceca. oí Echino- because of its size, can permeate through
paryphium rediae. Furthermore, there the cecal wall; hence it is believed that
is a similar concentration of phosphatase the glycogen molecule is first hydrolyzed
activity associated with the presumably to monosaccharides which permeate
egested material andthe sites of cytolysis. through the cecal wall, as in the case of
This again strongly suggests that enzymes the sporocyst wall (Cheng and Snyder,
normally found in the redial ceca are 1963), and are later resynthesized as
regurgitated along with indigestible debris glycogen in the bodies of developing cer-
and that local histolysis is affected by cariae. Whether this mechanism actually
these enzymes. does apply to the redial cecal wall is
In the case under consideration, me- being investigated currently.
chanical damage to the hepatopancreatic In an earlier paper (Cheng, 1963b) it was
cells of H. trivolvis is much more severe reported that the cercarial tail of Gor-
than that resulting from lysis of cells. In godera amplicava Looss includes less
comparing the degree of mechanical glycogen than the body proper, when found
damage caused by sporocysts with that within its molluscan host. This obser-
caused by rediae, the latter is muchmore vation was interpreted to mean " that
severe. This can be attributed to the since the body proper is the only portion
ability of rediae to ingest host cells. of the cercarla which continues to develop
It is now confirmed that the presence in the second intermediate host as the
of larval trematodes, be these sporocysts metacercaria while the tail is lost, the
or rediae, causes hypersecretion on the physiological need for a stored carbo-
part of the hepatic cells of hepatopancreas. hydrate source in the tail is not present
The secreted material forms yellowish and hence very little glycogen storage is
globules. appreciated in this structure." Since
The degree of glycogen reduction in less glycogen is found also in the cer-
Helisoma trivolvis infected with rediae is carial tail of Echinoparyphium, the same
considerably less than it would be if interpretation applies.
sporocysts were present. This is attri- As indicated, the PAS-positive material
buted to the fact that rediae actively ingest present in the redial wall is not completely
glycogen - containing hepatopancreatic composed of glycogen, since the diastase
cells and most probably acquire their it. The remaining
will only partially digest
carbohydrate requirement in this manner substance hence must be one of the follow-
rather than by absorbing glucose which ing: a neutral mucopolysaccharide, a
has resulted from the breakdown of the muco- or glycoprotein, a glycolipid, an
host's glycogen (Cheng and Snyder, 1962a, unsaturated lipid, a phospholipid, or
1963). The relatively small amount of combinations of these (Pearse, 1961).
stored cytoplasmic glycogen that is deleted Similarly, the PAS-positive but diastase
from intact cells in the presence of rediae resistant material in the cercarial
is also due to its digestion to simpler parenchymal gland cells must be of this
sugars which diffuse out of the cells, as category.
300 T. . CHENG
The low oxygen tension present within Looss. Proc. Helminth. Soc. Wash.,
the mollusk's hepatopancreas, coupled 30: 101-107.
with the relatively large size of rediae CHENG, T. C, and H. A. JAMES, 1960,
creates an essentially anaerobic environ- The histopathology of Crepidostomum
ment. For this reason, the presence of sp. infection in the second intermediate
glycogen in rediae and in the cercariae host, Sphaerium striatinum. Proc.
they enclose is significant, since in all Helminth. Soc. Wash., 27: 67-68.
probability carbohydrate metabolism is CHENG, T. C, and R. W. SNYDER, Jr.,
the primary source of energy in these 1962a, Studies on host-parasite re-
endoparasitic larval stages. Although lationships between larval trematodes
fatty acids are present (Cheng andSnyder, and their hosts. I. A review. II. The
1962b), these are generally not satis- utilization of the host's glycogen by the
factory for anaerobic energy production, intramolluscan larvae of Glypthelmins
because their carbon atoms, with the pennsylvaniensis Cheng, and related
exception of carboxyl carbons, are largely phenomena. Trans. Amer. Microsc.
reduced (von Brand, 1952) and hence do Soc, 81: 209-228.
not lend themselves to the internal oxi- ,
1962b,Studies on host-para-
dation-reductions characteristic of an- site relationships between larval trema-
aerobic processes.
ACKNOWLEDGEMENTS
todes and their hosts. III.
aspects of lipid metabolism in /5

Certain
trivolvis (Say) infected with the larvae

of Glypthelmins pennsylvaniensis Cheng
The author is grateful to Mr. Randall and related phenomena. Trans. Amer.
W. Snyder, Jr., School of Medicine, Uni- Microsc. Soc, 81: 327-331.
versity of Virginia, for technical as- , 1962c, Phosphatase activity in
sistance, and to Dr. Emile A. Malek, hepatopancreatic cells oi Helisoma tri-
Medical School, Tulane University, for volvis infected with rediae of Echino-
verifying the taxonomy of the moUusks paryphium sp. (Trematoda: Echino-
mentioned in this paper. stomatidae). Amer. Zool., 2: 513.
, 1963, Studies on host-para-
REFERENCES site relationships between larval trema-
todes and their hosts. IV. A histo-
BARFURTH, D., 1883, Über den Bau chemical determination of glucose and
und die Thätigkeit der Gasteropoden- metabolism of molluscan
its role in the
leber. Arch. f. Mikr. Anat., 22: 473- host and parasite. Trans. Amer.
524. Microsc. Soc, (In press).
CHENG, T. C, 1962, The effects of HARPER, V/. F., 1929, On the structure
parasitism by the larvae of Echino- and life histories of British freshwater
paryphium Dietz (Trematoda: Echino- trematodes. Parasitol., 21: 189-219.
stomatidae) on the structure and HURST, C. T., 1927, Structural and
glycogen deposition in the hepatopancre- functional changes produced in the
as of Helisoma trivolvis (Say). Amer. gastropod mollusk, Physa occidentalis,
Zool., 2: 513. in the case of parasitism by the larvae
,
1963a, Studies on phosphatase of Echinostoma revolutum. Univ. Cal.
systems in hepatopancreatic cells of the Publ. Zool., 29: 321-404.
molluscan host of Echinoparyphium sp, MALEK, E. A., 1962, Laboratory Guide
and in the rediae and cercariae of this and Notes for Medical Malacology.
trematode. Parasitol., (In press). Burgess Publ. Co., Minneapolis, Minn.
1963b, Histological and histo-
,
154 p.
chemical studies on the effects of para- MATHIAS, P. 1926, Sur le cycle évolutif
sitism of Musculium partumeium (Say) d'un trematode de la famille des Echino-
by the larvae of Gorgodera ampiicava stomidae Dietz, Echinoparyphium re-
curvatum Linstow. C. R. Acad. Sei., 1936, The effect of parasitism

,
183: 90-92. by larval trematodes on the tissues of

y 1927, Cycle évolutif d'un trêma- Littorina littorea (Linné). Proc. Zool.
tode de la famille des Echinostomatidae Soc. London, p 357-368.
(Echinoparyphium recurvatum Lin- REICH, F., 1927, Beitrag zur Kenntnis
stow). Ann. Sei. Nat. Zool., ser. 10, der Echinostomiden. 1. Der Leben-
10: 289-310. szyklus von Echinoparyphium aconiatum
Me COY, . R., 1927, Life history Dtz. 2. Cercaría latícaudata . sp.
studies on trematodes from Missouri. Zbl. f. Bakt. I, 103: 279-290.
J. ParasitoL, 14: 127-128. SUZUKI, M., 1932, On several cer-
najarían, H. H., 1954, Developmental cariae infesting lymnaeas in the sur-
stages in the life cyele of Echino- roundings of Taichu. Taiwan Igakkwai
paryphium flexum (Linton, 1892) Dietz, Zasshi, Taihoku, 31: 151-154. (Japanese
1910. J. Morph., 94: 165-197. text).
PEARSE, A. G. E., 1961, Histochemistry : TSUCHIMOCHI, K., 1924, On the life
Theoretical and Applied. Little Brown cycle of two species of echinostomatid
and Co., Boston. 998 p. trematodes. I. Studies on trematodes
REES, F. G., 1934, Cercaría patellae of domestic fowls in Formosa. Dobut-
Lebour, 1911, and its effects on the sugaku Zasshi, 36: 245-248.
digestive gland and gonads of Patella VON BRAND, T., 1952, Chemical Physi-
vulgata. Proc. Zool. Soc. London, p 45- ology of Endoparasitic Animals Aca-.
53. demic Press, N. Y. 339^^
ZUSAMMENFASSUNG
DER EINFLUSS VON ECHINOPARYPHIUM LARVEN (ECHINOSTOMIDAE) AUF DIE

STRUKTUR DES HEPAIOPANKREAS VON HELISOMA TRIVOLVIS SOWIE AUF
DIE GLYKOGENSPEICHERUNG, UND DIE GLYKOGENESE
IN DEN PARASITENLARVEN
Histologische Untersuchungen an hepatopankreatischen Geweben der planorbiden

Schnecken Helisoma trivolvis, die von Redien des Trematoden Echinoparyphium sp.
befallen waren, zeigten, dass die Schädigung der Wirtszellen in erster Linie darauf
zurückzuführen ist, dass diese von den Redien verschlungen werden und nur in zweiter
Linie auf deren Histolyse durch Verdauungsenzyme, die von den Redien zusammen mit
Zelltrümmern ausgespieen werden. Dafür, dass eine solche sekundäre lytische Wirkung
auch besteht, zeugt folgende Beobachtung: es finden sich angegriffene Zellen um dicht
konzentrierten Zelldetritus herum, der auffallend mit dem Inhalt der Darmblindsäcke
übereinstimmt. Die Anwesenheit der Redien löst in den hepatischen Zellen eine Hyper-
sekretion in Form gelber Tröpfchen aus. Histochemische Studien zeigten, dass aus den
unbeschädigten Leberzellen der mit Echinoparyphium sp. infizierten H. trivolvis
weniger Glykogen verlorengeht als dies bei Infektionen mit Sporocysten von Glypt-
helmins pennsylvaniensis der Fall ist. Es wird angenommen, dass diese Redien ihren
Karbohydratbedarf eher durch direkte Ingestion der glykogenhältigen hepatopankrea-
tischen Wirtszellen decken, als durch die Absorption von Monosacchariden, die aus dem
Abbau des Glykogens in unzerstörten Zellen herrUliren, wie es bei den genannten
Sporocysten der Fall ist.
Das Glykogen wird hauptsächlich in der Körperwand und dem Mundsaugnapf der
Redien gespeichert. Jedoch besteht nicht das gesamte PAS-positive Material aus
Glykogen, da es nur teilweise durch Diastase verdaut wird. In den sich entwickelnden
Zerkarien speichert sich das Glykogen in den Primordien der Salznäpfe, des Pharynx,
des Ösophagus, der Darmblindsäcke und in den 2 auffälligen seitlichen Sammelkanälchen
des Ausscheidur^sapparates an. Ausserdem entnalten die Parenchymzellen, mit
Ausnahme von einigen grösseren Zellen, die wahrscheinlich in Entwicklung begriffene
cystogene Drüsen sind, ebenfalls PAS-positives Material, dass nur teilweise von
302 T. . CHENG
Diastase verarbeitet wird. In den Zerkarienschwänzen findet man nur wenig Glykogen,
das hauptsächlich der zentralen Achse entlang konzentriert ist; dass die vorhandene
Glykogenmenge nur gering ist, lässt sich vielleicht dadurch erklären, dass diesses
temporäre Organ keinen anhaltenden physiologischen Bedarf an Karbohydraten hat. Es
wird angenommen, dass die Hauptquelle für die Energieerzeugung dieser intramollusken
Larven im Karbohydratmetabolismus des gespeicherten Glykogens zu suchen ist, denn,
obwohl Fettsäuren vorhanden sind, eignen sich die im wesentlichen anaerobischen
Bedingungen innerhalb des Hepatopankreas nicht zur Energieproduktion durch Lipoid-
metabolismus.
RESUME
L'EFFET DES LARVES D'ECHINOPARYPHIUM (ECHINOSTOMATIDAE) SUR
LA STRUCTURE DE L'HEPATOPANCREAS DE HELISOMA TRIVOLVIS ET SUR
LE DEPOT DE GLYCOGENE DANS CET ORGANE; ET ETUDE DE LA GLYCOGENESE
DANS LES LARVES DU PARASITE
L'examen histologique de tissus hépatopancréatiques du mollusque planorbe Helisoma,

trivolvis envahi par les rédies de Echinoparyphium sp. montre que la destruction des
cellules de l'hôte est attribuable enpremier lieu à une ingestion directe par les rédies
et secondairement aune histolyse par des enzymes digestives redíales qui sont expulsées
avec des débris cellulaires. La preuve morphologique d'une telle activité lytique
secondaire consiste en ce que l'on trouve des cellules attaquées autour d'amas denses
de débris cellulaires pareils au contenu des coecums intestinaux rédiaires. La présence
de rédies dans l'hépatopancréas provoque une hypersécrétion de globules jaunes par
les cellules hépatiques. L'étude histologique a montré que la perte de glycogène des
cellules intactes hépatopancréatiques des .trivolvis infectés par les rédies de Echino-
paryphium sp. est moindre qu'elle ne l'est quand ces mollusques sont infectés par les
sporocystes de Glypthelmins pennsylvaniensis Nous croyons que ces rédies obtiennent
.
les hydrates de carbone qui leur sont nécessaires, principalement par ingestion directe
des cellules hépatopancréatiques contenant du glycogène, plutôt que par absorption de
monosaccharides résultant d'un fractionnement du glycogène dans les cellules intactes
de leurs hôtes, comme le font les sporocystes.
Dans les rédies, la paroi du corps et la ventouse buccale sont les lieux principaux
de dépôt de glycogène. Mais, dans la paroi du corps, tout le matériel positif-PAS
n'est pas fait uniquement de glycogène, (il n'est que partiellement digéré par l'enzyme).
Les lieux de dépôt de glycogène dans les cercaires en cours de développement com-
prennent les ébanches de la ventouse, du pharynx, de l'oesophage, des caecums in-
testinaux, et les 2 conduits latéraux du système excréteur. En plus, les cellules du
parenchyme aussi contiennent du glycogène, à l'exception de quelques grandes cellules
qui sont probablement des glandes cystogènes en voie de développement; celles-ci
contiennent du matériel positif-PAS qui n'est que partiellement digéré par la diastase.
Dans les queues des cercaires on trouve assez peu de glycogène, situé surtout axiale-
ment. Sans doute ce taux réduit s'explique-t-il par le manque de besoin physiologique
durable d'hydrates de carbone dans cet organe temporaire.
Nous croyons que le principel mode de production d'énergie dans ces larves intra-
mollusques est un métabolisme carbohydratique du glycogène déposé, car, bien que les
acides gras soient présents, l'ambiance essentiellement anaerobique de l'hépatopancréas
ne favorise pas la production d'énergie par métabolisme lipidique.
RESUMEN
EL EFECTO DE LAS LARVAS DE ECHINOPARYPHIUM SOBRE LA ESTRUCTURA
DEL HEPATO PANCREAS DE HELISOMA TRIVOLVIS,'LA DEPOSICIÓN DE
GLYCOGENO EN ESE ÓRGANO Y LA GLUCOGENESIS EN LAS LARVAS PARASITAS
Investigaciones histológicas sobre el tejido hepatopancreático del caracol planórbido
Helisoma trivolvis infestado por redias del trematode Echinoparyphium sp. revelaron
que el dáno causado en las células del huésped puede atribuirse primariamente a la
ingestion y secundariamente a histólisis por enzimas digestivas rediales descargadas
junto con detritos celulares. La lisis de células en áreas donde hay gran concentración
de detritos idénticos a aquellos encontrados en el ciego intestinal de las redias constituye
evidencia morfológica de la actividad lítica secundaria.
E FFECTS OF ECHINOPAR YPHIUM ON HELISOMA 303
La presencia de redias en el hepatopancreas induce la hipersecreción de glóbulos

amarillentos por las células hepáticas. Estudios histoquímicos revelan que la substra-
ción de glucógeno de las células hepatopancreáticas intactas de H. trivolvis es menor
en la infestación por redias de Echinoparyphium que en aquélla por esporocistos de
Glypthelmis pennsylvanicus. Créese que esas redias satisfacen sus exigencias de
hidratos de carbono primariamente por ingestión directa de células hepatopancreáticas
que contienen glucógeno, qntes que por la absorción de monosacáridos resultantes de
la desintegración del glucógeno en células intactas, como en el caso de los esporocistos.
La pared del cuerpo redial y la ventosa oral son los sitios primarios en que se
almacena el glucógeno en este estado larval. Sin embargo, no todo el material PAS-
positivo en la pared redial es glucógeno, desde que éste sólo en parte es digerido por
diastasa. Los sitios de deposición de glucógeno en cercarlas en desarrollo incluyen
los primordios de la ventosa, faringe, esófago, ciege intestinal, y el par de tubos
colectores laterales, muy conspicuos, del sistema excretor. Además, células paren-
quimales también contienen glucógeno, excepto unas pocas células grandes, probable-
mente glándulas cistogénicas en desarrollo, las cuales incluyen material PAS-positivo,
que es digerido sólo parcialmente al ser tratado con diastasa. Comparativamente hay
poco glucógeno en la cola cercarial, concentrado principalmente a lo largo del eje. Esta
escasez de glucógeno podria explicarse por no haber una necesidad fisiológica prolongada
de hidratos de carbono en este órgano temporario. Créese que el metabolismo de las
reservas de glucógeno sirve come fuente de energía en estas larvas, desde que, aunque
ellas contienen ácidos grasos, el ambiente esencialmente anaeróbico en el hepatopan-
creas no se presta a la producción de energía a través del metabolismo de los lípidos,
ECHINOPARYPHIUM KA
HELISOMA TRIVOLVIS -
, -, , . (
-, -
Helisoma trivolvis,
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3)
,
Echinoparyphium -
.
sp.,
,,,, .
, -
,
, . , , -
-
--
Echinoparyphium
Glypthelm,ins
sp.,
pennsylvaniensis.
. ,
,
,
.
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.
A FLATWORM PREDATOR OF THE GIANT AFRICAN SNAIL
ACHATINA FÚLICA IN HAWAÜ^
Albert R. Mead
University of Arizona
Tuscon, Arizona, U. S. A.
ABSTRACT
The endemic, terrestrial, triclad turbellarian flatworm, Geoplana septemlineata

Hyman, 1939, has been found to prey upon the introduced Giant African Snail Achatina
fúlica in Hawaii killing even the largest specimens. The slender worms, attaining a
length of 40 - 60 mm, often attack in groups; 51 worms, totaling a length of nearly 2,000
mm, were removed from a single 50 mm giant snail specimen. The intense suction of
the worm's proboscis removes flesh from the exposed parts of the snail; and the invasion
of the lung cavity subjects the vital palliai organs to attack. Probably their greatest
efficacy in biological control lies in the destruction of the newly hatched snails. These
worms have also been observed to kill the introduced predatory snails Euglandina rosea
and Gonaxis quadrilateralis and the slug Deroceras laeve. A related form is reported
to attack A. fúlica in Java. The geoplanid worms and their allies unquestionably form
an important factor in the ecology of terrestrial mollusks. In Hawaii, a chain-reaction
is in progress: The introduction of foreign snails has increased the population of Geopl-
ana; Geoplana is in consequence providing a greater threat to the indigenous snails; and
since Geoplana has been found to carry eosinophilic meningocephalitis to humans, its
greater numbers might moreover intensify a public health problem.
An unsuspected endemic predator of the ussac), Gonaxis quadrilateralis (Preston)

Giant African Snail, Achatina fúlica and G. kibweziensis (E. A. Smith). When
Bowdich, has been found in Hawaii. Oddly these predatory snails attack a medium or
enough, it is not another beetle or "cannibal large size Achatina, the latter will in-
snail", of which several have been intro- variably keep crawling and evading in an
duced recently in Hawaii (see Mead 1961: attempt to escape. The predator will
102-45), but a terrestrial flatworm a— remove a considerable amount of mucus
triclad turbellarian or so-called "planar- and epidermal and dermal tissue; and then
ian"— which Hyman (1939) has described with its appetite satiated, it allows its prey
as Geoplana septemlineata and which is to escape. Within a very few weeks, new
apparently limited to the Hawaiian Islands. tissue regenerates and the snail appears
Swezey (1907: 54) and Williams (1931:339) no worse for its experience. In effect, then
probably refer to this species. these predatory snails together are able
In contrast to all other known inverte- to destroy the eggs and the young African
brate predators of the snail pest, Ac/zaiina snails up to about four months of age (also
fúlica, the worm can kill even the largest the older "pygmy" forms that remain es-
snail specimens; e.g. one experimental sentially at the four month size); but, with
specimen 128 mm long and weighing 168 relatively few exceptions, they merely
gm died from the attack of these worms. "harvest" the tissue of the body wall of
All but the smaller Giant African Snails the older and larger specimens.
(less than ca, 50 mm long) usually survive At first glance, it would not seem possi-
the attacks of the purposely introduced ble for Geoplana to be an effective and even
predatory snails Euglandina rosea (Fêr- lethal predator of a host several hundred
^This work was supported by a research grant, E

(in part) - 1245(C4), from the National
Institutes of Health, U. S. Public Health Service.
(305)
306 A. R. MEAD
times itsmass. In fact, although these is unduly sensitive to physical contact with
worms had frequently been observed in the these worms, since it elaborates a con-
field previously, their close association siderable amount of heavy, greenish,
with Achatina fúlica was not considered frothy mucus. This discharge does not
anything more than incidental. Specimens discourage the worms. The proportion-
of A. fúlica dead and covered with the ately long, white proboscis of the worm
worms did not raise any more serious is extruded from near the mid-ventral
question than did the numerous maggots surface of the body; it appears capable of
and adult flies, of several species, busily of strong suction, for many deep holes
consuming the carcass. But when fifty (ca. 0.75 mm in diameter) and grooves
full-grown, caged, experimental speci- appear at the sites of attack. Under the
mens of the Giant African Snail died, one microscope the translucent proboscis, in
after another, over a period of a very few carpet sweeper fashion, is observed to be
weeks, the suspicion that the worms were sucking in everything in its path-- mucus,
the direct cause of death steadily grew. It fluids, debris, air bubles —as it moves
was then a simple experiment to place a and probes about. The sensitivity of the
couple of snails in a one-gallon terrarium snail grows more acute with more worms
with a number of these worms. The obser- moving into position; and the victim with-
vations conclusively confirmed the fact draws into its shell, dragging the worms
that the worms would not only consume a in with it and embracing them in the folds
crushed or dying snail, but that they would of the invaginated head and tentacles. When
attack and kill a healthy, vigorous snail in many worms are present these all move
a matter of a very few hours, at the most. onto the exposed parts of the snail (the
These shiny -black, flat, leech-like, noc- mantle and left side of the foot) until
turnal worms usually measure about 40 x nothing but worms can be seen. The
2 mm in the extended state, although speci- harassed snail opens the pneumostome in
mens up to 60 mm are not infrequently a desperate effort to get more air; and
encountered. In spite of their extremely some of the worms crawl into the lung
tenacious slime, they move with remark- cavity. Soon the irritation and congestion
able speed and agility over even a dry sub- in the lung, as evidenced by the copious
strate. Very frequently their anterior end amount of mucus produced in the lung and
is attenuated, elevated, and flailed about in the bubbling from the pneumostome,
an apparent effort to locate prey. Both in causes the pneumostome to remain open,
the field and in the laboratory they seem only to permit still more worms to enter
sensitive to the slime trail of a snail; and, until a veritable webbing of black worms
in the vicinity of these worms, a snail is can be seen within.
soon seen with a number of worms rawling One 50 mm Achatina fúlica specimen
almost frantically in its wake. The di- was found in the field with 51 worms on it,
rective motion of the worms permits them totaling a length of nearly 2,000 mm, or
soon to overtake the snail with its hesitant, somewhat over six feet of worms. It is
probing locomotion. One after another little wonder that the snails are quickly
crawls upon the hapless victim until it is killed under such an attack. Attempts
apparent that one of the most effective have been made to rescue snails by re-
factors in the attack is the "ganging up" of moving all the worms as quickly as pos-
the worms on a snail whose escape reaction sible. If caught in time, the snail will
succeeds only in carrying its attackers survive and eventually regenerate the lost
with it and picking up still more en route. tissue; however, if the snail was under
î'roehlich (1955) suggests that in the heavy attack, it will not survive. Such a
attack of geoplanidsan extro-gastrovascu- snail appears emaciated, drawn, and al-
lar digestive enzyme may be released at most "dry" on the surface of the body,
the feeding site. This suggestion seems undoubtedly from the great loss of mucus;
reasonable, for it is obvious that the prey it seems exhausted and appears to move
PREDATOR OF ACHATINA FÚLICA 307
voluntarily only with the greatest effort; slightest stimulus, contributes sub-
it retracts violently on the slightest stimu- stantially to the increase in these worms.
lation; and it finally remains lethargic Picking up the worm with forceps will
and partly extended from its shell until almost invariably cause the worm to break
overtaken by death. When only one or two in two or more parts; and careless handling
worms attack a snail, the damage is often may cause it to fragment into many small
limited to the removal of tissue from the pieces. Even with gentle handling, the
exposed mantle and the posterior margins worm may seem to remain intact, only to
of the foot, parts which are vulnerably ex- autotomize seconds or minutes later.
posed to the marauding worm when the Geoplana is most abundant in the more
snail is in resting position on the ground. moist sections of Oahu and Kauai islands,
In environments where the worms are most although it has been encountered in some
abundant, Achatina specimens are often areas that are comparatively dry the year
seen with either freshly removed tissue around. Achatina fúlica tends to become
or regenerating tissue in these regions of nearly ubiquitous in Hawaii, but it still
the body, thus probably offering a fair index remains conspicuously unsuccessful in its
of the incidence of attack by the worms. attempts to invade some of the more lush,
The strongest preference is shown for deep valleys and higher peaks, in which
the newly hatched achatinas; and Geoplana areas the worm abounds. It is significant
undoubtedly is having its greatest effect that nearly 50% of the giant snails found
in biological control by destroying the alive in the very wet upper Manoa Valley
juveniles. Time after time, it has been in Oahu have had one or more worms on
observed both in the field and in experi- them. The predators Gonaxis quadri-
mental cages that these worms will Uate ralis and the smaller G. kibweziensis
congregate in great numbers in the egg reflect adaptations to their native East
masses. In fact, they are found laced all Africa by settling in the drier areas; in
through the eggs and adjacent debris; and contrast, Euglandina rosea seeks the more
even the intact soil around the "nest" in- moist areas and henc e is brought in greater
variably contains a few more specimens. contact with Geoplana. Of the four snails,
In attacking the newly hatched snail, the Euglandina doubtless suffers the greatest
worm either embraces it in its folds so that loss from attacks by Geoplana. Gonaxis
the proboscis can enter the aperture of the tends to burrow into the ground and the
shell, or it crawls into the shell and out worms therefore probably encounter it
again, forming a U-shaped fold that carries more frequently than the normally drier
the proboscis deeply into the body whorl. environment would suggest. Other intro-
The small shell characteristically is left duced snail pests, Bradybaena similaris
intact and completely clean of any flesh. (Ferussac), Subulina octona (Bruguiere)
Just how important is Geoplana in the and Opeas sp., within the past few years
economy of snail populations? There is have virtually vanished in some areas of
no question that this worm is amazingly Oahu. The explanation for the disap-
hardy and persistent, in spite of its appearance probably rests in a combination
parently delicate nature. It has been found of disease, predatory snails and these
in very dry environments curled up in the predatory worms» At the Kalalau Lookout
deep folds of leaf debris. In a given area, in Kauai, this species of Geoplana was
even with diligent searching none may be found feeding in characteristic fashion on
found; but soon after a rain, the large, full- a live specimen of the introduced slug
grown worms may be found in quantity — Deroceras reticulatum (Müller).
suggesting that they had successfully Although Froehlich (1955) mentions the
weathered the dry period by secreting snail-eating habits of Brazilian geoplanids
themselves in the ground and in deep re- and refers to earlier, more brief, accounts
cesses. Asexual reproduction through in the literature, the terrestrial turbel-
autotomy, apparently triggered by the larian flatworms, in general, have been
308 A. R. MEAD
almost completely unsuspected as an im- cardium. Further, since the worm is

portant ecological factor in snail popula- cannibalistic, transmission from one
tions. The geoplanids and their allies are worm to another appears easy. On the
widespread and there is little doubt that in other hand, the attacks by Geoplana could
many places, their presence has a pro- conceivably provide sufficient stress to
nounced effect upon snail distribution and cause an enzootic disease in the snails
abundance. These worms must be taken to go from the chronic to the acute -lethal
into consideration in the analysis of land phase. In this connection it should be noted
snail ecology. But their presence does not that, after removing the worms, some
necessarily mean snail prédation, for snails would appear to be recovering from
Froehlich indicates that some species have the attack and then suddenly go into a
a greater affinity for isopods and other decline and die, despite isolation and ample
arthropods; and in Hawaii, the introduced food and moisture^
Bipalium kewense so far has not been It is provocative to contemplate the
implicated in attacks on the giant snail, al- ecological "chain reactions" that are
though probably the scarce, endemic Geo- taking place as a result of the changes
plana subpalida Hyman (1939) eventually that have taken place in the past few years.
will be. However, in a recent communi- Geoplana septemlineata in its unaltered
cation Dr. Ir. J. Ruinard, of the Institute endemic state is apparently not a common
for Agricultural Research in Manokwari, animal. It has been seen feeding on earth-
Dutch New Guinea, reports seeing large, worms and small insects. Snails originally
black, leech-like worms on the bodies of did not figure importantly in its diet as
A. fúlica in that area. Specimens of these approximately half of the Hawaiian en-
worms have been examined by the author, demic snails are tree dwellers; and the
and while they are being definitely identi- geophilic forms are characteristically
fied by the proper authorities, it is already sparse in their distribution. This ecolo-
quite apparent that these large worms (ca. gical picture was changed with the arrival
75 X 7 mm
in the contracted state) are of Achatina fúlica when an abundant supply
geoplanids that could indeed be formidable of acceptable food became available» With
predators of the giant snail. Dr. Ruinard the introduction of the predatory snails,
interestingly reports, "We have found still more food became available, par-
some of these black animals inside the ticularly with Euglandina quickly invading
shell of a living giant snail; however the the deeper valleys and higher areas not
body of the snail was damaged. Another yet reached, and perhaps never to be
time [it] was seen that such a black 'leech' reached, by the Giant African Snail. The
was crawling on the back of a healthy population of Geoplana has unquestionably
looking slug.... One or two hours later on increased considerably as a result. This
the back of the slug 3 or 4 big, light colored opinion is supported by experimentation on
blisters appeared, which turned to black. a small scale and by my observations in
The next day the slug was dead. We ob- the field. One cannot but wonder what the
served that the leech pierced into the slug increased worm population is doing to the
[with] a white cylindric organ." scarce ground-dwelling endemic snails.
But the effect of this worm upon the And, as an interesting sidelight, it should
snail populations may have its more subtle be noted that Geoplana, along with the in-
aspects. A disease of unknown etiology troduced snails and slugs, has been shown
occurs in the giant African snail (Mead to be the intermediate host of the nematode
1956). Geoplana looms as a possible worm Angiostrongylus cantonensis, which
incidental or secondary vector of this causes the frequently fatal eosinophilic
infection particularly in view of its affinity meningocephalitis in humans (Alicata
for crawling into the pneumostome where 1962; see Mackerras and Sandars 1954).
it can come in direct contact with such The greater numbers of Geoplana and the
vital organs as the lung, kidney and peri- proclivity in this species for resting on
PREDATOR OF ACHATINA FÚLICA 309
the leaves of lettuce, automatically in- Exper. Gen., Paris, Notes et Rev., 80:
creases the danger of humans accidentally 116-24„
up on a bit
/
ingesting infected
grown Geo/)
worms. In fact, a full-
was recently found curled
of lettuce in a tossed salad
MACKERRAS, M. J. and D. F. SANDARS,
1954,
and
Life-history of the rat lung-worm
migration through the brain of
its
served at one of the better Waikiki restau- its host. Nature, 173(4411): 956-57.
rants. MEAD, A. R., 1956, Disease in the giant
African snail Achatina fúlica Bowdich.
Science, 123(3208): 1130-31.
REFERENCES , 1961, Ihe Giant African Snail:
a problem in economic malacology.
ALICATA, J. E., 1962, Angiostrongylus Univ. Chicago Press, xvii + 257 p.
cantonensis (Nematoda; Metastrongyl- SWEZEY, O. H., 1907, The sugar cane
idae) as a causative agent of eosino- leaf-roller (Omoides accepta) with an
philic meningocephalities of man in Ha- account of allied species and natural
waii and Tahiti. Canad. J. Zool., 40(1): enemies. Hawaiian Sugar Planters'
5-8. Assoc, Exp. Sta. Bull. No, 5, 60 p.
FROEHLICH, 1955, On the biology
G., WILLIAMS, F. X., 1931, Handbook of the
of land planarians. Bol. Fac. Fil. Cien. insects and other invertebrates of Ha-
Letr., Univ. S. Paulo, Zool., 20:263-72. waiian sugar cane fields. Hawaiian
HYMAN, L. H., 1939, Land planarians Sugar Planters' Assoc, Exp. Sta., 400p.
from the Hawaiian Islands, Arch. Zool.
ZUSAMMENFASSUNG
EIN PLATTWURM ALS FEIND DER AFRIKANISCHEN RIESENSCHNECKE
ACHATINA FÚLICA AUF HAWAH
Es wurde beobachtet, dass der auf Hawaii einheimische triclade Landturbellarier

Geoplana septemlineata Hyman, 1939, die eingeschleppte afrikanische Riesenschnecke
Achatina fúlica anfällt und sogar die grössten Individuen dieser Art zu töten vermag.
Diese schlanken Würmer, die eine Länge von 40-60 mm erreichen, greifen oft gruppen-
weise an: 51 Wtlrmer, mit einer Gesamtlänge von fast 2000 mm, wurden von einer
einzigen 50 mm langen Riesenschnecke entfernt. Das Fleisch exponierter Körper-
stellen wird durch die intensive Saugkraft des WurmrUssels abgetragen; auch die
lebenswichtigen pallialen Organe sind durch die Invasion der Lungenhöhle dem Angriff
ausgesetzt. Wahrscheinlich aber üben diese Würmer ihren wichtigsten Einñuss in der
biologischen Kontrolle dieser Schnecken dadurch aus, dass sie die frischgeschlUpften
Jungschnecken vernichten. Ebenso töten sie die eingeschleppten Raubschnecken Eu-
glandina rosea und Gonaxis quadrilateralis sowie die Ackerschnecke Deroceras laeve.
Eine verwandte Form greift auf Java A. foilica an. Die geoplaniden Würmer und ver-
wandte Formen stellen zweifelsohne in der Ökologie der Landmollusken einen ganz
unerwartet wichtigen Faktor dar. Auf Hawaii ist eine Kettenreaktion im Gange: die
Einschleppung der ausländischen Schnecken hat die Geoplanidenbevölkerung vermehrt,
welche daher eine grössere Gefahr für die einheimischen Schnecken bildet. Da Geoplana
sich auch als ein T. räger der menschlichen eosinophilen Meningocephalitis erwiesen
hat, könnte weiters die Vermehrung dieser Gattung einen gewissen Einfluss auf eine
Frage der öffentlichen Gesundheit des Landes ausüben.
310 A. R. MEAD
RÉSUMÉ
UN VER PLA1' PREDATEUR DU PULMONE GEANT AFRICAIN

ACHATINA FÚLICA A HAWAH
Nous avons pu établir que le turbellarié terrestre triclade Geoplana septemlineata

Hyman, 1939, indigène à Hawaii, attaque le Pulmoné géant Achatina fúlica importé de
l'Afrique, tuant les spécimens même les plus grands. Ces vers sveltes, qui atteignent
une longueur de 40 à 60 mm, attaquent souvent en groupe; 51 vers, mesurant au total
2000 mm, ont été recueillis sur un seul spécimen d'AcMtina de 50 mm de long. La chair
des parties exposées de l'animal est ôtée par la succion intense du probóscide de ces
vers; les organes vitaux palléaux sont aussi exposés à l'attaque par une invasion de la
cavité pulmonaire. Mais c'est sans doute la destruction des Acliatines nouvellement
éclos es qui agit le plus efficacement dans le contrôle biologique de l'espèce. De même
ces vers tuent aussi les Pulmones prédateurs importés Euglandina rosea et Gonaxis
quadrilateralis ainsi que la limace introduite Deroceras laeve. Une forme alliée attaque
Achatina fúlica à Java. Les vers géoplanides et leurs alliés sont incontestablement
un facteur important et inattendu dans l'écologie des mollusques terrestres. A Hawaii
une réaction en chaîne est en cours: l'introduction des Pulmones étraiigers a augmenté
la population de Geoplana qui, en conséquence, devient une menace plus grave pour les
Pulmones indigènes. D'autre part, puisque Geoplana a été constatée être un vecteur
de la méningocéphalite eosinophile humaine, la multiplication de cette espèce contribue
à aggraver ce problème de l'hygiène publique.
RESUMEN
UNA PLANARIA PREDADORA DEL CARACOL GIGANTE DE AFRICA,

ACHATINA FÚLICA, EN HAWAU
El turbelario tricládido terrestre endémico, Geoplana septemlineata Hyman, 1939,

se indica como predador del caracol gigante africano Achatina fúlica, introducido en
Hawaii, siendo capaz de matar hasta los más grandes individuos. Estas delgadas plana-
rias, que alcanzan una longitud de 40 a 60 mm, frecuentemente atacan en grupos: 51 de
ellas, totalizando una longitud de cerca de 2.000 mm, fueron extraídas de un solo
caracol gigante de 50 mm. La intensa succión proboscidal del turbelario arranca
pedazos de las partes expuestas del caracol, y la invasión de la cavidad pulmonar abre
el ataque a los órganos paléales vitales. Probablemente su mayor eficacia en el control
biológico reside en la destrucción de caracoles recién salidos del huevo. Estas plana-
rias también se han observado matando los caracoles predadores Euglandina rosea y
Gonaxis quadrilateralis, introducidos en Hawaii, y la babosa Deroceras laeve. Se ha
observado que una forma emparentada ataca Aclmtina fúlica en Java. Los geoplánidos
y sus aliados constituyen incuestionablemente un factor de importancia insospechada
en la ecología de los moluscos terrestres. En Hawaii está progresando una reacción
en cadena: la introducción de caracoles exóticos ha incrementado la población de
Geoplana, la cual, en consecuencia, se convierte en una amenaza creciente para los
caracoles endémicos; y desde que se ha verificado que Geoplana es transmisora de la
meningoencefalitis eosinofílica al hombre, su incremento produciría además la intensi-
ficación de un problema de salud pública,
,
ACHATINA
FÚLICA
.
lineata
.;. ,,
Hyman, 1939
50
Achatina fúlica
51
,
;
40
2,000
-
Geoplana
,60 ,-
septem-
1
.
, ,
PREDATOR OF ACHATINA FÚLICA
.
31
, . Deroceras
Evglandina rosea
.
Gonaxis
.
quadrilateralis , laeve.
:
;,
;
. fúlica,
Geoplana
--
CYTOTAXONOMIC STUDIES OF FRESHWATER LIMPETS
II. THE NEW ZEALAND RIVER LIMPET, LATIA NERITOIDES^
J. B. Burch and C. M. Patterson2
ABSTRACT
Latia neritoides Gray is a freshwater limpet restricted to New Zealand. It is generally
considered to be one of the most primitive freshwater basommatophorans and somewhat
related to the South American Chilina, but is usually placed in its own family, the Lati-
idae.
The haploid chromosome number of Latia neritoides is 18, which, although charac-
teristic of the Basommatophora in general, is different from that found in all other
freshwater limpets (Ancylidae) yet investigated (x=15, n=17), except Acroloxus lacustris
(Acroloxidae). However, on cytological grounds no relationship between Acroloxus and
Latia could be demonstrated, regardless of suggestions to the contrary based on mor-
phological comparisons. In fact, it would appear that they are not closely related.
INTRODUCTION The apparent primitiveness of Latia

makes it an interesting animal to study.
Latia neritoides Gray is a freshwater This pertains particularly to its cytology,
limpet restricted to streams of New since it has been suggested (Burch 1961,
Zealand. It has been variously placed with 1962; Burch and Heard, 1962) that almost
the Ancylidae (e.g., see Tryon, 1884; Suter, all changes in chromosome numbers of
1913), in its own family, the Latiidae phylogenetic significance have occurred
(Hutton, 1882; Thiele, 1931; Zilch, 1959; through aneuploidy, with an increase in
Taylor and Sohl, 1962) or with the Chilin- number in the direction of morphological
idae (see Boettger, 1955). specialization. It was then especially per-
Pelseneer (1901) was the first to de- tinent to compare the chromosome number
scribe the anatomy of Latia in any detail. of Latia with what is already known about
He considered be very different from
it to chromosome numbers in the other Basom-
the ancylids Ancylus, Acroloxus and
(i.e., matophora. Also, since Hubendick (1962)
Gundlachia) and less specialized than they suggests a connection between Latia and
are because it has a large palliai cavity Acroloxus, from the structure of their
and a long visceral commissure separated radulae and reproductive tracts, it is de-
into distinct ganglionic centers. In ad- sirable to compare cytological details of
dition, he pointed out Latia' s apparent Latia with what has already been demon-
relation to the South American snail strated for Acroloxus.
Chilina {"Latia paraissant avoir son plus Grateful acknowledgement is made to
proche voisin dans Chilina"). Since the Dr. R. K. Dell of the Dominion Museum,
time of Pelseneer, Latia has been gener- Wellington, New Zealand for kindly sup-
ally considered to be a primitive Basom- plying us with the cytological material of
matophoran. Latia neritoides used in this study.
^This investigation was supported (in part) by a research grant, 5 Tl AI 41-04, from the
National Institute of Allergy and Infectious Diseases, U. S. Public Health Service.
^Museum and Department of Zoology, University of Michigan, Ann Arbor, Michigan,

U. S. A.
(313)
314 BURCH AND PATTERSON
MATERIALS AND METHODS bivalent is 4.0 miera; the longest di-
mension of the smallest is 1.6 miera. The
Specimens used in this study were ob- sizes of Metaphase I chromosomes range
tained by Dr. R. K. Dell from a stream from 1.1 miera to 1.5 miera.
flowing into Otaki River near Otaki Falls, The gross morphological characters of
Wellington, New Zealand. The material the sperm of Latia neritoides when stained
examined consisted of ovotestes from 6 with acetic -orcein do not differ signifi-
specimens killed, fixed and preserved in cantly from those previously observed for
Newcomer's (1953) fluid. The repro- other Basommatophora (with the exception
ductive gland tissues were stained by the of Acroloxus lacustris). A minor differ-
acetic -orcein squash technique (La Cour, ence mentioned in passing is that the sperm
1941). Observations were made with heads have finer, longer and somewhat
Nippon Kogaku microscopes using lOOX sharper points at their anterior ends.
(n.a. 1.25) oil immersion objectives and The sperm heads (Fig. 2) measure 0.9 x
10-30X oculars. Photographs (Figs. 1-4) 3.6 miera and are turnip shaped. They are
were taken using a 20X ocular, oil im- followed by long, thin tails which are non-
mersion objective, a Kodak Wratten 57A staining, or are only faintly stained, with
(green) filter, and Kodak High Contrast acetic -orcein.
Copy film.
DISCUSSION
OBSERVATIONS
Among the most recent detailed con-
Spermatogonial chromosomes of Latia siderations of the phylogenetie placement
neritoides are metacentric as is charac- of Latia are those of Hubendiek (1945) and
teristic of the chromosomes of other Boettger (1955). They both consider (as
Basommatophora. The primary con- did Pelseneer) that Latia and Chilina are
strictions of the chromosomes can readily closely related"^ ;in addition, they both
be observed at metaphase. In addition, think that these two genera are primitive.
the larger chromosomes are bent at the Boettger regards Chilina to be the most
positions of their respecitve primary con- primitive of the freshwater Basom-
strictions (centromeres) and appear as matophora because in its nervous system
V's and J's when seen at metaphase in ithas a crossed chain of visceral ganglia.
polar view or when observed in side view As a result of its flattened limpet-shape
during anaphase. The sizes of spermato- Latia has apparently lost this visceral
gonial metaphase chromosomes corre- chain.
spond to those reported from similar Latia has 18 pairs of chromosomes.
squash preparations for other Basom- This is the number characteristic of
matophora (Burch, 1960a). Exact counts of Basommatophora in general, with the
of the chromosomes of spermatogonial exception of the Aneylidae (Burch, 1960a;
cells were not made. Burch, Basch and Bush, 1960), in which
Eighteen bivalents were observed during the basic numbers are 15 and 17. The
late prophase (diakinesis) (Fig. 4) and chromosome number then gives no indi-
metaphase of the first meiotic divisions cation of its relationships except that it
of spermatogenesis. At diakinesis nine does not deviate from that of most other
bivalents are held together by one chiasma, basommatophorans. Additional details of
eight by two chiasmata, and one by at least cytology observed in this study also do
two but probably three chiasmata. The not distinguish Latia from other Basom-
longest dimension of the largest diakinesis matophora.
"^Boettger places the two genera in the same family, although he distinguishes Latia by a
separate subfamily.
.
CYTOTAXONOMY OF LATÍA 315
> »*
#
*'*
Fig. 1. Sperm head oi Acroloxus lacustris (indicated by arrow).

Fig. 2. Seventeen sperm heads of Latia neritoides
Fig. 3. Late Prophase I (diakinesis) chromosomes oí Acroloxus lacustris.

Fig. 4. Late Prophase I (diakinesis) chromosomes of 2 cells oi Latia neritoides.
Compare with Fig. 3 for size.
Fig. 5. Drawing of the chromosomes shown in Fig. 4.
All figures X2025.
Since the European freshwater limpet placed at the opposite end of the basom-
Acroloxus lacustris in contradistinction matophoran phylogenetic scale from Laita.
to the ancylid limpets also has 18 pairs But Bondesen (1950), Burch (1962) and
of chromosomes (Burch,1962) and since Hubendick (1962) have shown that Acro-
Hubendick (1962) has suggested that it has
an origin in common with Latia, it will be
of interest to compare these 2 snails.
Acroloxus was until rather recently con-
loxus is an aberrant form which should
be separated from the ancylids. Bondesen
(1950) contended that the origin of
loxus "must be. ..sought nearer to more
forms within the Basom-
-
sidered a member of the Ancylidae and primitive
TABLE I. Comparison of the Morphology of Latia and Acroloxus
Feature Latia neritoides
Shell dextral
thick
with laminate septum
Apex coiled
Animal dextral
Pseudobranch on right side and

bearing medianly
located anus.
Dorsal adductor
muscles and
adhesive epithelium
Eyes
-
CYTOTAXONOMY OF LATÍA 317
matophora''8. Hubendick(1962) advocates lacustris. An attempt has been made, in

that Acroloxus had an origin in common Table I, to compare these 2 limpets by
with Latia. reviewing morphological information a-
Certain cytological aspects in Acroloxus vailable, mainly from Pelseneer (1901) and
lacustris have been given in a previous Hubendick (1962). The data shown in Table
study (Burch, 1962). Although it was found I indicate that any relationship advocated
that this species has the same chromosome for Latia and Acroloxus is still hypo-
number as that regarded as basic for thetical. In addition, until it is definitely
freshwater "pulmonates", it differs on shown justwhat the links between Acro-
other points from previously studied loxus and other Basommatophora are then
Basommatophora, The differences ob- perhaps Acroloxus should not be tied to
served consisted in the large size of the any special group.^ It will be of interest
various cells of spermatogenesis, the to see the anatomical details of Latia
greater volume ratio of chromatin to cyto- described in as thorough a manner as has
plasm, the relatively large size of the been done for various other freshwater
chromosomes and the morphology of the "pulmonates".
mature sperm, whose heads are long and
"thread -like", not bullet- or turnip LITERATURE CITED
shaped.
The comparative sizes of the chromo- BOETTGER, R., 1955, Die Systematik
somes of Acroloxus and Latia can be der euthyneuren Schnecken. Verhandl.
seen in Figs. 3 and 4 (all three cells are Deutsch. Zool. Ges. Tübingen, 1954:
at diakinesis and aliare magnified 2025X). 253-280.
Although the chromosomes of Latia (Fig. BONDESEN, P., 1950, A comparative
4) are in the size range normal for Basom- morphological-biological analysis of the
matophora, those of Acroloxus (Fig. 3) egg capsules of freshwater pulmonate
are noticeably very much larger. The gastropods. Natura Jutlandica,3 1 -208, :
relatively small amount of cytoplasm in pis. 1-9.

the cell of Acroloxus is indicated in Fig. BURCH, J. ., 1960a, Chromosome mor-
3 by the more compact mass of the chromo- phology of aquatic pulmonate snails
somes, i.e., the chromosomes are not (Mollusca: Pulmonata). Trans. Amer.
spread out as much as those of comparable Microsc. Soc, 79(4): 451-461.
cells of Latia. ,
1960b, Chromosome studies of
Even more striking differences are ob- aquatic pulmonate snails. Nucleus, 3(2):
served in the gross shapes of the sperm 177-208.
heads when viewed in chromosome squash , 1961, The chromosomes of
preparations. In Acroloxus the sperm Planorbarius comeus (Linnaeus), with
heads are relatively thin and very e- a discussion onthe value of chromosome
longated (Fig. 1). The peculiar shape of numbers in snail systematic s. Basteria,
Acroloxus sperm was first pointed out by 25(4/5): 45-52.
Retzius (1906). But in Latia the sperm 1962, Cytotaxonomic studies of
,
heads are much shorter and more compact freshwater limpets (Gastropoda:Basom-
(Fig. 2), and they are of essentially the matophora). I. The European Lake
same shape as found in all other Basom- Limpet, Acroloxus lacustris. Mala-
matophora (i.e., Ancylidae, EUobiidae, cologia, 1(1): 55-72.
Lymnaeidae, Physidae and Planorbidae). BURCH, J. ., BASCH, P. F. and BUSH,
From cytological considerations then L. L., 1960, Chromosome numbers in
one would not suspect that La í¿a neritoides ancylid snails. Rev. Portuguesa Zool.
has any close relationship with Acroloxus Biol. Ger., 2(3/4): 199-204.
8 The Ancylidae are usually considered to be the most phylogenetically advanced
basommatophorans (e.g., see Hubendick, 1947), but Burch (1962) has suggested that
they are perhaps among the most primitive.
^Taylor and Sohl (1962) created a separate supe rfamily for Acroloxus, the Acroloxacea.
BURCH, J. B. and HEARD, W. H., 1962, logical and histological fixing fluid.
Chromosome numbers of two species of Science, 118 (3058): 161.
Vallonia (Mollusca: Stylommatophora: PELSENEER, P., 1901, Etudes sur des
Orthurethra). Acta Biol. Acad. Sei. Gastropodes pulmones. Mém. Acad.
Hung., 12(1): 305-312. Roy. Sei., Lettr. Beaux-arts Belg., 54:
HUBENDICK, ., 1945, Phylogenie und 1-76, pis. 1-14.
Tiergeographie der Siphonariidae. Zur RETZIUS, G., 1906, Die Spermien der
Kenntnis der Phylogenie in der Ordnung Gastropoden. Biol. Untersuch., N. F.,
Basommatophora und des Ursprungs der 13(1): 1-36, pis. 1-12.
Pulmonatengruppe. Zool. Bidrag Up- SUTER, H., 1913, Manual of the New
psala, 24: 1-216. Zealand Mollusca. John MacKay, Wel-
,
1947, Phylogenetic relations lington, N. Z.,p 1-1119.
between the higher limnic Basom- TAYLOR, D. W. and SOHL, N. F., 1962,
matophora. Ibid., 25: 141-164. An outline of gastropod classification.
, 1960, The Ancylidae of Lake Malacologia, 1(1): 7-32.
Ochrid and their bearing on intra- THIELE, J., 1931, Handbuch der system-
lacustrine speciation. Proc. Zool. Soc. atischen Weichtierkunde. Jena 1929-35,
Lond., 133(4): 497-529. 2: 377-778.
, 1962, Studies on Acroloxus TRYON, G. W., 1884, Structural and
(Moll. Basomm.). Göteborgs Kungl. systematic conchology: an introduction
Vetensk. Vitterh.-Samh. Handl. Sjätte to the study of the Mollusca. 3: 1-453,
Följden. Ser. ,
9(2): 1-68. pis. 1-140.
HUTTON, F. W., 1882, Notes on some WALKER, ., 1925, New species of North
pulmonate Mollusca. Trans. New Zea- American Ancylidae andLancidae. Occ.
land Inst. 1881, 14: 150-158, pis. 3 and Papers Museum Zool., Univ. Mich., No.
4. 165: 1-13.
LA COUR, L., 1941, Acetic-orcein: A ZILCH, A., 1959, Gastropoda. 2. Euthy-
new stain -fixative for chromosomes. neura. In: Schindewolf, Handbuch der
Stain Techn., 16: 169-174. Paläozoologie. Lief. 1,6: 1-701. Born-
NEWCOMER, E. H., 1953, A new cyto- traeger, Berlin.
ZUSAMMENFASSUNG
ZYTOTAXONOMISCHE STUDIEN ÜBER DIE NAPFSCHNECKEN DES SÜSSWASSERS
II. DIE NEUSEELANDISCHE FLUSSNAPFSCHNECKE LATIA NERITOIDES
Latia neritoides Gray ist eine auf Neuseeland beschränkte SUsswassernapfschnecke,

die man allgemein zu den primitivsten unter den SUsswasserbasommatophoren rechnet
und fUr verwandt mit der südamerikanischen Chilina hält, aber gewöhnlich in eine
eigene Familie, die Latiidae, stellt.
Die haploide Chromosomenzahl von L. neritoides is 18, eine Zahl, die, obwohl
sie für die Basommatophoren im allgemeinen kennzeichnend ist, von der bisher
bei sämtlichen anderen SUsswassernapfschnecken (Ancylidae) gefundenen (15, 17)
abweicht, mit Ausnalime von Acroloxus lacustris (Acroloxidae). 1 rotz gewisser
gegenteiliger, auf anatomischen Vergleichenberuliender Hinweise, kann auf zytologischer
Grundlage keinerlei Verwandschaft zwischen Latia und Acroloxus festgestellt werden;
vielmehr scheint es, dass dieselben niciit verwandt sind.
YTOTAXO NOM Y OF LA TM 319
RÉSUMÉ
ETUDES CYTOTAXONOMIQUES SUR LES PATELLIFORMES D'EAU DOUCE

II. LA PATELLIFORME FLUVLATILE NEOZELANDAISE LATIA NERITOIDES
Latia neritoides Gray est une patelliforme fluviatile confinée à la Nouvelle Zélande.
Elle est généralement considérée comme un des basommatophores d'eau douce les
plus primitifs, apparenté en une certaine mesure aux Chilina de l'Amérique du Sud,
mais placé d'habitude dans une famille séparée, les Latiidae.
Le nombre haploïde de chromosomes de L. neritoides est de 18. Ce nombre, quoique
caractéristique des basommatophores en général, diffère de celui trouvé jusqu'à présent
chez les autres patelliformes d'eau douce (Ancylidae) examinées (15,17), s2mí Acroloxus
lacustris (Acroloxidae). Néanmoins, sur des bases cytologiques, au cune parenté
entre Acroloxus et Latia n'a pu être démontrée, contrairement à ce qu'indiqueraient
des comparaisons morphologiques. En fait, il semblerait qu'elles ne sont pas étroite-
ment apparentées.
RESUMEN
ESTUDIOS CITOTAXO NOM ICOS DE LAPAS FLUVIALES (GASTROPODA:
BASOMMATOPHORA). II. LA LAPA DE RIO NEOZELANDICA, LATIA NERITOIDES
Latia neritoides Gray es una lapita de agua dulce restricta a Nueva Zelandia. General-
mente se considera como uno de los basomatóforos dulceacuicolas más primitivos y en
alguna manera relacionado con las Chilina sudamericanas, pero constituye su propia
familia, Latiidae.
El número de cromosomas haploides en Latia es 18, y aunque éste es característico
de los basomatóforos en general, difiere de aquello que se encuentra en todas las otras
lapas de agua dulce (Ancylidae) investigadas (x=15, n=17), con excepción de Acroloxus
lacustris (Acroloxidae). Sin embargo, desde el punto de vista citológico, no se ha
).
podido demostrar relaciones entre Acroloxus y Latia, a pesar de que comparaciones
morfológicas parecieran sugerir lo contrario. En efeto no parece que tengan relación
cercana.
, latía NERITOIDES
(BProXOHOj-
.
Latia
.
neritoides
. .
Gray,
'-,-
-
,
Chilina,
Latiidae.
Latia neritoides 18,
.-
(Ancylidae),
(Acroloxidae).
. (=17, =15),
Acroloxus Latia,
Acroloxus lacustris
Malacologia will publish original monographs and Biological Sciences, 200 P Street, N.W., Washington
longer papers devoted primarily or exclusively to the 6, D. C. In particular, simplified practices, such
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DIRECTIVES
Malacologia publiera des monographies originales des rédacteurs.
et de longs mémoires consacrés principalement ou Manuscrits. est convenu pour le moment, qu'ils
exclusivement à l'étude des Mollusques. Il se propose pourront être en anglais, français, allemand,
de constituer un organe d'expression commune où espagnol ou russe. Y joindre un resume bref, mais
soient réunies les diverses branches de la suffisamment complet, pour traduction. Les manu-
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paléontologie, la physiologie, la taxonomie, la ligne et fournis en 2 exemplaires (le double sur
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possible, ou pour économiser de l'espace. Les Correspondance. Les manuscrits seront adressés
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phrase; les pourcentages seront représentés par le paiements et les demandes d'informations seront
signe %, et les abbreviations des mesures (après adressés soit au "Managing Editor",Dr. J. B. Burch,
un nombre), telles que mm, km, kg, n'auront ni Museum of University of Michigan, Ann
Zoology,
point, ni "s" au pluriel. Arbor, Michigan, A., soit à
U. S. .
J. Duncan,
Bibliographie. N'importe quel numéro récent de Department of Zoology, University of Liverpool,

Malacologie montrera la présentation désirée. England, U. K., ou à Mr. le professeur E. Fischer-
Separata. Les auteurs recevront gratuitement 25 Piette, Museum National d'Histoire, 55 rue de Buff on,
tirés à part, us pourront en obtenir davantage Paris V^, France,

Maíacoíog-M! publicará monografías originales, asi al uso de los siguientes factores simplificados: los
come estudios dedicados primera y exclusivamente al números no deben escribirse fuera, excepto al
estudio de los moluscos. Se propone proporcionar principio de una oración; los porcentajes (símbolo)
un medio común para los diferentes aspectos de la que siguen después de un número deben espresarse
Malacología, tales como Anatomía, Ecología, asi: %; las abreviaturas de medidas (después de
Malacología Médica, Paleontología Fisiología, y un número): mm, ml, kg, etc., no debiendo usarse
Taxonomía. Esta publicación tratará de combinar punto después de la abreviatura o eses en los
ambos, los artículos escolares con la rapidez en plurales.
la publicación. lodos los manuscritos serán En que se refiere a la bibliografía, véase un
lo
revisados por lo menos por dos editores. número reciente de Malacologia a cerca de la forma
Los manuscritos, por ahora, pueden ser en Inglés, de referencia.
Francés, Alemán, Ruso o Español. Deben ser con- Reimpresos. Los autores de artículos recibirán
cisos y adecuamente resanados para su traducción veinticinco copias gratis. Copias adicionales pueden
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leerse fácilmente aún después de reducidos. especialización o posición geográfica. Requisición
El estilo se cambiará tan poco como sea posible, de suscripciones, pagos e información deben ser
principalmente en lugares que se presten a ambigüe- enviadas al Director-Editor, J. B. Burch, Museo de
dades o donde la economía de espacio pueda Zoología, Universidad de Michigan, Ann Arbor,
:
asegurarse. Los autores de artículos deben remitirse Michigan, Estados Unidos de America.
,
, , - ,.,,- ; .;.., -
.,
^ : , ,-
, ..), -
, ,. , - ,(
.
KcLK
-
-
.:
..
%;
. .
25 -
, . -- . , ,
- -
,
-
. --
(
). .
-
.
Dr. J.
.
BURCH, Museum of Zoology, University of Michigan,
.
- -
-
Ann Arbor, Michigan, U. S. A., or to Dr. C. J.
DUNCAN, Department of Zoology, University of
Liverpool, Liverpool, England, U. K., or to Prof. E.
FISCHER-PIETTE, Museum National d'Histoire
Naturelle, 55 rue de Buffon, Paris V^ , France.
,
. .
..
Ij m?.
. .
. .
, 1963 .
.
161
163
(Bivalvia: Unionidae) 179

. M.
Lampsilis
(Bivalvia: Unionacea) 215
.
. . ,
, .. . X. . .
- 227
Hinkleyia
(Lymnaeidae, Stagnicola)
. .
..
. .
237
Taphius glabratus
-
(Say) ,
-
283
. .
Helisoma trivolvis
--
Echinoparyphium
291
. .
-
).
Achatina fúlica
. - 305
. .
2.
(, -
Latia neritoides 313
Vol. 1, No. 2 MALACOLOGIA July 1963
CONTENTS
Page
MALACOLOGIA, An International Journal of Malacology 161
G. L. WARMKE and L. R. ALMODOVAR

Some associations of marine mollusks and algae in Puerto Rico 163
J. J. PARODIZ and A. A. BONETTO

Taxonomy and Zoogeographie relationships of the South American
Naiades (Pelecypoda: Unionacea and Mutelacea) 179
A. M. CVANCARA
Clines in three species of Lampsilis (Pelecypoda: Unionidae) 215
H. VAN DER SCHALIE

Mussel distribution in relation to former stream confluence in
northern Michigan, U. S. A 227
D. W. TAYLOR, WALTER BURCH

H. J. and
Freshwater snails of the subgenus

from the western United States
J. B.
// (Lymnaeidae: Stagnicola)
237
H. W. HARRY and D. V. ALDRICH

The distress syndrome in Taphius glabratus (Say) as a reaction
to toxic concentrations of inorganic ions 283
T. C. CHENG
The effects of Echinoparyphium larvae on the structure of and
glycogen deposition in the hepatopancreas of Helisoma trivolvis and
gly oogenesis in the parasite larvae 291
A. R. MEAD
A flatworm predator of the Giant African Snail Achatina fúlica
in Hawaii 305
J. B. BURCH and C. M. PATTERSON

Cytotaxonomic studies of freshwater limpets (Gastropoda:
Basommatophora) II. The New Zealand River limpet, Latia
neritoides 313
"^
MUS. COMP. 200L
^^
I
LIBRARY
VOL 1 NO. 3 j^jj^g
I
!
JUIN 1 1 )4
HARVARD
university;
MALACOLOGIA

ANNE GISMANN,
MALACOLOGIA
General Editor B,BURCH, Managing Editor M. HUBER, Business Manager
19, Road 12 Museum of Zoology Museum of Zoology
Maadi, Egypt The University of Michigan The University of Michigan
U.A.R, Ann Arbor, Michigan 48104 Ann Arbor, Michigan 48104
U.S.A. U.S.A.

SCHRIFTLEITUNGSRAT CONSEIL DE REDACTION
P. . AGOCSY . HATAI W. L. PARAENSE

Magyar Nemzeti Múzeum Institute of Geology and Centro Nacional de Pesquisas
Természettudományi Múzeum Paleontology Malacológicas
Baross .13 Tohoku University . P. 2113, Belo Horizonte
Budapest, VIII., Hungary Sendai, Japan Minas Gérais, Brazil
J. J. PARODIZ
K. H. BARNARD N. A. HOLME Carnegie Museum
South African Museum Marine Biological Association
Pittsburg Pennsylvania
13,
Cape Town of the United Kingdom
U. S. A.
Republic of South Africa The Laboratory, Citadel Hill
Plymouth, Devon, England R. D. PURCHON
R. BOETTGER Dept. of Botany and Zoology
Technischen Hochschule
G. P. KANAKOFF Chelsea College of Science
Braunschweig
Los Angeles County Museum and Technology
Pockelsstrasse 10a
900 Exposition Boulevard London, S. W. 3, England
Germany
Los Angeles 7, California
S. G. SEGERSTRÂLE
U. S. A.
A. H. CLARKE, JR. Zoological Museum of
National Museum of Canada A. M. KEEN Helsinki Universit/
Ottawa, Ontario, Canada Department of Geology P.-Rautatiekatu 13
Stanford University Helsinki, Finland
C. J. DUNCAN Stanford, California, U. S. A.
F. STARMÜHLNER
Departmefat of Zoology
Zool. Institut der Universität
University of Liverpool Y. KONDO Wein 1, Lunegerring 1
Liverpool, England Bernice P. Bishop Museum
Austria
Honolulu 17. Hawaii, U. S. A.
E. FISCHER-PIETTE
N. MACAROVICI
J. STUARDO
Museum National d'Histoire Departamento de Zoologie
Laboratoire de Géologie
naturelle Instituto Central de Biologia
Université "Al. I. Cuza"
55 rue de Buff on Universidad de Concepción
lasi, Romania
Paris ye, France Cas. 301, Concepción, Chile
D. F. McMICHAEL W.S. S. VAN DER FEEN - VAN

A. FRANC The Australian Museum BENTHEM JUTTING
Faculté des Sciences
College Street Zoologisch Museum
55 rue de Buff on
Sidney, Australia Amsterdam, The Netherlands
Paris ye, France
J. E. MORTON CM. YONGE
P. GALTSOFF Department of Zoology Department of Zoology
Box 167
P. O. The University of Auckland The University
Woods Hole, Massachusetts Auckland, New Zealand Glasgow, Scotland
A. ZILCH
T. HABE W. K. OCKELMANN Senckenberg-Anlage 25
National Science Museum Marine Biological Laboratory
6 Frankfurt am Main 1
Ueno Park, Daito-ku Grjíínnehave, Helsing^r
Germany
Tokyo, Japan Denmark
MALACOLOGIA was established with the aid of a grant (NSF-G24250) from the National Science Foundation,
Washington, D, C, U. S. A.
MALACOLOGIA wurde unter Beihilfe einer Unterstützung (NSF-G24250) der National Science Foundation,
Washington, D. C, U. S. A., gegründet.
MALACOLOGIA fut établi avec l'aide d'une subvention (NSF-G24250) de la National Science Foundation,
MALACOLOGIA
Foundation, Washington, D. C, U. S. A.
,
fue establecida con la ayuda de una subvención (NSF-G24250) de la National Science
. (NSF - G24250) -
MUS. COMP. ZOOL
LIBRARY
JUIM 111964
HARVARD
UNIVERSITY
MALACOLXDGIA is prepared in the MoUusk Division, Museum of Zoology, The Univerisity of Michigan and
published at irregular intervals by the Institute of Malacology, 2415 South Circle Drive, Ann Arbor, Michigan,
U. S. A. The Sponsor Members of this Institute, also serving as editors are:
MALACOLOGIA wird in der "MoUusk Division, Museum of Zoology" der Universität von Michigan fertiggestellt
und in unregelmässigen Zeiträumen vom "Institute of Malacology", 2415 South Circle Drive, Ann Arbor, Michigan,
V. S. A. herausgegeben. Die gründenden Mitglieder dieses Institutes, die auch als Schriftleiter fungieren sind:
MALACOLOGIA est élaboré au Département de Mollusque de Musée de Zoologie du l'Université de Michigan
et publié à intervalles irréguliers par l'Institut de Malacologie, 2415 South Circle Drive, Ann Arbor, Michigan,
,
E. . A. Les membres du Comité de Patronage de cet Institut constituent en même temps le Comité de Publication
de MALACOLOGIA. Ce sont:
MALACOLOGIA es elaborada en la División de Moluscos del Museo de Zoología de la Universidad de Michigan
y publicada a intervalos irregulares por el Instituto de Malacología, 2415 South Circle Drive, Ann Arbor, Michigan,
.
. R. CARRIKER, President
Systematics-Ecology Program
:
S. A. Los miembros patrocinadores de este Instituto, quienes también sirven como editores, son:
Drive, Ann Arbor, Michigan, U. S. A.
E. G.
Laboratory
BERRY,
-
President-Elect-
Diseases
of Parasitic
2415 South Circle
R. ROBERTSON,
The Academy
Vice-President
of Natural
Marine Biological Laboratory National Institutes of Health Sciences of Philadelphia
Woods Hole, Massachusetts, U. S. A. Bethesda 14, Maryland, U. S. A. Philadelphia 3, Pennsylvania,
U. S. A.
A. G. SMITH, Vice-President D. W. TAYLOR, Vice-President N. F. SOHL, Secretary
Department of Invertebrate Zoology U. S. Geological Survey U. S. Geological Survey
California Academy of Sciences U. S. National Museum U. S. National Museum
San Francisco 18, California, U. S. A. Washington 25, D. C, U. S. A. Washington 25, D. C, U. S. A.
BURCH, Treasurer
J. B.
Museum of Zoology
The University of Michigan
Ann Arbor, Mich., 48104, U. S. A.
Participating Members of the Institute of Malacology are:

Aktiv teilnehmende Mitglieder des Malakologischen Institutes sind:
Les membres actifs de l'Institut de Malacologie sont:
Son miembros activos del Instituto de Malacología:
P. O. AGOCSY . HATAI J. J. PARODIZ

K. H. BARNARD N. A. HOLME R. D. PURCHON
R. BOETTGER G. P. KANAKOFF S. G. SEGERSTRÂLE
A. H. CLARKE, JR. A. M. KEEN F. STARMÜHLNER
J. DUNCAN Y. KONDO STUARDO
J.
E. FISCHER-PIETTE N. MACAROVICI W. S. S. VAN DER FEEN
A.
P.
A.
FRANC
GALTSOFF
GISMANN
D.
J.
W.
F.
E.
K.
McMICHAEL
MORTON
OCKELMANN
.
H. VAN DER SCHALIE
Z.
YONGE
ZILCH
T. HABE W. L. PARAENSE
Members-at-Large of the Institute of Malacology include all subscribers to MALACOLOGIA.
Alle Abonnenten von MALACOLOGIA gelten als ordentliche Mitglieder des Institutes,
Tous les abonnés de MALACOLOGIA sont Membres Associés de l'Institut de Malacologie.
Todos los suscriptores de MALACOLOGIA son miembros asociados del Instituto de Malacología.
Subscription price for MALACOLOGIA is Í5.00 (U. S. A.) or X. 1.16.- or fr. 25.00. Subscription requests, pay-
ments and enquiries can be sent to any of the Editors listed below:
Der Abonnementspreis für MALACOLOGIA beträgt jährlich S5.00 (V. S. A.) oder £1.16.- oder französische
fr. 25.00. Abonnementsbestellungen, Einzahlmigen und Anfragen sind an einen der unten angegebenen Schrift-
leiter zu richten:
Le prix de l'abonnement à MALACOLOGIA est de 5 dollars ou 1.16.- livres sterling ou 25 francs (français). Les
demandes d'abonnement, les paiements, et les demandes de renseignements, peuvent être envoyés a l'un des
membres de la liste suivante:
La suscripción de MALACOLOGIA es de US|5.00, o£l.l6, 25 francos franceses. Los pedidos de suscripción
y de información y los pagos pueden ser dirigidos a cualquiere de los Editores abajo mencionados:
J. B. BURCH J. DUNCAN E. FISCHER-PIETTE

Museum Zoology
of Department Zoology
of Museum National d'Histoire
The University of Michigan University of Liverpool naturelle
Ann Arbor. Mich. 48104, U. S. A. Liverpool, England, U.K. 55 rue de Buff on
Paris ve, France
Special INQUA issue of MALACOLOGIA
The Malacology plans a special issue of MALACOLOGIA on the occasion

Institute of
of the VII Congress of the International Association for Quaternary
International
Research (INQUA), Boulder and Denver, Colorado, U. S. A,, August 30-September 5,
1965. The issue is scheduled to appear in July 1965. Papers devoted primarily to
Pleistocene mollusks must be submitted to an appropriate editor by January, 1965,
and meet regular standards of the journal.
Numero especial INQUA de MALACOLOGIA
El de
Instituto Malacología planea para julio de 1965 un número especial de
MALACOLOGIA, en conexión con el VU Congreso Internacional de la Asociación
Internacional de Investigación del Cuaternario (INQUA), Boulder y Denver, Colorado,
. UU., 30 agosto - 5 septiembre de 1965. Los artículos, dedicados primariamente
a los moluscos del Pleistoceno y ajustados a los padrones de esta revista, deberán
ser sometidos a un editor apropiado alrededor de enero de 1965.
--
MALACOLOGIA, 1(3): 321-330
ONTOGENETIC CHANGES OF FORM AND MODE OF LIFE

IN TRESUS NUTTALLI (BIVALVIA: MACTRIDAE)!
Ross H. Pohlo2
ABSTRACT
There is a gradual change in mode of lifeduringthe ontogeny oiTresus nuttalli.
Young individuals inhabit the upper, unstable layers of the substrate, andaré
more subject to being exposed by water currents, while larger animals may be
buried to a depth of one meter and are more protected. Burrowing times reveal
that there is aprogressive loss of burrowing ability in the larger forms. Because
animals about 60 mm
long and over were slow burrowers, this size was taken as
a convenient limit to divide the individuals into 2 groups, those over and under 60
mm long. Shell morphology was examined to determine whether changes in shell
proportions are associated with the changes in mode of life.
A means of arranging correlated measures into groups, called ^ groups (Olson
and Miller, 1958), reveals that the small individuals have ahigher degree of inte-
gration than do the large. This is illustrated by the fact that diagrams repre-
sentingthe small forms show more bonds at a higher level of P than do the large
specimens. Features of shell shape are more highly interelated in the young indi-
viduals, indicating an association between shell shape structures and mode of life.
The young active forms possess a tightly knit pattern of shell features which is
associated with their greater burrowing ability.
Deformed rectangular coordinates show that there is an ontogenetic change in
orientation of the body and siphons in relation to the umbo. The long axis of the
siphons alters its position in such a way that the posterior tip moves dorsally
with respect to the umbo. Related to these changes in the body- siphons orien-
'
tation is a postive allometry of the posterior shell portion with relation to the
anterior. This allometric growth is apparently associated with the space re-
quirements of the large siphons and the change in placement of the body and
siphons. The position of the large animal wdthin the burrow is related to the
change in body orientation. An upward pull is exerted on the shell when the
siphons are being retracted. The shell is situated obliquely in the burrow, thus
apparently offering greater resistance to that force.
INTRODUCTION in juveniles and that it decreases with

increasing size. Accompanying the re-
Changes in mode of life are often production in burrowing activity, is a change
nounced between small and large individu- in form from juvenile to adult cockles.
same species of bivalves. Young
als of the Young Tresus nuttalli (Conrad, 1837)
burrowing forms live near the surface of {-Schizothaerus nuttalli) occupy the upper
the substratum, and are subject to being unstable layers of sand while larger indi-
exposed by water currents. Because of viduals may be buried to a depth of one
this danger, the power of reburial is meter. This paper deals with changes in
criticalat this stage, and small indi- form that are associated with this change
viduals are usually quite active, rapid in life habit. Quantitative and qualitative
burrowers. Kristensen (1957) described methods have been used to demonstrate
this phenomenon in Cardium edule L. and modifications in morphology that ac-
showed that burrowing ability is critical company the changes in life habits.
^Research supported (in part) by research grants, B-8755 and G-18767, from the National Science
Foundation, Washington, D. C. U. S. A. ,
^Department of Biology, San Fernando Valley State College, Northridge, California, U. S. A.
(321)
322
R. H. POHLO
angle of about 75 degrees with respect to
MODE OF LIFE
the surface (Fig. 2, arrow 1). Next, tne
Forty-three Tresus nuttalli, ranging in posterior retractors contract, pulling the
from 23 to 148 mm were collected
length posterior edge downward until the long
axis of the shell makes an angle of about
from mud flats in Tómales Bay, Cali-
fornia. 30 degrees with respect to the soil surface
This animal is a suspension feeder, (Fig. 2, arrow 2), During both of these
having long united siphons which make movements the animal is being drawn into
contact with the water-soil interface. the substratum towards the anchored foot.
Larger individuals live in deep burrows
where they rest on tneir antero-ventral
margin (Fig. 1). The antero-posterior
axis is thus slightly inclined to the surface
plane of the substratum.
FIG. 2. Burrowing movements. After the foot

is anchored, the. retractors contract
and cause the posterior edge to rise
(arrow 1) and then to move dowTiward
(arrow 2).
To assess relative speed of burial, the
burrowing times of 14 Tresus were taken.
Animals were placed under water (13°C)
on the mud flat near tne burrow from wliich
they were removed. Burrowing time was
calculated from the moment the foot
probed the soil until the shell disappeared
beneath the surface. Table 1 lists the
FIG. 1. The living position of a small (25 mm burrowing times for size ranked indi-
long) and large (85 mm
long) Tresus
viduals. Five specimens did not com-
nuttalli.Drawings from preserved
pletely rebury themselves within a period
specimens.
of 2 hours, although they made an attempt.
When an unearthed animal is placed on The observed burrowing times were con-
the mud flat, it reburies itself in much verted to rates, in millimeters per second
the same manner as Jordan (1915) de- (mm/sec). The rate "zero" was assigned
scribed for Mactra inflata. Tresus first to those animals which were not buried
probes the soil with the tip of the foot in 2 hours' time. The Spearman rank
and repeats this action until the foot is correlation test (r^) (Dixon and Massey,
deep in the substratum. It appears that 1951: 260-61) was employed to compare
the anterior retractor muscles then the ranks of shell size in mm
and the
contract raising the posterior shell edge rate of reburial in mm/sec. An r^ of
until the long axis of the saell makes an -0.770 (significant at the 0.01 level) was
ONTOGENETIC CHANGES IN TRESUS 323
obtained, indicating that larger animals point to divide the individuals into 2 groups,
burrow less rapidly. Also, in contrast to those over and under 60 mm
long. A total
large individuals, small forms begin of 20 small and 23 large individuals was
burrowing shortly after being unearthed. examined by quantitative and qualitative
Because animals around 60 mm
long and methods in order to illustrate differences
over usually were not buried after 2 hours, in morphology that occur in the rapid
this size was regarded as a convenient (small) and slow (large) animals.
TABLE 1. Burrowing rates of 14 Tresus nuttalli
Length in mm
324 R. H. POHLO
Length of the protractor (PL). Maximum those structures found to increase

dimension of the anterioi' protractor im- allometrically are discussed below.
pression. Bartlett's line of best fit was used to
not represent calculate the regression equations (see
These measurements do
Simpson et al., 1960: 232-37).
the shell morphology in all its complexity,
The techniques of morphological inte-
but they give a representative coverage
gration developed by Olson and Miller
of observed changes in those features
(1958) were used to show differences in
concerned with burrowing, and at the same
correlation patterns between the two size
time keep the number of measurements
groups. This method assumes that highly
within manageable limits.
correlated structures are the result of
ôc functional interdependence, developmental
and residual effects. Thus, if groups of
morphological dimensions can be shown to
vary together, an underlying biological
interrelationship can be ascribed to these
associations. Briefly, the steps in for-
mation of what is termed a p group {p
being the population correlation co-
efficient) are as follows (see Olsen and
Miller, 1958, for details). First, the
selected dimensions of the organism are
measured, and correlation coefficients for
each pair of measurements are calculated.
In the present study, this process resulted
in 105 correlation coefficients for each
size group, or 210 coefficients in all. All
of these correlations were positive. Next,
the sample correlation coefficient, r, is
translated into the lower limit of the
FIG. 3. Measurements taJ<en on Tresus population value/?. The 95% confidence
nuttalli.
interval was used in this study. Groups
Methods employed to analyze mor- of correlated measures are then obtained
phology are deformed coordinates, at any arbitrarily selected level of p (but
regression analysis, and correlation co- p must be greater than zero) by noting
efficients. which pairs of measures have p values
Deformed coordinates (Thompson, 1942) equal to or greater than the selected level.
are used to portray the differences in size The pairs that exceed this level are said
and shape of homologous regions of related to be bonded and are interconnected by a
organisms. Yonge(1952, 1953, 1955, 1958) series of lines. A grouping such as that
considered that the use of deformed in Figure 6 results, and comparison of
rectangular coordinates is incorrect when the resulting patterns of large and small
applied to both the mantle/shell and body specimens of Tresus, at the same level
of the Bivalvia, but Stasek (1963) used of p, is then possible. The p groups can
deformed coordinates and demonstrated be studied at any level of complexity
that, in his opinion, the criticisms of this merely by raising or lowering the value
method are invalid. of p. For example, if p is raised to a
Regression analysis was employed to high enough level all bonas wouia disap-
depict changes in proportion between pear. Conversely, ii p is just above zero
dimensions taken two at a time. Only many measurements are bonded.
MORPHOLOGY to lie with its long axis in a more antero-

ventral direction. In fact, the anterior
Deformed Coordinates region of the organism pivots in relation
Figure 4 illustrates the differences in to the posterior adductor and umbo. The
form observed in small and large indi- siphons also move from an original orien-
viduals of Tresus nuttalli. This shows that tation in which their long axis is directed
in the larger form the posterior edge of antero-posteriorly to a position where this
the shell is truncated in the region of the axis becomes more oblique.
siphons. Although the posterior edge Figure 4 also shows that the small
appears truncated, it grows with positive animal has a relatively elongated, pointed
allometry when compared to the anterior anterior shell margin while in the large
shell portion (shown below under Relative form this edge is rounder and not as
Growth). This aspect of shell form is the elongated.
result of greater shell growth from the Relative Growth
umbo to the point where the ventral margin All pairs of measurements taken on
of the siphons intersects the shell edge. the shell were plotted on separate
A general reorientation of the antero- bivariate scatter diagrams to de-
posterior axis of the body in relation to termine whether any changes in growth
occurred during ontogeny. A straight
I 2 3 4 5 67 line appeared to be the best fit for
most of the scatter diagrams, indicating
that there were no changes in growth
gradients. A few seemed to indicate
some curvature and were tested for
this phenomenon. Only one regression
proved to be significantly curved, this
being the comparison of the anterior
shell length (U) and the posterior shell
length (P) (Fig. 5). The estimating
equation is U = 0.879pl-216 xhis
indicates that the posterior shell portion
increases with positive allometry in re-
lation to the anterior.
Correlation
Figure 6 show the type of bonding that
occurs at three levels of ^ and indicates
that, in general, the p groups do not
segregate the organism into differing
functional units. Instead, a general pattern
of morphology emerges. Although these
groupings are difficult to interpret, some
interesting elements occur within the
overall pattern.
FIG. The deformation coordinates from
4.
a small (30
of
mm long) (A) to large (90
Specimens less than 60 mm long possess
a fairly complex type of bonding at tne p
mm long) (B) Tresus nuttalli.
- 0.950 level, while no bonds occur in
Drawings from preserved animals.
the large forms. A rather complete
the umbo accompanies this growth. The representation of shellrshape dimensions
posterior adductor retains the same occurs at this level (by shell-shape
position relative to the umbo, but the dimensions I refer to tne measures of
anterior adductor grows with a more length (L), height (H), and width (W) ),
postero-ventral component, so that this for all three measuresof shell shape enter
muscle, originally having its long axis the bonding. This group is not completely
oriented in a dorso-ventral position comes interbonded, because height (H) is not
Í
326
R. H. POHLO
ANTERIOR SHELL LENGTH
POSTERIOR SHELL LENGTH
20 30 60 80
FIG. 5. The relation ofanterior to posterior shell length using Bai'tletfs line of best fit. The
estimating equation is U = 0. 879Pl- ^16 (measurements in mm).
complex pattern; and width (W) and length

from the umbo to the anterior edge (U) do
UNDER 60 MM OVER 60 MM not enter the bonding although they are
bonded in the main pattern in the small
forms.
Although no index that expresses the
integration of the organism by a single
number, such as the index of integration
(Olson and Miller, 1958), was calculated,
it is evident from an inspection of Fig. 6
that in the young forms more measure-

ments are correlated at higher levels than
in the large specimens. This indicates a
higher degree of interdependence or inte-
gration of the organs associated with the
measurements.
DISCUSSION
In the early post-larval stages, many

PAW AAW bivalves are byssally attached; thus they
resist being swept away by water currents
FIG. 6. /ögToups for small and large Tresus (Yonge, 1962). But they soon lose their
nut ta Hi. byssus, and burrowing ability becomes
their means of maintaining positional
bonded to width (W). At p = 0.930 all stability. The capability to rebury thus
three of these measurements are inter- becomes critical for these juvenile clams.
bonded in the small forms. Fromunfigured Tresus nuttalli follows this trend, in that
data, this group is not completely formed the smaller forms are active and rebury
until p = 0.605 in the large animals. At themselves relatively quickly when
p = 0.920 the small individuals have a unearthed. Larger forms live at great
ONTOGENETIC CHANGES TRESUS 327
depths and are protected from dis- meets the ventral shell margin). This
turbances. Associated with their deep area is associated with the large siphons,
burial is a gradual loss of burrowing and with the change in orientation of the
activity and speed. body and siphons relative to the umbo.
The p groups reveal that features of The anterior portions of the body and
shell shape (L-H-W) show a greater inter- siphons grow with an increasing postero-
dependence in the small forms. It is ventral component in relation to the umbo.
suggested that adaptation to relatively The postero-ventral shell growth ac-
rapid burrowing is a prime factor in the commodates this change in orientation of
control of this pattern. The larger forms the body and siphon. It appears that
do not have the same selective pressures this change is, in part, an adaptation which
acting on features of shell shape as do the more securely anchors the animal in the
active small individuals. Also, the kinds substratum. A comparison of the orien-
of measurements comprising the pattern tation of the large and small individuals
are different in the two size groups. In within the burrow is shown in Fig. 1.
the large animals, the highest correlation This figure illustrates that the long axis
occurs for shell length (L) and anterior of large T. nuttalli is not perpendicular
adductor length (AAL), and the rest of the to the water-soil interface, and the least
pattern forms around these two elements resistance dimension (dotted line. Fig, 1)
(Fig. 6), suggesting that internal anatomy is not parallel to the long axis of the
is more highly associated in the large siphons. When the siphons are being
individuals. It is conceivable that, if retracted, this must exert an upward pull
more features of the internal anatomy, on the shell, tending to move the animal
such as the ctenidia or labial palps, were toward the surface. To counteract this
measured and the correlations de- upward motion, the animal is so oriented
termined, the large individuals might in the burrow that more resistance is
have higher correlations for these di- offered to the surrounding soil.
mensions than the small forms. In the The orientation of the animal within the
small animals, the central pattern- burrow (Fig, 1) and the mode of burrowing
forming elements (the two structures with (Fig, 2) are interrelated. Because the
the highest correlations) are two measures angle at which the shell is moved (Fig, 2)
of shell shape, namely length (L) and when reburying is similar to the position
height (H). This further points to the of the animal within the burrow, the
importance of shell shape, and thus organism seems merely to cease its
burrowing, features in the small forms. burrowing and is then in its final orien-
Thus the pattern and intensity of inte- tation within the burrow.
gration does change in the ontogenetic The change in form that occurs during
groups of T. nuttalli, and these variations ontogeny is illustrated in Fig. 4. Young
are associated with alterations in life specimens have a more pointed, elongated
habits. end, and appear to be more streamlined.
The regression diagram (Fig. 5) shows This shape is an advantage in burrowing,
that the posterior edge increases with for the anterior edge is more wedge-like
positive allometry relative to the anterior and helps penetrate the soil,
shell portion. The area of greatest
marginal increment (see Fig, 4) is that ACKNOWLEDGMENTS
between two lines, one extending from the
umbo to the point where the ventral margin I wish to thank Drs. Ralph Johnson, E,
of the siphons intersects the shell edge, W, Eager, and Charles R, Stasek, for
the other extending from the umbo to the critically reading this manuscript. This
posterior-most extension of the pedal gape work was completed while the author was
(where coordinate number 4 in Fig, 4 a Sverdrup Fellow in Oceanography at
328 R. H. POHLO
the Scripps Institution of Oceanography, 112(3): 195-214.

THOMPSON, D'Arcy W., 1943, On growth
LITERATURE CITED and form. Cambridge Univer. Press,
1116,
DIXON, W. J., and MASSEY, F. J., 1951, YONGE, M,, 1952, Studies on Pacific
Introduction to Statistical Analysis, Coast Mollusks IV. Observations on
McGraw-Hill, X + 370. Siliqua patula Dixon and on evolution
JORDAN, J,, 1915, Über die Art wie within the Solenidae. Univer. Calif.
Mactra inflata sich den Sand einwühlt. Publ. Zool., 55: 421-438.
Zool. Jb. Abt. Zool. Physiol., 35: 289- ,1953, The monomyarian con-
300. dition in the Lamellibranchia. Trans.
KRISTENSEN, I., 1957, Differences in Roy. Soc. Edinb., 62: 443-478.
density and growth in a cockle population ,1955, Adaptation to rock boring
in the Dutch Wadden Sea. Arch. Neer- in Botula and Lithophaga (Lamelli-
land. Zoologie, 12(3): 315-453. branchia, Mytilidae) with a discussion
OLSON, E. C. and MILLER, R. L., 1958, on the evolution of this habit. Quart.
Morphological Integration. Univer. of J. Micr. Sei., 96: 383-410.
Chicago Press, xv + 317. ,1958, Evolution within the bi-
SIMPSON, G. G., ROE, A., and LE- valve Mollusca. Proc. XV. Intern.
WONTIN, R. C, 1961, Quantitative Congr. Zool., sect. IV, Paper 26.
Zoology. Harcourt, Brace and Co., vii , 1962, On the primitive sig-
+ 440. nificance of the byssus in the Bivalvia
STASEK, R., 1963, Orientation and and its effect in evolution. J. Mar.
form in bivalved Mollusca. J. Morphol., Biol. Assoc, 42: 113-125.
ZUSAMMENFASSUNG
ONTOGENETISCHE VERÄNDERUNGEN IN GESTALT UND LEBENSWEISE
VON TRESUS NUTTALLJ (BIVALVIA: MACTRIDAE)
Man beobachtet beiTresus nuttalli eine allmähliche Veränderung der Lebensweise:

junge Individuen bewohnen die oberen und unbeständigen Schichten des Substratums und
sind daher dem Einfluss von Wasserströmungen eher ausgesetzt als die grösseren
Muscheln, die bis zu einer Tiefe von einem Meter gelagert sein können und dadurch
mehr geschützt sind. Messungen der Eingrabungsgeschwindigkeiten haben ergeben, dass
bei den grösseren Formen die GrabtUchtigkeit progressiv abnimmt und dass Tiere von
60 mm und darüber sich nur mehr sehr langsam eingraben. Es wurde daher dieses
Mass als geeigneter Ausgangspunkt für eine Unterteilung in 2 Gruppen angesehen für
Individuen über und unter 60 mm. Gleichzeitig wurde die Morphologie der Schale
untersucht, um festzustellen ob mit den Veränderungen der Lebensweise auch solche
in den Proportionen der Schale einhergehen.
Anwendung einer Methode (Olson und Miller, 1958), die wechselseitig bedingte
Messungen in Gruppen, / Gruppen genannt, zusammenfasst, hat ergeben, dass die
kleinen Individuen einen höheren Grad der Einordnung aufweisen als die grossen. Dieses
ist dadurch ersichtlich, dass die Diagramme für die kleinen Formen mehr Bindungen
auf einer höheren Stufe von ^ zeigen als die grösseren. Dass die einzelnen Merkmale
der Schalengestaltung bei den jungen Individuen in stärkerem gegenseitigen
Zusammenhang stehen besagt, dass zwischen Lebensform und Schalenform eine Beziehung
besteht: das fester gefügte Formenbild der jungen aktiven Formen hängt mit ihrer
grösseren Grabtauglichkeit zusammen.
Das Wachstum ist allometrisch. Deformierte rechtwinkelige Koordinaten zeigen
eine im Verlauf der Ontogenie auftretende Änderung der Orientierung des Körpers und
der Syphone mit Bezug auf den Umbo. Die lange Achse der Syphone verlagert sich
solchermassen, dass deren hinteres Ende mit Bezug auf den Wirbel dorsal verschoben
erscheint. Gleichzeitig besteht eine positive Allometrie der hinteren Muschelpartie
im Verhältnis zur vorderen. Dieses allometrische Wachstum steht augenscheinlich
im Zusammenhang mit der Notwendigkeit für die grossen Syphone Raum zu schaffen
wie auch mit der Veränderung der Lage des Körpers und der Syphone zueinander.
Ebenso steht die Lage der ausgewachsenen Muschel im Bohrloch in Zusammenhang
mit der Orientierungsänderung der Körperachse. Bei Einziehung der Syphone tritt
ein Zug nach oben auf. Die Muschelschale liegt dann schräg im Loch verankert wobei
sie offenbar diesem Zug einen grösseren Widerstand entgegenzusetzen vermag.
RESUME
CHANGEMENT DE FORME ET DE MODE DE VIE PENDANT L'ONTOGENIE
DE TRESUS NUTTALLI (BIWALVIA: MACTRIDAE)
Durant l'ontogénie de Tresus nuttalli l'on observe un changement graduel du mode de

vie: les individus jeunes habitent les couches supérieures et instables du substrat,
plus exposés ainsi aux courants d'eaux tandis que les individus plus grands peuvent se
trouver jusqu'à 1 m de profondeur et sont ainsi mieux protégés. Des observations sur
les durées d'enfouissement montrent chez les grandes formes und perte relative
progressive dans leur capacité de s'ensevelir. Comme les individus d'une longueur
d'à peu près 60 mm ou plus se terrent déjà lentement, cette taille fut prise comme
point de départ pour la division des spécimens en 2 groupes ("petits" et "grands")
selon que les individus avaient plus ou moins de 60 mm. La morphologie de leur coquille
fut étudiée pour voir s'il y avait, associée au changement du mode de vie, une
modification des proportions de la coquille.
Une méthode servant à disposer des mesures corrélatives en groupes, appelés
groupes /^ (Oison et Miller, 1958), a démontré que les petits individus ont un degré
d'intégration supérieur à celui des grands, du fait que les diagrammes montrent plus
de liens pour une valeur élevée de P chez les petites formes que chez les grandes.
Les traits structuraux de la coquille chez les individus jeunes montrent des inter-
relations plus étroites, indiquant un rapport entre forme de la coquille et mode de
vie. Les formes jeunes et actives possèdent un ensemble serré de caractères
conchyliologiques qui est relié à leur capacité de creuser.
La déformation de coordonnées rectangulaires indique un changement ontogénique
de l'orientation du corps et des siphons par rapport à l'umbo. Le grand axe des siphons
change de position de sorte que son bout postérieur se trouve déplacé dorsalement par
rapport à l'umbo; en même temps il existe une allométrie positive de la portion
postérieure de la coquille par rapport à sa portion antérieure. Cette croissance
allométrique est apparamment reliée aux besoins en espace des grands diphons et
aux changements de position du corps et des siphons. La position du grand animal dans
son trou est également en rapport avec ces changements d'orientation. Quand il rétracte
les siphons, une traction vers le haut est exercée sur la coquille: celle-ci se trouve
alors placée obliquement dans le trou, offrant ainsi une résistance accrue à la traction.
RESUMEN
CAMBIOS ONTOGENÉTICOS EN LA FORMA Y MODO DE VIDA EN
TRESSUS NUTTALLI (BIVALVTA: MACTRIDAE)
Un cambio gradual en el modo de vida es observado durante la ontogenia de Tressus

nuttalli. Los individuos jóvenes habitan las capas superiores e inestables del substrato
y están más sujetos a ser expuestos por las corrientes de agua, mientras que los
mayores se pueden enterrar a profundidades de un metro, quedando así mejor protegidos.
Existe una pérdida progresiva de capacidad excavatoria en las formas grandes. Debido
a que individuos de 60 mm
de largo, o mayores, son excavadores lentos, este tamaño
se indica como conveniente para limitar los dos grupos de individuos. La morfología
de la concha fué examinada para determinar si los cambios en sus proporciones están
asociados con los cambios en el modo de vida.
Un método de disponer medidas correlacionadas en grupos, denominados grupos /?
(Olson y Miller, 1958), revela que los individuos pequeños tienen un grado más elevado
de integración que los grandes. Esto se ilustra por el hecho que, diagramas repre-
sentando las formas pequeñas, muestran más vínculos a un alto nivel de p
que en las
grandes. Las características determinantes de la configuración de la concha están
330
R- H. POHLO
más altamente interrelacionadas en los individuos jóvenes, indicando una asociación

entre la configuración de la concha y el modo de vida. Las estructuras de la concha
en las activas formas jóvenes exhiben una disposición compacta que está asociada con
su mayor capacidad excavatoria.
Coordinadas rectangulares deformadas muestran que ocurre un cambio ontogenético
en la orientación del cuerpo y de los sifones en relación al umbón. El eje largo de los
sifones altera su posición de tal manera que la extremidad posterior se desplaza
dorsalmente con respecto al umbón. Relacionada con estos cambios de orientación
cuerpo-sifon, hay alometria positiva de la parte posterior de la concha con relación a
la anterior. Este crecimiento alométrico aparentemente está asociado con los
requerimientos de espacio de los grandes sifones y el cambio de ubicación del cuerpo
y de los sifones. La posición del animal grande dentro de la excavación está relacionada
con el cambio en orientación del cuerpo. Cuando los sifones se contraen, la concha
es empujada hacia arriba, y como está situada oblicuamente en la excavación, aparente-
mente ofrece mayor resistencia a esa fuerza.
A NEW ANAESTHETIZATION TECHNIQUE FOR FRESH WATER SNAILS,

TESTED ON LYMNAEA STAGNALIS
J. Lever, J. C. Jager and A. Westerveld
Department of Zoology
Free University
Amsterdam, The Netherlands
ABSTRACT
Current methods of narcotizing snails prior to fixation and dissection require
prolonged immersion (12-30 hours) in the various anaesthetic solutions. Faster
methods are desirable in biological experimentation. A recently developed
technique, requiring only 40 minutes of exposure, unfortunately caused swelling
and decrease of heart-beat in Lymnflea sía¿7UzZ¿s and moreover involved a long
period of recuperation.
Here an improved rapid anaesthetization technique is described, that is ef-
fective within 15 minutes in adult L. stagnalis and has but slight and brief after
effects. The substances used are, in combination, nitrogen gas to drive dis-
solved oxygen out of the water, and the narcotics nembutal, M. S. 222 (Sandoz)
and carbon dioxide. Optimum dosages and timing regarding 2 depth stages of
narcosis, the first adequate for the injection of drugs and the second suitable for
performing biopsies, are given for 2 temperatures.
At270(resp. 20°) stage I is reached after a 10 minute immersion in a 0. 08%
,
(resp. 0. 1%) nembutal solution, previously aerated with N2 for 3 minutes, if CO2
is bubbled through the solution during the latter 5 minutes of immersion. Stage
is attained in another 5 minutes by transferring to solutions of M. S. 222 (0. 07%
at 27° C; 0. 3%, in connection with 0. 1% nembutal, at 20° C).
For recovery, the snails need only be kept in tap water, preferably aerated.
At the temperatures quoted, the snails behave normally in 20 minutes to 2 hours.
It is expected that this method, perhaps with some modifications, will also
work with other species.
INTRODUCTION The long period needed for complete

anaesthetization is a disadvantage for
Several new methods for narcotizing biological experimental work, e.g. in
fresh water snails have recently been physiological or endocrinological work
described: VAN DER SCHALIE (1957) involving injections and in extirpation
used nembutal after stimulation with (see HEKSTRA and LEVER, 1959; LEVER
alcohol, McCRAW (1958) combined et al. ,
Therefore a more rapid
1961).
nembutal with menthol, while VAN EEDEN method was developed by JOOSSE and
(1958) prescribed a mixture of menthol LEVER (1959). It was found that when
and chlor alhydrate. After 12-24, 16- Lymnaea stagnalis that have previously
18, and 15-30 hours respectively, the been treated with a 0.08% nembutal so-
snails are insensitive and well-extended, lution for 20-30 minutes, at 21-25^0,
and thus are suitable for anatomical are then immersed in a solution of 0.08%
dissection and for fixation. nembutal + 0.3% M. S. 222^ for about 10
The anaesthetic M. S. 222 isa meta- amino- benzoic acid-ethylester in the f orm of a methan-
sulphonate (Sandoz, Basle), often recommended for use with cold-blooded animals (e.g. ROTHLIN,
1932; McGOVERN andRUGH, 1944; RANDALL, 1962).
(331)
332 LEVER, JAGER AND WESTERVE LD
minutes, they are anaesthetized so deeply are unfavourable. The prolonged after-
that they are ready for operation. For treatment is another difficulty.
full recovery after excision, the snails It was therefore attempted to find a
are left in running tap water till the technique having a minimum effect upon
following morning. For the next 2 days heart -beat and body weight, and a short
they are kept in aerated tap water that is recovery period. Satisfactory results
renewed twice a day. Thereafter aeration were obtained by using not only aqueous
is discontinued, but the water is changed solutions of nembutal and M. S. 222, but
daily. also the gases nitrogen and carbon dioxide.
During and shortly after the anaes-
thetization procedure the body weight of DESCRIPTION OF METHOD
the snails was found to increase by ap-
proximately As already shown by
10%. The procedure is divided into 2 stages:
HUF (1934), narcotization-swelling
this after the first treatment the snails (adult
is a result of a disturbance of the water specimens of Lymnaea stagnalis with a
balance. Most probably this effect is shell-length of 30 - 35 mm) are suf-
caused mainly by a decrease in heart ficiently insensitive for an injection. But
activity: during anaesthetization, the for an operation the second stage is
frequency of the heart -beat often de- necessary to obtain the required degree
creases from about 30 to 5 per minute, of anaesthesia.
while in some cases the heart even stops The dosages and timing that appear to
completely (LEVER et. al., 1961). give the best results were arrived at
It is clear that, especially for experi- after many experiments and are summa-
mental studies on organs which influence rized in Table I. The speed of narco-
the water balance, these phenomena tization was found to be markedly
(swelling and decrease of heart activity) dependant on temperature. Indications
TABLE L Scheme of the anaesthetization technique

A NEW ANAESTHETIZATION TECHNIQUE 333
are here given for 2 temperatures: 20^ skin is exposed to the nembutal solution
and 27^C. The snails were treated in in a uniform way. After 5 minutes the
400 ml glass jars, placed in a water- snails are anaesthetized to such an extent
bath at constant temperature. that they do not retract when CO2 is bubbled
Stage I, appropriate for injections . through the solution, again by means of a
Initially an attempt was made to obtain porous stone. After another 5 minutes,
sufficiently deep anaesthesia with carbon or 10 minutes in all, the snails are ready
dioxide alone. It was found that, after a for injection.
stay of 15 minutes in a closed vessel Stage II, appropriate for operations .
entirely filled with soda-water, many pond Although injections can be given to snails
snails are relaxed enough for injection. after a 10-minute exposure, their anaes-
When placed in tap-water, these snails thesia is not sufficiently deep for
return to normal within another 15 minutes operations: incision of the skin or even
and no special after-treatment is neces- manipulation causes them to retract
sary. However, not all snails react in slowly into the shell. This can be avoided
this manner. A number of them retract by additionally treating the snails with
into the shell and never assume the well- M. S. 222. After studying the effect of
extended position needed for a successful various concentrations and mixtures, it
injection. Results are even less acceptable was found that highly satisfactory results
when the snails are put in tap -water that are obtained by using a mixture of 0.1%
is then aerated with carbon dioxide, nembutal + 0.3% M. S. 222 at 20^ C, and,
because the snails retract as soon as at 270 C, a solution of 0.07% M. S. 222
aeration starts. Moreover, aC02-anaes- for 5 minutes (see Table I). In this
thesia is never deep enough to allow manner the snails are ready for operation
operations. in 15 minutes. They are completely anaes-
Thus, with carbon dioxide alone, good thetized, extend out of their shells and do
anaesthesia can not be obtained in these not show movements of any kind when
fresh water snails. It was therefore manipulated or operated upon.
decided to investigate whether a combi-
nation of CO2 -treatment with the earlier TREATMENT FOR RECOVERY
nembutal - M. S. 222 method would shorten
the reaction time of that method, and Recovery from anaesthesia by the above
therefore also the period required for method is prompt and is achieved by the
recovery. This was indeed the case. simplest of treatments: the snails are
The method was further considerably merely kept in tap water, preferably
improved when nitrogen-gas was also aerated, and will resume normal activities
used. within a short time. As an example, mean
A nembutal solution (0.1% at 20^ C, recovery times at temperatures ranging
0.08% at 270 C) is aerated with nitrogen from 16° - 27° in stagnant, running
for 3 minutes. The gas is introduced via and aerated tap water are given in Table
a porous stone, so that it escapes in . In this experiment 6 groups of 4 snails
very small bubbles. In this way the each were anaesthetized at 20° and then
dissolved oxygen and carbon dioxide are given an injection of 0.1 ml Ringer so-
driven out of the solution. When the snails lutions The figures demonstrate that,
are immersed in this solution immediately with equal conditions of anaesthesia,
after nitrogenation, the lack of oxygen temperature of the medium is the most
forces them all to extend out of their shells. important factor in speed of recovery,
In this manner a considerable area of the but that at equal temperatures, renewal
2c omposition in gram per liter: NaCl 1.37, KCl 0.21, CaCl2 0.33, MgClg u.49, NaHCOgl.g?,
NaH2P04 0. 104.
334 LEVER, JAGER AND WESTERVELD
TABLE II. Effect of medium on the recovery period of Lymnaea stagvalis after
anaesthesia (stage I at 20° C) and injection of 0. 1 ml Ringer solution
Medium after
the treatment
Table demonstrates that the periods

III counting the operating period itself, to
for heart recovery after anaesthesia, 40 minutes. It is expected that this method
whether it has been followed by treatment of anaesthetization can be applied, with
or not, entirely tally at 27°C (groups 3 some modifications if necessary, to other
and 4; 7 and 8), and correspond well species of fresh water snails.
enough at 20° C.
Comparison of data given in Tables III LITERATURE CITED
and II shows that mean recovery times
following stage I anaesthesia at 20° C, HEKSTRA, G. P. and LEVER, J., 1959,
in stagnant water at 20° (Table 1, Some effects of ganglion-extirpations in
group 2) correspond fairly well with Limnaea stagnalis. Proc. kon. ned.
findings in running water (Table II), heart- Akad. Wetensch., Amsterdam, C63:
beat returning to normal in 36 - 42 271-282.
minutes in the former case, while crawling HUF, ., 1934, Über den Einflusz der
was resumed after 44 minutes in the latter Narkose auf den Wasser- und Mineral-
case. When the narcotization
either haushalt bei SUszwassertieren. Pflüg-
temperature (Table or the temperature
III) ers Arch. ges. Physiol., 235: 129-140.
of the recuperation medium (Table II) is JOOSSE, J. and LEVER, J., 1959, Tech-
raised by 5-7°C, recovery times are niques of narcotization and operation
practically halved, amounting to 21 for experiments -with Limnaea stagnalis
minutes for heart-beat and 20- 24 minutes (Gastropoda P\ilmonata). Proc. kon.
for crawling. ned. Akad. Wetensch., Amsterdam, C62:
It is noteworthy (Table III) that recovery 145-149.
from rapid anaesthesia (Stages I and II), LEVER, J., JANSEN, J. andDE VLIEGER,
done at 27°C, is rapid (21-25 minutes) even T. A,, Pleural ganglia andwater
1961,
ifthe temperature of the recovery medium balance in the fresh water pulmonate
is only 20°C) and the water stagnant. The Limnaea stagnalis. Proc. kon. ned,
effect of the high anaesthetization temper- Akad. Wetensch,, Amsterdam, C64: 531-
ature (27 ° C) on the speed of recovery is 542,
especially marked in stage II anaesthesia, McCRAW, B. M., 1958, Relaxation of
but even when done at 20° recovery is snails before fixation. Nature, 181: 575.
almost complete in about 2 hours. McGOVERN, B. H. and RUGH, R., 1944,
Efficacy of M-amino ethyl benzoate as
CONCLUSION an anaesthetic for amphibian embryos.
Proc. Soc. exp. Biol, and Med., 57:
With the method described above it is 127-130.
possible to inject drugs into adult speci- RANDALL, D. J,, 1962, Effect of an
mens of Lymnaea stagnalis after 10 anaesthetic on the heart and respiration
minutes and to carry out operations after of teleost fish. Nature, 195: 506,
15 minutes. The duration of the after- ROTHLIN, E., 1932, M. S. 222 (lösliches
effects is always short. At 27° snails Anaesthesin), ein Narkotikum für Kalt-
are back to normal in 20-25 minutes, blüter. Schweiz, med, Woch.-Schr,, 62:
when anaesthesia is followed by an 1042,
adequate and simple after-treatment. VAN DER SCHALIE, H,, 1957, Nembutal
Under optimum conditions the time during as a relaxing agent for mollusks. Amer.
which the snails are disturbed by the Midland Naturalist, 50: 511-512,
injection process can be reduced to a VAN EEDEN, J, A., 1958, Two useful
minimum of 30 minutes and the corres- techniques in fresh water malacology,
ponding disturbance for biopsy, not Proc, malacol, Soc, London, 33: 64-66,
336 LEVER, JAGER AND WESTERVELD
ZUSAMMENFASSUNG
EIN NEUES ANAESTHESIEVERFAHREN FÜR SÜSSWASSERSCHNECKEN,

AN LYMNAEA STAGNALIS ERPROBT
Die bisher angewendeten Betäubungsmethoden für Schnecken vor Fixierung und

Sektion benötigen längeres Eintauchen (12-30 Stunden) in den verschiedenen anaest-
hesierenden Lösungen. Für biologische Versuche sind raschere Methoden
wünschenswert. Ein vor kurzem entwickeltes Verfahren, dass nur 40 Minuten Eintau-
chen erfordert, verursachte leider Schwellung und verlangsamten Herzschlag in Lymnaea
stagnalis und benötigt ausserdem eine lange Erholungszeit.
Hier wird nun ein verbessertes schnelles Verfahren beschrieben, das bei
ausgewachsenen L. stagnalis innerhalb von 15 Minuten wirkt und nur kurze und leichte
Nachwirkungen hat. Es werden dabei die folgenden 4 Substanzen angewendet: zuerst
Stickstoffgas, um den im Wasser gelösten Sauerstoff auszutreiben, sowie dann die
Anaesthetica Nembutal, M. S. 222 (Sandoz) und KohlensSuregas. Optimale Dosen und
Zeiten sind hier für 2 Temparaturen und ftlr 2 Narkosetiefen angegeben, deren Erste
für die Injektion von Drogen ausreicht und deren Zweite genügend tief für Biopsie ist.
Bei 27°(bzw. 20°) wird das 1. Stadium bereits nach 10 Minuten in einer Nembutal-
Lösung von 0.08% (bzw. 0.1%) erreicht, durch die man vorher 3 Minuten lang NÍ2
geleitet hat, wobei man aber in den letzten 5 Minuten der Immersion auch CO2 durch
die Lösung führen muss. Das 2. Stadium wird in weiteren 5 Minuten erreicht, wenn
man die Schnecken in M.S. 222 Lösungen überführt, und zwar in eine von 0.07% bei
27° und eine von 0.3%, zusammen mit 0.1% Nembutal, bei 20°
Zur Wiederherstellung genügt Aufenthalt in vorzugsweise durchlüftetem Leitungs-
wasser. Bei den oben genannten Temperaturen verhalten sich die Schnecken in 20
Minuten, bzw. 2 Stunden wieder normal.
Es ist anzunehmen, dass diese Methode, vielleicht mit leichten Abänderungen, bei
anderen Arten ebenfalls wirksam sein wird.
RESUME
UNE NOUVELLE TECHNIQUE D'ANESTHESIE POUR MOLLUSQUES D'EAU DOUCE,
ESSAYEE SUR LYMNAEA STAGNALIS
Les méthodes courantes d'anesthésie des mollusques avant fixation et dissection

exigent une immersion prolongée (de 12-30 heures) dans les diverses solutions anes-
thésiantes. Pour l'expérimentation biologique, des méthodes plus rapides sont
préférables. Un procédé récemment développé, ne requérant que 40 minutes
d'immersion, provoquait malheureusement le gonflement du corps et le ralentissement
des battements du coeur chez Lymnaea stagnalis et, en plus, nécessitait une période
de rétablissement assez longue.
Ici est décrite une méthode améliorée rapide, qui agit en 15 minutes sur les spécimens
adultes de L. stagnalis, et dont les effets accessoires ne sont que légers et de courte
durée. Les substances utilisées, en combinaison, sont: de l'azote gazeux (pour chasser
de l'eau l'oxygène dissout) et les narcotiques "Nembutal", "M.S. 222" (Sandoz) et le
dioxyde de carbone. Les dosages optimaux et temps requis pour 2 stades de
narcotisation, dont le premier convient à l'injection de drogues et dont le second est
suffisamment profond pour permettre la biopsie, sont donnés pour 2 températures.
A 27° (resp. 20°) le premier stade est atteint 10 minutes après immersion dans
une solution de Nembutal de 0.08% (resp. 0.1%), aérée au préalable à l'azote gazeux
pour 3 minutes, si l'on y fait passer du CO2 pendant les 5 dernières minutes de
l'immersion. Le second stade est atteint 5 minutes après, par simple transfert à
des solutions de M. S. 222 (d'un taux de 0.07% à 27° C: et de 0.03%. avec 0.1% de
nembutal, à 20°).
Pour récupérer, les mollusques sont simplement placés dans de l'eau de conduite,
aérée de préférence. Aux températures données, ils se comportent normalement
après un délai de 20 minutes à 2 heures.
Il est probable que sette méthode, peut-être avec quelques modifications, s'avérera
efficace chez d'autres espèces de mollusques aquatiques.

RESUMEN
NUEVA TÉCNICA DE ANESTESIA PARA CARACOLES DE AGUA DULCE
ENSAYADA EN LYMNAEA STAGNA LIS
Los métodos corrientes para narcotizar caracoles antes de la fijación y disección
requieren inmersión prolongada (12-30 horas) en las varias soluciones anestésicas.
Aqui se describe una técnica rápida que permite anestesiar en 15 minutos los adultos
de L. stagnalis y cuyos post-efectos son leves y breves. Las substancias usadas son,
en combinación, gas nitrógeno, para acarrear fuera del agua el oxígeno disuelto, y los
narcóticos nembutal, M. S. 222 (Sandoz) y dióxido de carbono. Las dosis y duración
óptimas con respecto a dos grados de narcosis, adecuados el primero para la inyección
de drogas y el segundo para ejecución de biopsias, se dan para dos temperaturas.
A 270 (resp. 20°) C. el grado I se alcanza después de 10 minutos de inmersión en
solución de nembutal al 0,08% (resp. 0,1%) previamente tratada con N2 por 3 minutos,
si se burbujea CO2 a través de la solución durante los últimos 5 minutos de inmersión.
El grado se alcanza en otros 5 minutos por transferencia a soluciones de M.S. 222
(0,07% a 27° C; 0,3% en combinación con 0,1% nembutal a 20° C).
Para recuperación, los ejemplares sólo necesitan ser colocados en agua corriente,
de preferencia aireada. A las mencionadas temperaturas los caracoles reasumen su
comportamiento normal entre 20 minutos y 2 horas. Es de esperar que este método,
quizá con algunas modificaciones, sea aplicable a otras especies.
THE UPTAKE OF RADIOACTIVE ZINC, CADMIUM AND COPPER BY

THE FRESHWATER SNAIL, TAPHIUS GLABRATUS
Charles M. Yager^ and Harold W. Harry^
ABSTRACT
Laboratory reared Ta phius glabratus (ûstralorbis or Planorbina of authors)
were exposed to concentrations of zinc 65, cadmimn 115M or copper 64 which
allowed normal behavior, and to concentrations of the same metals which re-
sulted in the snail showing the distress reaction (incapacity to retract into the
shell). Snails were exposed in groups of 6-10, to volumes rangingfrom 10-200ml
of dosing solution per snail, and for exposuretimes of 6-72hours. Much-variation
was found in the uptake of metal ions by the snails exposed in each experiment,
and this is thought to be due to some intrinsic property of the snails perhaps
,
depending on the activity of each snail while in the dosing solution. Pairs of
snails showing counts of the intact body closest to the average were selected for
analysis of the uptake of various body parts. After removing the shell and
dividing the protoplasmic body into definite sections ,these were weighed and
digested in concentrated nitric acid, following which counts of radioactivity were
made using appropriate apparatus. An average difference of 42.5% between the
counts per minute per milligram of the intact, live snail and summation of counts
of the body parts after digestion was attributed to the difference in geometry of
counting. The difference was about equally divided between loss and gain after
dissection, and showed no correlation with the weight lost in dissection, which
averaged 18.1%. The uptake patterns were essentially the same for all 3
metals. In normal snails, more metal ions were taken up by snails exposed to
larger volumes of dosing solution than was taken up by snails exposed to only 10
ml of solutions of the same concentration for equal periods of time. The uptake
by the shell, in counts/min. /mg, decreased in experiments in
which snails were exposed to 10 ml of dosing solution per snail, as time in-
creased. In a given experiment, the uptake was of the same order of magnitude
for all of the protoplasmic body except the liver, which showed 4-7 times the
amounts present in other tissues. The liver in distressed snails had only twice
the amount of metal found in other tissues. Snails which showed normal activity
often had much larger amounts of metal present in the protoplasmic tissues than
did distressed snails. It was concluded that distress is dependent on the concen-
tration of an ion, rather than the amount of an ion absorbed, and that the ions
are effective in producing distress by disrupting membrane permeability rather
than by interfering with some internal metabolic mechanism.
In previous studies (Harry and Aldrich, found to be 0.1 to 1.0 ppm. Lower con-
1963) was found that
it zinc, cadmium centrations of these ions allowed normal
and copper were among the more toxic behavior, i.e., the snail was extended and
of 22 ions tested on the planorbid snail, crawling. The present study was under-
Taphius glabratus (Say) {Australorbis or taken to gain additional information on
Planorbina of authors). At concentrations the effect of these 3 heavy metal ions in
of 0.050 to 0.10 ppm ( parts per million, relation to the normal and distressed
or milligrams per liter ) cadmium and states of this snail, by using radio-
copper produced a condition termed active isotopes. Since this study was
distress, in which the snail was extended completed, an important paper on the
but unable to attach the foot or crawl. uptake of radioactive sodium, strontium,
The distressing range of zinc ions was rubidium and iron by this snail has been
Physics Department, North Georgia College, Dahlonega, Georgia, U.S.A.
^Research Associate, Biology Department, Rice University, Houston 1, Texas, U, S. A.
(339)
340 YAGER AND HARRY
published by Johnson et al. (1962), in 424.7 - 5.4 mg (SEM for 192 snails).
which they review the few previous papers No food was given to the snails during
on the uptake of radioactive ions by fresh- the experiments. Snails were removed
water snails. from the aquaria, rinsed briefly in dis-
Our work was supported in part by tilled water, and placed directly in the
grants from the Atomic Energy Com- dosing solution.
mission and the World Health Organi- In these studies constant concentrations
zation. We are indebted to Dr. Clark of radioactive metal ions were used with
P. Read and Dr. David V. Aldrich for 2 patterns of exposure: (1) Snails were
much constructive criticism on the manu- exposed to a constant volume of dosing

script, and to Mr. Donald M. Denney and solution, 10 ml per snail, for periods
Mr. Bruce Duncan for technical varying from 6-72 hours, to study the
assistance. variation of uptake with time. (2) With
constant exposure time, usually 48 hours,
MATERIALS AND METHODS the volumes of dosing solution per snail
was varied from 10 - 200 ml, to study the
All snails were raised in the laboratory effect on the uptake of larger amounts of
from a stock originating in Puerto Rico. ions in the environment. Usually the snails
Initially they were maintained in charcoal used (6 or 10) in any one experiment were
filtered tap water (50 to 70 ppm total exposed in common in one beaker of
dissolved solids) in plastic or enamel solution.
rectangular pans each containing about 4 After exposure to the dosing solution,
liters water and 50 snails.
of Snails each snail was dried on soft paper tissue
raised under these relatively poor and and weighed. The count of the whole, live
crowded conditions weighed an average of snail was then determined, using a
210.3 ±7,0 mg per snail (SEM, or standard scintillation counter for zinc and cadmium,
error of the mean, of a sample of 80 and by placing the snail near the window
snails). These were used in only a few of a Geiger counter in the case of copper.
experiments, as specifically noted in the After determining the counts per minute
text, below. per milligram for each snail, pairs of
Healthier stock was later obtained by snails closest totheaverage were selected
maintaining colonies of several hundred for further analysis.
snails in frame aquaria of 80 liters The differential uptake of the ions in
capacity, covered with glass plates. Again different parts of the snail was determined
tap water was used, filtrated with charcoal as follows. To remove the animals from
before the introduction of the snails, and the shell they were plunged into distilled
with continuous filtration thereafter. A water heated to 65 -70° for 30 seconds,
small amount of reagent grade calcium after which the cephalopedal mass was
carbonate powder was added to the water grasped with fine forceps, and the soft
weekly (several grams, estimated). Only tissues could be removed from the shell
fresh lettuce was used as food. No sand intact. With care, this extraction can be
or other substrate material was present made without breaking the soft tissues or
in the aquaria, nor any aquatic plants. leaving blood in the shell. We are indebted
Fecal material was siphoned from the to Dr. Bengt Hubendick for suggesting this
bottom of the tanks about monthly, and useful technique.
charcoal filtered tap water used to re- Snails with shells removed were placed
place the water removed. Pieces of thin in a dry Petri dish. Under a dissecting
polyethylene plastic sheets were placed microscope they were divided into
in the aquaria because the snails show a cephalopedal mass, pulmonary complex
predilection for depositing eggs on this and upper visceral complex, as shown
material. Snails raised in these aquaria in Fig. 1 B. The cephalopedal mass
and used for the experiments averaged contains the bulk of the nervous system.
UPTAKE OF RADIOACTIVE ZN, CD AND CU BY TAPHIUS 341
-GONAD
^ HERMA PHROD I TIC
DUCT
-LIVER
STOMACH
ALBUMEN GLAND
HEART-
PULMONARY
CAVITY-
RECTUM
KIDNEY-
HYPOPEPLAR
CAVITY-
KIDNEY
PULMONARY
CAVITY
VAS DEFERENS
VI DUC
RECTUM
-ISTHMIAN HEMOCOEL
ESOPHAGUS
-COLUMELLAR MUSCLE
with shell removed.
FIG. 1. Diagram of the major aspects of the topographic anatomy of Taphius glabratus
A: Lateral view; B: Dorsal view; C: Cross section of the pulmonary complex about
midway of its length.
Heavy dashed lines (A and C) indicate the sections separated for analyses.
342 YAGER AND HARRY
much cutaneous musculature, the buccal also used in the dosing solutions.
mass, salivary glands, part of the eso- The radioactive isotopes were obtained
phagus, penis, preputium and part of the from Oak Ridge National Laboratory, and
vas deferens. The pulmonary complex were Cu64 as Cu(N03)2 in HNO3, Cd ll^M
has 2 parts: (1) The large pulmonary as Cd(N03)2 in HNO3 and Zn65 as ZnCl2
cavity lined with epithelium which is in HCl. Data on amounts of metals in
continuous with that of the mantle through this paper refer to the weights of the
the pneumostome. This cavity holds metal ions, rather than to the compounds
varying amounts of water or air. In its containing them. Data on amounts of
roof are the heart and kidney. (2) The radioactivity are in counts per minute
isthmian hemocoel, which connects the per milligram of snail tissue or shell,
hemocoel of the upper visceral complex unless specifically noted. Corrections for
with that of the cephalopedal mass and decay of the isotopes, as well as for
extends laterally into the roof of the background count were made on all data
pulmonary cavity. This part contains the reported herein. In some experiments,
spermathecal sac, the columellar reas noted, non-radioactive reagent grade
tractor muscle, oviduct and uterus, most CuSO^ and Cd(N03)2were added to provide
of the vas deferens and prostate, the a greater concentration of metal ions than
esophagus and most of the anterior aorta. could be conveniently produced using only
Most of the major organs of the upper radioactive material. The anions included
visceral complex are shown. The mantle are known from previous work to be non-
epithelium of this area has very little toxic at the concentrations used. The
musculature underlying it. All 3 major pH of the dosing solutions was checked
body parts contain blood and connective initially, and adjusted to pH 4.5 to 6.0
tissue (see also Malek 1962). In some with reagent grade KOH when necessary,
experiments the upper visceral complex but no buffer was added. During the
was further subdivided into Sparts: gonad, experimental exposure, beakers holding
liver (with hermaphroditic duct and upper the dosing solution and snails were placed
loop of the intestine) and stomach-albumen in a water bath held at 27 ± 1°C.
gland complex (containing the lower loop The dosing solutions were prepared by
of the intestine) (Fig. lA), In a few mixing enough of the radioactive stock
experiments the pulmonary complex was solutions with distilled water to produce
also subdivided longitudinally, as shown in counts within a reasonable range. Suitable
Fig. 1 (see also Malek, 1962). amounts were found to be of the order of
The comparable body parts from the 2 0.25 microcuries per milliliter of the
snails of a selectedpair were then weighed dosing solution when the scintillation
and digested together in 2 ml of concen- counter was used, and of the order of 0.1
trated nitric acid, with gentle heat. In ixc/ml when the Geiger counter was
the zinc and cadmium experiments the employed. These amounts in terms of
liquid digests were placed in a sodium radioactivity represented amounts in
iodide, thallium activated, well-type terms of weight which were usually well
scintillation counter to count the gamma below the concentration of the metals which
radiation of these elements. To count produced distress.
the beta radiation of copper, the digests
were evaporated to dryness in planchettes RESULTS
under an infrared lamp, and counted with
a Geiger tube and suitable scaler. The Uptake of Zinc and Cadmium
All glassware used was Pyrex,
chemically cleaned and rinsed in distilled The counts made of the whole snail
water which had been passed through an immediately after the exposure period
ion exchange column after being distilled showed considerable variation in uptake
in a metal still. Water so prepared was of the isotope for all 3 metal ions. More-
TABLE 1. Uptake of radioactive zinc and cadmium by Taphius glabratus at varying exposure
times and volumes of solution.
Zinc 65 (at 0.019 ppm)

344 YAGER AND HARRY
to the glass Petri dish and stainless steel decrease in count after dissection, and
instruments used in dissection, as well there was no correlation with the loss in
as to evaporation, for there was obvious weight.
drying of the surface of the protoplasmic Despite the limitations noted above,
parts during the few minutes required for consistent differences were found in the
this work. To minimize this loss the uptake of the several body parts of the
tared 10 ml beakers in which the body snails, when they were exposed to concen-
parts were placed were covered with trations of metal ions which would allow
aluminum foil until they could be weighed, normal behavior. The snails of all
and care was taken to complete the experiments recorded in Table 3 were
dissections and weighings as rapidly as exposed to 0.019 ppm zinc, and showed
possible. normal behavior at the end of the time
A discrepancy was also found in the of exposure. In the series in which each
total count obtained from the intact snail snail was exposed to 10 ml dosing solutio
and that determined by summation of the for varying periods of time (upper half
counts of the body parts after digestion. of Table 3), the body of the snail was
This difference averaged 42.5 ± 5.0% for only subdivided into 3 major parts. There
a sample of 114 snails. It was about was an evident decrease in the uptake of
equally divided between increase and the shell as time increased in that series.
TABLE 3. Average uptake of zinc 65 by the various body parts of Taphius glabratus, in counts
per minute per milligram of tissue, at varying exposure times and volumes of dosing
solution (concentration 0. 019 ppm)
Intact
snail
The cephalopedal mass, however, showed by the individual parts than in smaller
a steady increase with time of exposure. volumes of the same concentration. The
The irregularity in the trend of decrease shell showed an increase in uptake pro-
in uptake, found in the pulmonary complex, portional to the volume of dosing solution.
may be attributed to errors of counting The amounts taken up by the cephalopedal
and chance variation. Of special interest mass and pulmonary complex were again
is the large uptake of the upper visceral of the same general order of magnitude
complex as compared to other parts, and in each experiment, and theamounts taken
the fact that the values for the cephalopedal up by the stomach-albumen gland and gonad
mass and pulmonary complex are always were slightly higher than those found in
of thesame general magnitude in a given the cephalopedal mass and pulmonary
experiment. complex. The liver showed an uptake of
In the experiments in which the snails 4-7 times the amount present in other
were exposed to varying volumes of zinc protoplasmic body parts.
solution for 48 hours (lower half of Table Comparison of Tables 3 and 4 shows
3) the upper visceral complex was that the distribution of cadmium 115M
further subdivided (see Fig. 1). In larger within the snail is similar to that of zinc
volumes, larger quantities of zinc were 65 under the conditions of these experi-
usually taken up by the whole snail and ments. The data reported in Tables 1,3
TABLE 4. Average uptake of cadmium 115M by the various body parts of Taphius glabratus, in
counts/min/mg of tissue, at varying exposure times and volumes of dosing solution
(cone. 0. 045 ppm)
Intact
snail
346 YAGER AND HARRY
TABLE 5. Average uptake of cadmium 115M by the various body parts of />5 glabratus,
in counts /min/mg of tissue, during exposure to varying volumes of a dosing solution
of 0. 057 ppm Cd ions, of which only 91% were radioactive.
No. of
snails
used
UPTAKE OF RADIOACTIVE ZN, CD AND CU BY TA PHIUS 347
snail, although the concentration of zinc The Uptake of Copper

was a
little lower in this series of
experiments. This concentration, 0.0169 The short half life of Cu^^ (12 hours) and
ppm, was less than 1/5 of the concen- the fact that it is a beta emitter neces-
tration (0.10 ppm) of zinc previously sitated certain modifications of technique.
reported as producing distress in ^/5, Since the concentration of radioactive
when one snail was exposed to 40 ml copper used to provide suitable counting
of dosing solution. As in the case of levels amounted to only 0.001 ppm, and
the cadmium experiments, distressed we were interested in determining the
snails showed no more than about twice uptake in both the normal and distressed
as much metal in the liver as in other snail, the dosing solution was fortified
protoplasmic body parts, as against 4 by adding 0.030 ppm of non-radioactive
or more times as much in normal snails. copper in CUSO4. It was not possible,
Some normal snails had more metal in in this series, to duplicate the durations
all body parts than did the distressed of the zinc and cadmium experiments,
snails. but the time ranges applied roughly
TABLE 7. Average uptake of copper 64, in counts/min/mg of tissue, by the various body parts
of Taphius glabratus, in dosing solutions of 0. 031 ppm Cu ions of which 3. 2% (0. 001
ppm) only were radio-active, at varying volumes of solution and exposure times.
Total
body
parts
348 YAGER AND HARRY
correspond (Table 7). ments.
All snails exposed to 10 ml of dosing Shells from which the animals had been
solution showed normal, active behavior. removed by heat extraction, that had then
The liver again contained several times been filled with lead shot and whose
as much of the metal ions as did the aperture had been plugged with paraffin,
other body parts. All snails exposed to were found to take up 55.6 counts/min./
larger volumes of the same concentration mg Zn^S^ when placed in volumes of 10
showed distress, and their livers con- ml ofdosing solution of 0.019 ppm Zn,
tained, on the whole, smaller amounts at a pH of 7.0, for 48 hours. A parallel
than the other body parts. The livers series of experiments was done under
of distressed snails contained less copper conditions similar in all respects except
than did the livers of the normal snails, that the hydrogen ion concentration was
but analogous comparisons for the other varied (by increments of about pH 1.0)
body parts do not hold well in the data to test the uptake of Zn^S under acid and
of Table 7. alkaline conditions. KOH and HCl were
used to obtain the desired pH, but the
Incidental experiments with Zinc 65 solutions were not buffered.
The blood of a few of the normal snails
Satisfactory methods have not yet been used in the zinc experiments was examined
devised for determining the uptake of by drawing it up in weighed fragments
metal ions by the feces and mucus of capillary tubing, reweighing the filled
produced by the snails during exposure tube and counting it in the scintillation
to the dosing solution, despite numerous counter. The amount of zinc present in
attempts in which these materials were 4 such samples averaged 45.5 counts/
filtered and dried. Some preliminary min./mg. Each snail had been exposed
experiments indicated that about 40% of the to 10 ml dosing solution containing
of
radioactive zinc from a 0.019 ppm solution 0.0169 ppmzinc for 48 hours. This
may be adsorbed by the glass surface amount of zinc 65 was about half that
of the small Pyrex beakers used as present in the various protoplasmic parts,
experimental containers for exposing exclusive of the liver, of snails exposed
snails. Fresh lettuce, the only food given to the same conditions (see Table 6).
to the snails in the culture tanks, was However, the snails from which the blood
found to have an uptake, in light, of was taken had lower total body counts
342.5 counts/min./mg (average of 4 than did the snails used in the uptake
samples) of zinc 65 after 48 hours of experiments reported in that table. In
exposure (0.0169 ppm Zn, in 100 ml a few experiments on zinc uptake the
volumes, containing several grams of pulmonary complex was subdivided longi-
lettuce). Consequently no food was tudinally into 3 parts, one containing the
provided for the snails during the experi- heart and kidney, another the organs of
TABLE 8. Progressive release of zinc 65 from 4 normal Taphtusglabratus snails in 50 ml oi

water each after exposure for 48 hours to 10 ml of 0. 019 ppm Zn^S ion
distilled ,
solution.
Hours
the isthmian hemocoel and the third con- components in the uptake of these metal
taining the rectum (Fig. 1 C). The counts ions by the snail. Adsorbed metal may
of all 3 parts agreed very closely, showing be present on the shell, the cephalopedal
no difference in uptake among them. mass and that part of the pulmonary
The limited data obtained on the complex (the hypopeplar cavity and pulmo-
retention of zinc are presented in Table 8. nary cavity: see Fig. 1) which come
Four snails exposed individually to 10 ml into direct contact with the dosing solution.
of 0.019 ppm zinc for 48 hours were then From the present data it is not possible
placed in individual beakers each con- to differentiate the amount of metal
taining 50 ml of distilled water. Counts adsorbed and that absorbed by the
on the snails and water were taken after cephalopedal mass and pulmonary com-
17, 68 and 91 hours. The snails lost an plex. Probably both phenomena are
average of only 31.2% of the metal in operative concurrently in those 2 parts
91 hours. of the body.
Evidence for the adsorption of metals
DISCUSSION by the shell was furnished in the experi-
ment in which the empty shells were
was not feasible to determine the
It found to take up zinc. The uptake was
amounts of heavy metals present in the found to be a function of the pH of the
snails before they were subjected to dosing solution. That the shells of snails,
experimental doses of the same. Whether which were placed in 10 ml only of dosing
such intrinsic metal, or some metabolic solution, took up less metal as time
analogue of it, present perhaps in varying increased (very clear in upper part of
amounts, may have been a factor in the Table 3; see also Tables 4 and 7), while
variable uptake observed in the experi- those placed in increasing amounts of
mental lots can not be determined at dosing solution of the same concentration,
present. However, it is well known that for a constant period of time, took up
snails are inconsistent in their behavior, larger amounts (lower part of same
moving about and feeding very irregularly. tables), may also be taken as evidence
It has been regularly observed in our for adsorption. Perhaps the initial concen-
laboratory cultures that snails of the tration of the metal was rapidly and
same post hatching age, maintained under significantly reduced in the former case,
uniform conditions, show much variation but remained relatively constant in the
in size. It is therefore reasonable to latter, for reasons to be discussed below.
assume that the irregular uptake is also If this be so, then the adsorptive capacity
a function of their irregular behavior, and of the shell is a function of the concen-
that a snail, while crawling,may incorpo- tration of metal ions in the solution bathing
rate more of these metals than during the shell.
periods when it is quiescent, retracted to Absorption may take place by one or
varying degrees, or has crawled above the more of the following routes, of which
water line (as occasionally happened). the first seems to be most probable: (1)
Whatever the various causes, the amount through the epithelium of the cephalopedal
of uptake of the whole snail also. seems mass and pulmonary complex; (2) through
to depend, in part, on some intrinsic the wall of the digestive tract after the
property of the snail. The differences fluid enters through the mouth; (3) through
obtained in counts per minute of the the shell, or between the shell and mantle
whole snail and the summation of counts epithelium, and thence through the latter.
per body part of the digested parts may cases the liver of normal snails
In all
be attributed to the geometry of counting showed a marked propensity for
as well as to the material lost in dis- accumulating the metal ions and that of
section. distressed snails one much less pro-
Both adsorption and absorption may be nounced, less in the cadmium than in the
350 YAGER AND HARRY
zinc series and the reverse in the copper We chose a dosage of 0.031 ppm copper
series. No regular pattern of differential because it lies slightly below the minimal
concentration in other tissues was found. concentration previously determined to
That the amounts were slightly higher in produce distress when one snail was
the stomach-albumen gland and gonad exposed to 40 ml of dosing solution (Harry
preparations than in the cephalopedal mass and Aldrich, 1963). This choice was
and pulmonary complex may be due to fortunate, and again distress was
bits of liver tissue adhering to the former differentially produced, according to the
2 parts, for the soft consistency of the volume of dosing solution to which the
liver renders it difficult to remove that were exposed.
snails
organ intact.Mantle epithelium, con- The influence of the volume of the
nective tissue and blood were present in dosing solution may be merely incidental
all 3 of the subdivisions of the upper in producing distress. Possibly there
visceral complex. It is unlikely that the is a more rapid decrease in the concen-
upper loop of the intestine or the herma- tration of metal ions in the smaller
phroditic duct, both present in the samples volumes of dosing solution than in the
designated "liver" in the several tables, larger.
were responsible for the high concen- In theory, the amount of ions adsorbed
tration of metal in that section. on the beaker's surface would be pro-
The stunted snails used in the initial portionally less in the larger than in the
zinc experiments (Table 6) were not smaller volumes of the dosing solution.
distressed in small volumes of dosing The adsorption by feces and mucus would
solution, but were distressed in larger be approximately constant in both cases.
volumes of the same concentration. The The absorption by the snail would also
distressed condition as determined by remove ions from the medium. Since the
inspection was substantiated by the pattern larger volumes (50 ml or more) provided
of differential uptake of the zinc by the 5 or more times as much metal as the
livers of distressed and normal snails. smaller volumes (10 ml), the concen-
Our inability to repeat these results with tration of ions in the fluid would remain
another lot of unstunted snails and relatively more constant throughout the
approximately the same concentration of exposure period in the larger volumes.
zinc (Table 3) may be due to the difference Perhaps in the small volume tests,
in their size, those used in the latter sufficient ions were removed from the
experiments being twice as large as those medium soon enough to reduce the concen-
in the former. Similarly the snails, all tration to a level beloAv that which would
large, dosed with cadmium, were not produce distress, and in the larger
distressed when exposed to a range of volumes the concentration of ions was
volumes of dosing solution with a low not reduced below that threshold. Un-
concentration (Table 4). Presumably, a fortunately we have no counts on the test
slight increase in the concentration of solutions at the end of exposure time.
zinc, or cadmium, would produce distress However, the readings for the shell,
in the larger snails, differentially, i.e., particularly the decrease in count with
produce distress in the larger but not the increase of time in small volume tests
smaller volumes of dosing solution. Un- (and uniform initial concentration), give
fortunately, time did not allow us to test indirect support to this view. If this be
this hypothesis. The increased concen- so, we may tentatively conclude that the
tration of cadmium chosen (0.057 against concentration of the metal in the medium
0,045) was evidently beyond such a (and the length of time the snail is exposed
threshold for snails of that size (Table 5), to it) is the important factor in inducing
since it already produced distress in distress, rather than any absolute amount
relatively small volumes of dosing so- which may gain entry into the snail, and
lution. indeed it has been shown that in many
instances normal, undistressed snails {I.e.,Fig. 7, p 858). In the case of

contained more metal than the distressed sodium, strontium and rubidium,
snails, both in the liver and in other nearly all parts of the animal showed
tissues. an amount of radioactivity so large as
Regarding metabolic mechanism
the to allow little differentiation of structure
whereby these produce distress,
ions in the autoradiographs. However, there
one might initially have assumed that the is a suggestion that the gonad took up
liver is directly involved, since it shows less sodium and rubidium than did the
such a propensity for taking up the metal other body parts, for the apical tip of the
ions. This, however, does not explain protoplasmic mass is somewhat darker
the fact that normal snails often have than the rest of the body (compare with
several times more metal in their livers our Fig. 1). In the figure representing
than the distressed snails (assuming the the uptake of iron, there is more
initial amounts metals were the
of the differentiation, and we would suggest that
same). A more likely hypothesis would the liver and part of the pulmonary cavity
be that the permeability of the surface took up less iron than the rest of the
membranes through which the metal enters snail. The gonad (and seminal vesicle),
the snail is somehow impaired, if a the stomach-albumen gland, some organs
sufficient concentration of the effective of the ismian hemocoel and perhaps the
ion is maintained for a sufficient period of kidney all seemed to concentrate this
time. The impairment of permeability material.
is evidently not complete, for some ions
seem to get into the snail even in the BIBLIOGRAPHY
distressed state. Perhaps the impairment
of membrane permeability is none the HARRY, H. W.
and D. V. ALDRICH,
less sufficient to block the passage of 1963. The
distress syndrome in
materials which need to pass if the snail Taphius glabratus (Say) as a reaction
is to function normally (e.g., oxygen or to toxic concentrations of inorganic
waste products) and distress ensues. ions. Malacologia, 1(2): 283-289.
Taphius glabratus seems to react to the JOHNSON, C. R., R. ANGEL and D. G.
3 metals used in the present study in a ERICKSON, 1962. The uptake, dis-
very similar manner. However, Johnson tribution and excretion of four radio-
et al. (1962) have presented evidence nuclides in Australorbis glabratus
indicating that the uptake pattern found (Planorbidae). Amer. J. Trop. Med.
by us may not apply in the case of other andHyg. 11:855-860.
cations. They published autoradiographs MALEK, E. A., 1962. Laboratory guide
of the protoplasmic body removed from and notes for medical malacology.
the shell by heat extraction, after exposure Burgess Press, Minneapolis. 154 p.
to iron, sodium, strontium and rubidium
ZUSAMMENFASSUNG
AUFNAHME VON RADIOAKTIVEM ZINK, KADMIUM UND KUPFER BEI DER
SÜSSWASSERSCHNECKE TAPHIUS GLABRATUS (SAY)
Im Labor gezüchtete Taphius glabratus Ç\ustralorbts und Planorbirm anderer Autoren)

wurden 2 Konzentrationen von Zn^^, Cd H^M oder Cu^^ ausgesetzt, deren schwächere
noch normales Verhalten zuliessen, deren stärkere dagegen bereits einen ''Notzustand"
hervorriefen: die Schnecken waren nicht mehr fähig sich in ihre Schale zurückzuziehen.
Sie wurden in Gruppen von 6-10 in jeweiligen Dosierungslösungen von 10-200 ml
Volumen für 6-12 Stunden belassen. In jeder der Versuchsgruppen nahmen die
Schnecken die Metall-Ionen sehr verschiedentlich auf, was wahrscheinlich auf iiinen
ganz individuell innewohnenden Eigenschaften beruhen dürfte, die möglicherweise auch
mit der von ihnen entwickelten Aktivität in den metallischen Lösungen in Zusammenhang
352 YAGER AND HARRY
stehen. Um die Metallanreicherung nach einzelnen Körperteilen zu ermitteln, wurden
Schneckenpaare gewählt, bei denen die Radioaktivität des Gesamtkörpers dem Mittelwert
am nächsten lag. Nach Entfernung der Schale wurden die protoplasmischen Körperteile
in bestimmte Abschnitte zerlegt, gewogen, mit konzentrierter Salpetersäure versetzt
und daraufhin mit der entsprechenden Apparatur auf ihre Radioaktivität hin geprüft.
Die Messungen ergaben einen durchschnittlichen Unterschied von 42.5% zwischen den
Zählungen pro Minute pro Milligram Gewicht, für die lonversehrte lebendige Schnecke,
und der Summe der Zählungen der verschiedenen Körperteilenach Digerierung. Dieser
Unterschied war gleicherweise auf Gewinn und Verlust aufgeteilt und zeigte keinerlei
Zusammenhang mit dem Gewichtsverlust nach der Sektion, der im Mittel 18.1% betrug;
er dUrfte daher dem Untersuchungsverfahren zuzuschreiben sein.
Das Aufnahmebild für die 3 Metalle war im wesentlichen das Gleiche. "Normale"
Schnecken, die in grössere Volumen der radioaktiven Lösungen gebracht worden waren
nahmen mehr Metall auf als, bei gleicher Konzentration und gleicher Zeitdauer, solche
in nur 10 ml Lösung. In den Versuchen mit nur 10 ml Lösung pro Schnecke verringerte
sich, mit zunehmender Zeitdauer, die Aufnahme durch die Schale, gemessen in Zählungen
/Min/mg. In den jeweiligen verschiedenen Versuchen war die Metallanreicherung aller
Weichteile von derselben Grössenordnung, ausser in der Leber, in der sie 4-7 mal so
hoch war. Die Leber von Schnecken in "Nöten" enthielt hingegen nur 2 mal so viel
Metall als die übrigen Gewebe. Gewebe von Schnecken, die sich durchwegs normal bewegt
hatten, enthielten oft grössere Metallmengen als solche die schon toxische Symptome
zeigten. Wir folgern daraus, dass der Notzustand eher von der Konzentration der Ionen
in der Flüssigkeit abhängt als von der absorbierten lonenmenge und dass die
Wirksamkeit der Ionen eher auf Zerstörung der Membranpermeabilität beruht als auf
Störung eines internen metabolischen Mechanismus.
RESUME
ABSORPTION DE ZINC. CADMIUM ET CUIVRE RADIOACTIFS
PAR LE PULMONÉ TAPUIUS GLABRATUS (SAY)
Des Taphius glabratus (ûstralorbis et Planorbina des divers auteurs) d'élevage

furent soumis à des solutions de Zn°^, Cd ^^^^ et Cu""^ à taux suffisamment falibles
pour permettre encore un comportement normal, ainsi qu'à des taux suffisamment
élevés pour provoquer déjà un état de détresse (incapacité de se rétracter dans la
coquille). Les mollusques furent traités en groupes de 6 à 10 individus, par des
solutions dosées de volumes allant de 10 à 200 ml par mollusque, pour une durée de 6 à
72 heures. Dans chaque essai, l'absorption des ions métalliques par les divers individus
se montra fort variable, probablement en raison de quelque propriété intrinsèque du
mollusque, dépendant peut-être de son activité dans la solution radioactive. Pour
étudier l'absorption selon les diverses parties du corps, nous avons choisi des paires
de mollusques dont les "comptes" du corps intégral étaient les plus proches de la
moyenne. Après avoir ôté la coquille et divisé le corps en sections définies, celles-
ci furent pesées, digérées avec de l'acide nitrique concentré et ensuite mesurées quant
à leur radioactivité au moyen d'appareils appropriés. Une différence moyenne de
42.5% fut constatée entre "comptes" par minute par milligrammes du mollusque vivant
intact et la somme des "comptes" pour les diverses parties du corps après digestion.
Cette différence est attribuée à des différences de procédure car elle était distribuée de
façon égale entre gains et pertes et ne montrait aucune relation avec les pertes de
poids après dissection, qui étaient en moyenne de 18.1%. Les régimes d'absorption
étaient essentiellement les mêmes pour les 3 métaux. Pour les mollusques normaux.
à concentration de solution et durée de traitement égales, une quantité plus grande
d'ions métalliques fut absorbée par les mollusques en volumes plus grands de solution.
que par ceux se trouvant dans 10 ml de solution par mollusque seulement. Aussi, dans
les séries à volumes de 10 ml. l'absorption par la coquille, mesurée en comptes
min/mg. diminuait avec l'accroissement de la durée de traitement. Dans un essai donné,
l'abjorption était du même ordre de grandeur pour toutes les parties protoplasnuques
du corps, sauf le foie, où elle était de 4 à 7 fois supérieure. Quant aux mollusques
en détresse, leur foie ne contenait qu'une quantité double de métal. Des mollusques
ayant montré une activité normale, avaient souvent plus de métal dans leurs tissus
protoplasmiques que ceux en détresse. Nous déduisons donc que l'état de détresse
dépend de la concentration des ions en solution plutôt que de la quantité d'ions absorbée
et que leur action consiste à détruire la perméabilité des membranes plutôt que de se
rapporter à quelque mécanisme métabolique.
RESUMEN
LA ABSORCIÓN DE CADMIO Y COBRE RADIOACTIVOS POR EL CARACOL DE
ZINC,
AGUA DULCE TAPHIUS GLABRATUS (SAY)
Taphius glabratus (/íustralorbis o Planorbina de los autores) criados en laboratorio
fueron expuestos a concentraciones de zinc 65, cadnúoll5M o cobre 64 que no cambiaron
el comportamiento normal, y a otras concentraciones de los mismos metales que
produjeron la reacción depresiva (incapacidad para retraerse en la concha). Los
caracoles fueron expuestos, en grupos de 6- 10, por períodos de 6-72 horas, a volúmenes
de las soluciones variando entre 10-200 mi por individuo. Se comprobó amplia variación
en la absorción de los iones metálicos por los caracoles expuestos en cada experimento,
lo que se atribuye a alguna propiedad intrínseca de los animales, quizá dependiente
de la actividad de cada individuo durante su permanencia en la solución. Se seleccion-
aron parejas de caracoles, cuyos cuerpos completos proporcionaron recuentos los
más cercanos a la media, para análisis de la absorción por varias paurtes del cuerpo.
Después de quitar la concha y dividir el cuerpo en secciones definidas, estas fueron
pesadas y digeridas en ácido nítrico concentrado y luego se hizo el recuento de radio-
actividad. Una diferencia media de 42,5% entre los recuentos por min/mg del animal
entero y vivo y la suma de los recuentos de las partes después de la digestión fué
atribuida a la diferencia en la geometría del recuento. La diferencia se dividió más
o menos igualmente entre pérdida y ganancia después de la disección y no mostró
correlación con el peso perdido en la disección, que fué en media 18,1%. Los padrones
de absorción fueron esencialmente los mismos para los 3 metales. En caracoles
normales, más iones metálicos fueron tomados por aquellos expuestos a mayor volumen
de solución que por aquellos expuestos solamente a 10 mi de solución de la misma con-
centración por iguales períodos. La absorción por la concha, medida por min/mg,
decreció a medida que aumentó el tiempo en experimentos en que los caracoles fueron
expuestos a 10 mi de solución por individuo. En cada tipo de experimento la magnitud
de la absorción fué la misma para todo el cuerpo, con excepción del hígado, el cual
reveló 4-7 veces las cantidades presentes en otros tejidos. Caracoles que mostraban
actividad normal, con frecuencia tenian mucho mayores concentraciones de metal en
los tejidos del cuerpo que aquellos con reacción depresiva. Se concluyó que el síndrome
depresivo depende antes de la concentración de un ion en la solución, que de la cantidad
de un ion absorbido, y que la eficacia de los iones en la producción del síndrome
depresivo resulta de su capacidad para alterar la permeabilidad de las membranas
más bien que de su interferencia en algún mecanismo metabólico interno.
THE ANATOMY OF CHILINA FLUCTUOSA GRAY

REEXAMINED, WITH PROLEGOMENA ON THE PHYLOGENY OF THE HIGHER
LIMNIC BASOMMATOPHORA (GASTROPODA: PULMONATA)
Harold W. Harry
Biology Department
Rice University
Houston 1, Texas, U. S. A.
ABSTRACT
A reexamination of the gross anatomy of this species revealed several charac-
ters not previously noted in the Chilinidae, such as the post-tentacular lappet, the
presence of sand grains in the stomach, a spermatophore the adnate nature of
,
the colimiellar retractor muscle, the tesselated integument of the cephalopedal

mass, calcareous connective tissue granules and calcareous granules in the
lumen of the vagina.
A comparison is made of the anatomy oiChilina with that of the higher limnlc
Basommatophora, and incidentally with other major piilmonate groups. This
comparison is followed by a preliminary outline of the phylogeny of Chilina and of
the higher limnic Basommatophora, in which each stage in their evolution is de-
fined. The outline starts arbitrarily at the "Primitive Pulmonate" level, then
suggests which characters were added,, modified or lost in relation to the time
when other characters underwent similar changes. Multiple features are found
to characterize each of several hypothetical stages, each of which has given rise
to one or more families which survive. These hypothetical stages are named in
a manner which has no nomenclatorial validity.
The scheme shows evolution as an uninterrupted continum. It is not intended
to be an exhaustive definition of taxa, although some families are extensively
defined. The constriction of the mantle cavity to form an apical pulmonary
cavity and an apertural hJopeplar cavity, interconnected by a pneumostome, is
considered one of the most fundamental characteristics of the pulmonates. This
mechanism further evolved along one of 2 different lines: (1) one with a con-
tractile pneumostome, in which the pulmonary cavity is used exclusively for
aereal respiration; (2) one with a noncontractile penumostome, in which the
pulmonary cavity is used either exclusively for holding water, or water and air,
but never exclusively for air. Only the second type occurs in Chilina and the
limnlc Basommatophora. It has been modified by the addition of a pulmonary
lamella, and several pneumostomal appendages. It is suggested that the pseudo-
branch and siphon of the higher limnic Basommatophora are always distinct
entities, never fused, and not homologous to each other. But the homology of
siphon and pseudobranch to the pnexmiostomal lappet of the lower pulmonates
is an undecided question. The loss and modification of these in the higher limnlc
Basommatophora is discussed.
Other characters considered in the phylogeny are heterostrophy hyperstrophy
,
and inverted symmetry; the reversion to a patellate form; the movement of the
cerebral nerve ring posterior to the buccal mass, and constriction of the visceral
and cerebral nerve rings position of the eye and shape of the tentacles modi-
;
;
fication of the type of calcareous granules of the connective tissue; presence of

hemoglobin; apical punctation of the shell; shell dentition; distribution of the
intestinal loops; presence of a gastric cecum; number of major lobes of the
liver; several radvdar characters; disappearance of the external reproductive
groove; extent of separation of the male and female reproductive tiibes below the
(355)
356 H. W. HARRY
albumen gland; loss of the vesícula seminalis vaginae; type of gonadal atrium
and the attachment of the hermaphroditic duct in relation to it and the topo-
,
graphical relation of the gonad and upper liver lobe.
INTRODUCTION Baker (1925); that on Acroloxus from

Sharp (1883), André (1893), and Hubendick
the importance accorded to
Despite (1962); that on Latia and Amphibola is
Chilina the numerous outlines of
in from Pelseneer (1906). The several
phylogeny proposed for the pulmonate papers by Hubendick on Bulinus, the
gastropods, no new data on the anatomy Planorbidae, Lymnaeidae, Patelliformia
of these snails have appeared since the (Siphonariidae and Gadiniidae) and other
fundamental study of Haeckel(1911), other groups, and the extensive bibliographies
than a few observations recorded by which he has cited have provided a wealth
Hubendick (1945). My unpublished studies of information. The monumental but
on the anatomies of the higher limnic neglected compendium of the literature on
Basommatophora suggested that a number pulmonate snails provided by Simroth and
of anatomical features of systematic Hoffmann (1908-1928) has been of special
importance in this group have been ignored value. For convenience, this work is
or little emphasized. Reference to the often cited below to substantiate a point,
literature did not satisfy requirements rather than citing the dozens of original
for comparative data on Chilina, and a papers from which their work was drawn.
reexamination of the anatomy of Chilina I am indebted to Dr. Walter Biese for
fluctuosa showed several new points. This the several specimens of Chilina
species was one of the 4 studied by fluctuosa, and to Dr. Clark P. Read for
Haeckel. making it possible to complete this work,
In the present paper, the term "higher which was in part financed by National
limnic Basommatophora" is meant to Science Foundation grants G17943 and
include the Planorbidae, Bulinidae, ^ GB-820.
Ancylidae, Physidae, Lymnaeidae (in-
cluding Lanx) and Acroloxidae, but it ANATOMY OF CHILINA
excludes the Chilinidae and Latiidae. I
have had the opportunity of making The 10 specimens of Chilina fluctuosa
personal observations on the anatomies examined were from Chaira, Sano
of all known species of the EUobiidae Relourami, S. Puerto Monatt, Chile. They
from Florida, and a few species from the were all very contracted, having been
Pacific area; on the Carychiidae from dropped directly into alcohol when col-
Michigan; on the freshwater pulmonates of lected by Dr. Biese, in 1958. He informed
Puerto Rico and other Caribbean islands, me that they came from brackish water.
and from several places in North America
SheU (Fig. 1).
{Parapholyx, courtesy of Dr. Dwight
Taylor), on living and preserved Bulinus Dextral, ovate, imperforate, the
sp. from the Sudan (courtesy of Drs. apertural height about 3/4 the height of
Henry van der Schalie and J. B. Burch); the shell. Whorls of the spire evenly
as well as on Siphonaria sp., in the living and prominently convex. Suture moder-
state, from Texas. Reference to the ately indented. Body whorl rounded near
published literature on these groups is the suture and at the base, but the pe-
made where relevant. riphery is much less convex. Texture of
The information on Gadinia is derived the later shell is smooth but not polished,
from Schumann (1911); that on Lanx from with vague, flattened growth lines of
For reasons of separation of Bulinidae from Planorbidae, see p 380-381. ED,

ANATOMY OF CHILINA AND BASOMMATOPHORAN PHYLOGENY 357
^3./ PC
M s
FIG. 1. Shell of Chilina fluctuosa Gray.
FIG. 2. Abapertural view of animal removed from shell.
FIG. 3. Anterior view of animal removed from shell, with pvilmonary roof cut away (lined area).
LIST OF ABBREVIATIONS
A - Anus PARG - Parietal ganglion
AG - Albumen gland PC - Pulmonary cecum
BUG - Buccal ganglion PDG - Pedal ganglion
BUR - Buccal retractor of columellar muscle PE - Penis
BV - Blood vein PLG - Pleural ganglion
CD - Collecting duct of gonad PN - Pneumostome
CERC - Cerebral commissure PR - Prostate
CEFG - Cerebral ganglion PREP - Preputium
CS - Cardiac end of Stomach PRM - Protractor muscle
DPCL - Dorsal part of pulmonary cavity PS - Penis sheath
lamella PSB - Pneumostomal appendage
DSG - Duct of salivary gland PTL - Post-tentacular lappet
E - Eye PYL - Pyloric end of stomach
ERG - External reproductive groove R - Rectum
ES - Esophagus RAD - Radular sac
FG - Female reproductive groove RM - Retractor muscle
FRO - Female reproductive opening RP - Renopore
FT - Lateral area of foot RPCL - Rectal part of pulmonary cavity
GC - Gastric cecum lamella
HT - Heart SEMV - Seminal vesicle
ING - Innominate ganglion SG - Salivary gland
- Kidney SPOD - Spermoviduct
KM - Muscle band of kidney SPTH - Spermatheca
L - Liver STA - Statocyst
LLI - Lower loop of intestine T - Tentacle
LO - Opening of the liver into the intestine TA - Talon
LP - Labial palp UT - Uterus
MG - Male groove of spermoviduct VA - Vagina
MRO -
Male reproductive opening VD - Vas deferens
MS - Muscular stomach VSV - Vesicula seminalis vaginae
OS - Osphradium
358 H. W. HARRY
irregular prominence and spacing. No veloped sub-sutural, linear indentation.
spiral sculpturing. The internal partitions
External features of the animal removed
are not absorbed. Aperture ovate, lip
from the shell (Figs. 2, 3).
sharp and thin, not reflected. No teeth
in any part of the aperture. A thin, The bodies were much retracted, but
white callus on the inner surface of the not inverted. The animal is also dextral
outer lip, sometimes slightly elevated with anus, pneumostome and repro-
near the margin, to simulate a labial ductive openings on the right, and the
callus. Parietal area with a thin, white heart on the left side. Foot short, with
callus which continues about half way the a broad, oval sole gently rounded before
length of the columellar area in the and behind. The sole had on it numerous
umbilical region. Columellar area longitudinal, deep, discontinuous grooves
flattened, elongate, very narrow, slightly which appear to be preservation phe-
arched on its inner margin. Its upper nomena; no transverse or longitudinal
end continues as a projecting columello- grooves were present which might be
parietal lamella, twisting about the present in the living animal. According
columella and disappearing into the shell. to Haeckel (1911) the pedal gland cells
The apices of all larger shells were badly are diffusely scattered over the surface
eroded, and large erosion spots were of the sole, and there is no single gland
present on the spire and body whorl. with well defined duct opening at the
Color dark brown, with 4 spiral, maroon anterior end of the foot. No such gland
bands, of moderate width, equally spaced was present in the material which I
between the suture and the base. These examined, either.
maroon bands are interrupted at the more The integument of the side of the foot
prominent growth striae, and they are has large, round and oval bosses, forming
sometimes connected by thin, transverse an irregularly tesselated pattern. There
stripes of maroon between the 4 major are no supra-marginal grooves, nor
bands. In smaller shells the transverse median, dorsal, posterior groove nor
stripes predominate, and they may be posterior pedal gland. The integument
irregular and undulating, or sharply angled is dark brown. Numerous small orange
and chevron-shaped. A typical shell
measured: height, 18.3 mm; apertural
height, 13.7 mm; greater diameter, 10.3
mm; suture whorls, 3 1/2 (apex eroded).
One juvenile shell with apex intact is
present;
apertural
it measures:
height,
height, 4.7 mm;
3.6 mm; greater
diameter, 3.1 mm; it has 2 suture whorls
beyond the half whorl forming the hetero-
strophic apex. The aperture is as de-
scribed for the adult shell, including the
columellar callus and the columello-
parietal lamella. The maroon markings
are transverse, very irregular in width
^
and spacing, and undulating. The embry-
onic half whorl of the shell is distinctly
FIG. 4. Radula, to show general shape and
at a right angle to the first whorl of
shape of transverse rows of teeth
later shell growth, and the apex is there-
(only a few shown).
fore heterostrophic (see Harry, 1951).
The embryonic whorl is smooth, without FIG. 5. Jaw.
sculpture. The shoulder of the body whorl FIG. 6. Radular teeth; centrals and first few
is slightly flattened, with a vaguely de- laterals of each side.
flecks, clumped to form small patches of mantle cavity that is outside the
the
on the head, persisted even after 5 years pulmonary cavity) is deep and large, and
of preservation. Small white flecks were not occluded by an extensive thickening of
also present. the mantle margin. The mantle margin
The prominent labial palps (Fig. 3) are is smooth, without papillae, and apparently
rectangular. The lower margins are not does not extend beyond the margin of the
covered by a cuticle, and apparently they shell's aperture.
have no sensory pads. The labial palp on The pneumostome (Figs. 7 and 8) is a
the right side is separated from the head simple elongate slit, extending along the
by a groove which extends all the way shell lip from the apertural angle of the
up to the male reproductive pore, and the shell to a point about half the length of the
palp on the left has a separating groove outer lip. On its medial margin there
of comparable extent dorsally. The attaches a large, swollen, semilunar flap,
tentacles are very small, semilunar pro- the pneumostomal appendage. On it lies
jections from the head, and broader than the rectum which opens near the outer,
long. The eyes are situated at the outer or free, margin. There are no transverse
side of the base of the tentacles. Behind lamellae on this structure. Nothing
the eye is a second flap as large as the comparable to a true siphon (see below)
tentacle, which seems to be homologous was found in any of the specimens ex-
to the sensory flap at the base of the amined.
tentacle in the higher limnic Basommato-
The pulmonary complex (Figs. 2, 7 and 8).
phora.
On the right side of the body there is The external epithelium of the pulmo-
a genital groove leading from the female nary cavity uniformly dark black,
is
pore on the body stalk to the posterior occasionally with minute oval or round
end of the right post-tentacular flap, spots left unpigmented near the mantle
where the male reproductive pore opens. margin. The black epithelial layer extends
The groove is shallow and narrow, but to the apex of the visceral mass, but only
very distinct. in the shoulder area of the shell. This
The hypopeplar cavity (i.e., that part layer is very loosely attached to the
RPCL /DPCL
FIG. 7. Pneumostomal corner of pulmonary cavity, opened to show pulmonary cecum.

FIG. 8. Pneumostomal corner of pulmonary cavity, with pneumostomal appendage in natural
position.
360 H. W. HARRY
underlying musculo-connective tissue is filled with numerous transverse
layer of the body wall, and sloughs off trabeculae which extend from the roof
readily. to the floor of the kidney and anastomose
The pulmonary cavity extends inward to some extent. They are discontinuous
from the shell's aperture about 1/4 whorl, transversely in an irregular fashion, so
and probably farther in living specimens. that the movement of the kidney contents
At the base of the shell there is a very from the upper to the lower end is through
slight horn extending apically beyond the a labyrinthine passage, with no tube-
general area of the cavity, and this shaped central lumen. There is no sub-
contains the pericardium. Paralleling renal fold of tissue projecting into the
the suture of the shell there is a narrow, pulmonary cavity, but a prominent, narrow
elongate horn of the pulmonary cavity band of muscle tissue is present on the
which extends farther apically about 1/4 free surface, near the left margin of the
whorl. This structure was designated kidney.
as the pulmonary cecum byHaeckel(1911: A small blood sinus courses along the
100). The apex of the pulmonary cecum left margin of the kidney, entering the
is well above (apical to) the region of auricle of the heart. Similar blood
the stomach. sinuses (or vessels, though they are
The rectum enters the pulmonary nothing more than recessions in the
cavity slightly below the apical end of connective tissue of the pulmonary cavity
the pulmonary cecum, and takes its roof) take their course along the other 2
course to the pneumostome in a large margins of the triangular kidney, and
rectal wedge which bulges into the cavity. there have their origin from a single
On the rectal wedge is a broad, flat, tubular sinus which runs down the roof
opaque, white strip of tissue which appears of the pulmonary cavity cecum from the
glandular. Haeckel (1911) found it to be hemocoel of th« upper visceral complex.
ciliated. He described and figured this An osphradium is situated on the roof
strip, which is here termed the rectal of the pulmonary cavity at the left end
part of the pulmonary cavity lamella, as of the pneumostome, near the renal pore.
beginning near the tip of the "pseudo- It is a fleshy ridge, about 3 times as
branch" (= pneumostomal appendage). I long as broad, projecting into the pulmo-
was not able to confirm its presence nary cavity. On its free surface is a
beyond the edge of the pneumostome. It deep, narrow groove, about 3/4 as long
courses apically into the pulmonary as the organ itself. In the wall of the
cecum, and at the tip of the latter it is partition between the pulmonary cavity
reflected on the roof of the pulmonary and the hypopeplar cavity, at the left
cavity (the dorsal part of the pulmonary end of the penumostome and near the
cavity lamella. Figs, 7 and 8), where it osphradium, is the right parietal ganglion
runs parallel to the rectal part all the (Fig, 8), which sends a large nerve to
way back to the pneumostome. the osphradium.
The kidney is a large, triangular,
The upper visceral complex (i.e., the
flattened sac, which nearly fills the outer
part of the visceral mass apical to the
wall of the pulmonary cavity. The basal
pulmonary cavity).
corner abuts against the pericardium,
and there is a large renopericardial The upper visceral complex extends
opening at this place. The renal pore is through about 2 whorls. Its lower
at the apertural angle, near the pneumo- (aperturad) end is almost completely
stome (Figs. 7 and 8), This pore has a occluded by the albumen gland and
thickened margin. No true ureter is spermoviduct, which leaves merely a
present, the "ureter" described by small passage on the basal (left) side for
Haeckel being only the apertural end of the passage of the short esophagus. The
the kidney proper. Internally the kidney stomach is toward the base, and just above
the albumen gland- spermovi duct complex. forming the upper (apicad) loop of the
Apical to the stomach, the upper visceral intestine, and continues aperturad near
complex is filled by the liver, with the the suture of the shell. It enters the
upper loop of the intestine, the gonad and pulmonary cavity by way of the pulmonary
hermaphroditic duct imbedded in it. cecum, to become the rectum.
The digestive system of the upper visceral The reproductive system in the upper
complex. visceral complex (Fig. 11),
Near its upper end the esophagus The gonad consist of several bunches
expands somewhat to form the cardiac of rounded follicles which open
short,
end of the stomach. This part lacks into branched ducts.
small, These in
grossly evident circular muscles (Fig. 9). turn open into a major collecting duct,
The middle portion, or muscular stomach, along which they are distributed at widely
has a prominent sheet of circular muscle spaced intervals. The major collecting
tissue, which does not form a complete duct is thin, membranous and of uniform
sphincter around the tract, but is broken diameter throughout its length. The gonad
into 2 crescent-shaped parts, so that the follicles and collecting ducts are imbedded
muscular stomach appear bilobed. Apical in the surface of the liver, on the
to the muscular stomach is another thin- columello-parietal aspect of the visceral
walled part, the pyloric stomach, the di- mass, extending from the apex aperturad
ameter of which again narrows as it for about one whorl.
approaches the intestine. The interior The hermaphroditic duct emerges from
of the stomach of all specimens was the common collecting duct of the gonad
filled with sand grains. The intestine at a point distant about 1/4 of the gonad's
curves first toward the left; at its length from its lower end. It continues
junction with the pyloric stomach, it between the surface of the liver and the
receives the single opening of the liver mantle to the lower end of the upper
from above and, just opposite, the opening visceral complex, where it enters the
of the short gastric cecum. The liver "talon" (see below). Throughout most of
is limited the region apical to the
to the distance from the gonad to the "talon",
stomach. lobed and grossly similar
It is the hermaphroditic duct is swollen, con-
to that of the higher limnic Basommato- voluted, with numerous digitiform out-
phora, but since it opens into the intesti- pocketings. This modification of the
nal tract in only one place, it has only hermaphroditic duct constitutes the
one major lobe. The subdivisions of this seminal vesicle.
lobe open into a large, poorly defined
The lower visceral complex.
central lumen. The gastric cecum also
resembles that of the higher limnic It includes all of the cephalopedal mass
Basommatophora in form and position. and the organs in that partof thehemocoel
After running aperturad, close to the which subtends the pulmonary cavity (the
left side of the stomach, the intestine isthmian hemocoel). In Chilina the
turns to the right, crossing the lower end isthmian hemocoel is short and wide and
of the upper visceral complex along the thus poorly differentiated in comparison
apical margin of the albumen gland and to its shape in snails whose visceral
peripherally to the esophagus. This mass occupies more whorls.
portion is the lower (aperturad) loop of The body wall of the cephalopedal mass
the intestine. It then passes again apicad is thick and muscular, including the dorsal
along the right side of the stomach. Here region of the head. The columellar
it becomes imbedded on the surface of retractor muscle is short and broad,
the liver, and courses about a whorl originating about 1/4 of a whorl apically
above the stomach, swinging again toward of the shell's aperture, at the upper extent
the left in this passage. At its upper of the isthmian hemocoel. This muscle
extent it again swings to the right, thus is not divided longitudinally, but it does
362 H. W. HARRY
BUG
PYL-
RAD PRM
BUR
CERC
SEMV
PDG
CERG
PLG
PARG
ING
FIG. 9. Anterior part of the digestive tract. Dorsal aspect.
FIG. 10. Lateral view of buccal mass.
FIG. 11. Gonad and upper part of hermaphroditic duct, with initial portion of seminal vesicle.
FIG. 12. Upper 3 groups: calcareous granules from connective tissue; lower 2 gi'oups: calcareous
gi'anules from lumen of tne vagina. Scale applies to both groups and is 0. 03
, lon^. mm
FIG. 13. Central nervous system. Statocysts on pedal ganglia are not labled.
not extend beyond the columello-parietal of uniform diameter throughout its length.
plait of the shell toward its apertural Internally it has numerous low, longi-
angle. Underlying the rectal wedge, tudinal rugae which do not anastomose.
which actually lies on the parietal wall The salivary glands have short ducts.
of the pulmonary cavity, and in the wall, Each gland branches to form a flattened
there are muscle fibers which seem to mass, closely adhering to the dorsal
be short, oblique, and almost transverse surface of the esophagus. The anterior
(relative to the long axis of the rectum). nerve ring (see below) is anterior to the
The columellar muscle is completely major part of the buccal mass, and well
adnate to the body wall, and gives off in front of the origin of the esophagus
only 3 short branches to the visceral and salivary glands.
organs of the cephalopedal mass: a penis The anterior end of the buccal mass,
retractor, and a right and a left buccal or buccal atrium, is lined with a thin,
mass retractor. acellular cuticle, the anterior margin of
The diaphragm (or floor of the pulmo- which is at the animal's mouth. This
nary cavity) has prominent strips of margin shows darkly colored thickenings
transverse muscles in it, and is closely which are commonly referred to as the
attached to the underlying structures by jaw. The jaw in Chilina fluctuosa (Fig.
means of parenchymatous masses of 5) consists of numerous small plates,
connective tissue. The organ masses extending entirely across the dorsal
filling the isthmian hemocoel and its part of the cuticle, and well down on the
cephalopedal extension are: the buccal lateral margins, thus forming a "horse-
mass (with the anterior nerve ring), shoe pattern". The plates in the mid-
anteriorly and to the left; the penis dorsal region are somehwat longer than
complex, anteriorly and to the right; and the others, and seem to be partly fused.
the albumen gland, spermoviduct and Haeckel noted a much more solidified
vagina, forming a large triangular mass, jaw in C. patagónica (1911: 106, and PI.
posteriorly and to the right. 8, Fig. 22).
The radulais a large, broad membrane,
fhe digestive system in the lower visceral
complex (Figs. 9 and 10). about 2.4 mmwide by 4.2 mm
long. The
The buccal mass is elongate, and sides are parallel. The posterior end
gradually enlarges from the anterior to is slightly expanded, and here the 2 sides
the posterior end. The latter is sub- are rounded, with a median cleft. When
spherical in outline, thin and membranous. dissected out, there is no transverse
This is the radular sac. It does not
flexure and the posterior part is not
project from the buccal mass as a rolled inward.
cylindrical appendage, as drawn by The radula (Fig. 4) contains about 56
Haeckel (1911). The esophagus and ducts rows of teeth. Each transverse row has
of the salivary glands arise dorsally the right and left halves almost at right
from the middle of the buccal mass. The angles to each other, so that the rows are
salivary glands are attached behind to each chevron-shaped. The radular formula
other. The right and left buccal mass is 60-1-60. The central tooth (Fig. 6)
retractors, originating from the colu- is smaller than the first lateral, and
mellar retractor muscle, insert on the asymmetrical, with a prominent, large
buccal mass a little anteriorly and later- mesocone, a small ectocone on the left
ally to the attachment of the esophagus. side, and a mere indication of its corre-
There are 2 small protractor muscles late on the right side, somewhat closer
attaching on the ventral side of the buccal to the base than that on the left. There
mass, at about the level of the esophageal are also 2 small denticles near the base
origin, and these also attach to the wall of the left ectocone, which are not present
of the cephalopedal mass below the mouth. on the right.
The esophagus is short and apparently The first lateral tooth is slightly smaller
364 H. W. HARRY
SEMV
VD
RM
PREP-
FIG. 14. Reproductive system in lower visceral complex, from terminal portion of seminal
vesicle, above albumen gland, to point just prior to genital openings.
FIG. 15. Cross section of the lower part of the spermoviduct.
FIG. 16. The same part of the reporductive system as that shown in Fig. 14, lateral aspect, as
itappears in situ.
FIG. 17. Uterus, with size and position of an included spermatophore outlined in broken line.
Upper lateral tube is the vas deferens, lower (to right) appendage is the vesícula.
FIG. 18. Penis complex, lateral view, in situ.
FIG. 19. Penis complex opened.
FIG. 20. Cross section of penis.
than the second, but from the 2nd to the the sperm groove in the wall of the
50th, all laterals are of uniform size. spermoviduct, but only in its lower half.
The free plate of each tooth consists of They are not present beyond the point
a long, narrow stem, with an expanded, where the vas deferens leaves the
blade-like terminal portion which is spermoviduct.
obliquely oval. The latter is cleft to Below that point, the spermoviduct is
form the 4 cusps, which have curved continued by a thick-walled tube which is
margins and sharp tips. These are not membranous externally, but internally
sharply differentiated as endo-, meso- has longitudinal rugae arranged in an
and ectocones. There is no marked oblique fashion. This tube, without a
division of the teeth into laterals and muscular wall, is here termed the uterus.
marginals, but the last 10 teeth of each It is about as thick as the spermoviduct.
row are vestigial, with only 2 or 3 cusps, At lower end it narrows to become
its
whereas the other teeth have 4, or rarely the vagina. At this point arises a thin-
5 cusps. walled membranous duct, which ascends
The reproductive system of the lower the wall of the uterus to the spermoviduct,
visceral complex (Figs. 14-20). where it expands to form an elongated
The lower end of the hermaphroditic sac. Haeckel called this structure the
duct is a thin, narrow tube which enters vesícula seminalis vaginae.
laterally into a similar tube of slightly In 2 of the specimens a spermatophore
larger diameter, near its tip. This latter was present in the uterus (Fig. 17). It
tube is here called the talon. It is much consisted of a spherical sac, drawn out into
convoluted, and was not accurately a long acuminate tail directed downward
described and figured in Haeckel's toward the vagina. The spermatophore
account. The talon seems to join the was composed of a light brown, non-
upper end of the spermoviduct at the cellular cuticle, which was thickened at
same place where the albumen gland joins. the apical end. Internally, the sac was
The albumen gland is an elongate, flattened packed with a mass of white material,
sac, with glandular walls of moderate consisting entirely of spermatozoa.
thickness (very brittle in preserved The vagina is the part of the female
specimens). It is compressed over the reproductive tube between the uterus and
spermoviduct and traverses the visceral the female reporuductive pore. It is
mass from right to left. cylindrical, of about uniform diameter

The male and female ducts are not throughout its length, and twisted back
separated immediately after passing the and forth upon itself several times (Fig.
albumen gland but form a large tube, 16). Its wall is very muscular, the
the spermoviduct, which has thick, external layer being circular. Numerous
glandular walls, and is much folded upon large fleshy folds, directed longitudinally
itself (Fig. 16), Internally the duct is and frequently anastomosing, occlude the
differentiated into a sperm
somewhat lumen of the vagina. Within the lumen,
groove, with a large fold overlying it, secreted between these folds, are large,
while the rest of the lumen may be con- irregular, calcareous granules (Fig. 12,
sidered the oviducal, or female, part of lower group). Just before the vagina
the spermoviduct (Fig. 15). Very little opens externally, there arises a
could be determined about the possible spermatheca. It has a membranous duct
difference of glandular types in different of almost uniform diameter, which is
parts of the spermoviduct (Haeckel's many times longer than the maximum
account has details), but it was possible diameter of the spermathecal sac. The
to note what seems to be a prostate. latter is subspherical, and situated on
This organ consists of small glandular the left side of the cephalopedal hemacoel,
alveoli, only slightly elongate, arranged wedged in between the pericardium and the
in short, transverse rows. They overlie base of the shell (Fig. 2).
366 H. W. HARRY
The vas deferens is a thick-walled tube and the vas deferens opens terminally.
of uniform diameter throughout its length. The wall of the lower end of the penis
It is also convoluted throughout, and thus sac is swollen (sarcobellum), and there is
several times as long as the actual path a circular groove between it and the
it traverses. After running down the preputium, with a velum separating the
vagina to the area of the female pore, groove from the preputium proper. The
it becomes incorporated in the body wall preputium is thick-walled, with a promi-
-along the external reproductive groove, nent external layer of circular muscles
passing thus to the vicinity of the male surrounding an inner layer of longitudinal
reproductive pore, where it again becomes muscles. There are no pilasters or
free in the body cavity. It then continues preputial organs in the preputium. Its
upward along the penis complex, entering inner surface is almost smooth.
the penis sac at its apex (Fig. 19).
The nervous system (Fig. 13),
The penis complex consists of a pre-
putium and penis sac. The latter is bent The anterior nerve ring consists of
upon itself (Fig. 18), and the flexure is cerebral and pedal ganglia, their commis-
maintained by a sheet of connective tissue sures (transverse) and connectives
with a few muscle fibers radiating from (joining ganglia on the same side of the
the apex to the lower half of the sac on midline). These ganglia do not seem to
the medial side. There are 2 retractor be lobed. This nerve ring passes around
muscles, both attaching at the apex of the the anterior end of the buccal mass (Fig, 9),
penis sac. The larger originates as part well in front of the origin of the esophagus
of the columellar retractor muscle, the (an important point on which Haeckel
smaller (not shown in Fig. 19) originates agrees). The cerebral commissure is
on the body wall anteriorly to the penis, moreover joined to the head by a strong
on the right side. sheet of pigmented connective tissue. The
The penis sac is about 2/3 the diameter buccal ganglia are tucked behind the origin
of the preputium, and about twice as long. of the esophagus, and connected to each
The preputium is swollen at its upper end, other by a long commissure, from the
and tapers slightly to the male pore. The middle of which arises a large nerve which
wall of the penis sac is thin, but its supplies the posterior part of the buccal
inner surface is covered with numerous, mass.
fleshy papillae, which taper to acute points. The posterior nerve ring consists of 5
Some appear to be fused into lamellae, major ganglia joined by long connectives
which in turn anastomose. The papulation (Fig. 13). Close to the anterior nerve
is sparse and the papillae small in the ring on each side is a pleural ganglion,
upper end of the sac, particularly at the each joined to the cerebral and pedal
flexure. ganglia near it by short connectives. The
The penis is a fleshy, elongate structure left pleural ganglion is attached to the
of nearly uniform diameter.
Its external left parietal ganglion (sub-intestinal
surface is drawn out into transverse ganglion of Haeckel's account) by a very
lamellae which are triangular in cross long connective. The left parietal ganglion
section. Haeckel found these cuticularized is situated near the origin of the right
in C. dombeyana, but I was unable to buccal mass retractor muscle. Close to
confirm this in C. fluctuosa. These the left parietal ganglion is a second
papillae tend to anastomose, forming ganglion of the same size, here called
circular rugae. The vas deferens courses the innominate ganglion ("abdominal-2"
through the middle of the penis. It is not of Haeckel's account). This ganglion is
convoluted in that organ, nor does there connected to the left parietal ganglion by
seem to be a space developed between the a short connective, and to the right
vas deferens and penis wall. The penis parietal ganglion by a very long connective
tip is unarmed (i.e., not cuticularized), (or commissure). The right parietal
ganglion lies in the partition between the and these seem to be of themicrogranular
hypopeplar and pulmonary cavities at the type, comparable to those of Lymnaea.
anterior end of the pneumostome (Fig. 8). Several connective tissue membranes are
It is triangular in shape, receiving the present in the hemocoel of the cephalo-
connectives of the posterior nerve ring pedal mass, but these were not studied in
at 2 corners and at the third giving off a detail. Besides the membrane mentioned
nerve which supplies the osphradium. The above, connecting the cerebral commis-
right parietal ganglion and the connectives sure to the head, there is another, trans-
joining it are twisted in such a way (Fig. verse, thin, narrow one passing across the
13) as to give the distinct impression of body cavity immediately behind the buccal
chiastoneury. mass, and with the esophagus passing
The penis nerve arises from the right under it. This membrane may be compa-
cerebral ganglion. There is a prominent rable to the capito-cerebral membrane of
nerve arising from the left pleural the higher limnic Basommatophora (see
ganglion, but I could find none attached to Hubendick, 1962: 20).
the right pleural ganglion. Other nerves
were not studied in detail. Haeckel COMPARISON AND DISCUSSION
describes additional ganglia, which I did OF ANATOMICAL FEATURES
not find, and notes that the pattern of
the posterior (visceral) nerve ring varies The heterostrophic apex of the shell is
in the several species he studied, par- a feature which Chilina shares with the
ticularly in the length of the connectives EUobiidae (but not the Carychiidae), and
between the left pleural and parietal gan- one which is apparently not present in
glia. His figure 46, of Chilina fluctuosa, either the Stylommatophora or the higher
is much closer to what I found in that limnic Basommatophora. The colored
species than is his figure 38, the latter markings of the shell of Chilina represent
being the one copied by Simroth and a feature which is very rare in the higher
Hoffmann (1908-1928). In C. fluctuosa limnic Basommatophora, where the ancylid
a small strand of tissue arises from the Rhodacmea and "Planorbis" eucosmius
left pleuro-parietal connective, from about Bartsch are the only known examples.
where Haeckel (and Hubendick, 1945: 139) The pigment patterns of these latter 2 do
show the left parietal ganglion. This seems not agree with each other or with that
to be no ganglion, and the strand of tissue of Chilina. The markings of Chilina
arising from this part of the connective show a close similarity to the patterns
is only a bloodvessel or a bit of connective in the EUobiidae, as noted earlier (Harry,
tissue. 1951).
The dextral organization of the animal
The circulatory system. in Chilina is a feature found only in the
This system presents nothing unusual, Lymnaeidae and Acroloxus of the higher
i.e., does not differ from the general limnic Basommatophora. Also dextral
pattern found in the higher limnic are: Latia, most Stylommatophora (though
Basommatophora. The heart is 2- there are many exceptions), most EUo-
chambered, the aorta hooks over the lower biidae (exception: Blauneria) and the
loop of the intestine. The most unique Patelliformia (Siphonariidae and Gadini-
feature detected is the pulmonary vein idae).
descending the roof or the pulmonary The
lateral position ofthe eye in /7,
cecum and dividing to supply a vein along in relation to the base of the tentacle,
2 margins of the kidney. is a character shared with Latia
Nothing is yet known about the color of (according to Haeckel, 1911), but with
the blood in Chilina. A few calcareous none of the higher limnic Basommato-
granules could be found in the connective phora. group the eyes are
In the latter
tissue (Fig. 12, upper 3 granular masses). at the inner (medial) side of the base of
368 H. W. HARRY
the tentacles. In the EUobiidae the eyes limnic Basommatophora. Notable also is
are usually at the center of the base. the groove which separates the right
The general form of the tentacle in labial palp from the surface of the head.
Chilina resembles that in Lymnaea and In Chilina and the EUobiidae this groove
Lanx more than that in any other family begins at the lower end of the reproductive
of the higher limnic Basommatophora. groove, i.e., at the male reproductive
The post-tentacular lappet in Chilina pore, but in the higher limnic Basommato-
is probably homologous with the post- phora it never extends that far up on the
tentacular sense organ of the higher side of the head.
limnic Basommatophora, in which its The absence of an anterior pedal gland
under surface contains a sensory with a discrete duct opening at the
epithelium of unknown function. Such a anterior margin of the foot is a character
sense organ is present in all the genera which Chilina shares with the higher
of the higher limnic Basommatophora limnic Basommatophora: gland cells are
which I have examined and is probably concentrated in that area, but open
universal in this group, as (according individually to the exterior. In the EUo-
to Sharp, 1883) was recognized long ago biidae and the Stylommatophora a distinct
by Sarasin and Lacaze-Duthiers. It was gland with duct is present.
described by Sharp (1883) in Ancylus As in all the multi-whorled higher
fluviatilis and Acroloxus lacustris and limnic Basommatophora, the columellar
recently confirmed in the latter by retractor muscle is also adnate to the
Hubendick (1962). Stiglingh et al. (1962) body wall throughout its length in Chilina.
illustrated it in Bulinus, but did not seem In some EUobiidae and in the Stylommato-
to be aware of its function as a sense phora, the main columellar retractor
organ. The organ is not mentioned in muscle becomes free from the body wall
other anatomical accounts (e.g., Haeckel, immediately at its origin. This funda-
Simroth) of Chilina or higher limnic mental difference accounts for the
Basommatophora. In living animals the cephalopedal mass of the Stylommato-
under surface of this tentacular lappet phora being inverted on retraction, where-
has a longitudinal groove, which shows as that of Chilina and the higher limnic
a marked tendency to take up methylene Basommatophora is not. The EUobiidae
blue when it is applied as a vital stain. represent an intermediate stage in this
The post-tentacular sense organ is not respect.
present in the EUobiidae andCarychiidae, One most fundamental characters
of the
nor in the Stylommatophora. of thepulmonate snails is the division of
Whether or not the tesselated surface the original mantle cavity, such as the
of the cephalopedal mass, noted in the one present in the prosobranchs and
preserved Chilina, is evident in the living shelled opisthobranchs, into 2 compart-
specimens or is only a phenomenon of ments. This division was accomplished
preservation has yet to be determined. by a constricting partition of the body wall
The surface of the body of higher limnic and mantle parallel to and near the plane
Basommatophora is smooth in life, with- of the The upper, or
shell's aperture.
out tesselations, and this condition apical, chamber pulmonary cavity,
is the
persists in the preserved specimens. All and the lower, or apertural, remnant has
larger Stylommatophora and some larger been termed the hypopeplar cavity. They
EUobiidae (e.g., Pythia) show such are joined by a hole in the partition,
tesselations, which also persist in pre- the pneumostome. The modification of
served specimens. the size of these 2 cavities, the use to
The reproductive groove present in which they are put, and the several
Chilina is shared with the EUobiidae, pneumostomal mechanisms which have
where its condition varies considerably evolved in keeping with the function of
(Morton, 1955), but is absent in all higher these cavities are all factors which must
be considered in arriving at a sound the pseudobranch, and both are attached

theory of pulmonate phylogeny. along the medial margin of the lens-
Both cavities are present in most shaped pneumostome. When only one of
pulmonates, but the pulmonary cavity the 2 appendages occurs, it attaches along
seems to be lost or much reduced in the whole of the medial margin.
Vaginula and its relatives (Simroth, 1908: The question of homology of the siphon
385). The simplest manifestation of the and the pseudobranch of the higher limnic
division of the mantle cavity, devoid of Basommatophora with each other and
all pneumostomal appendages, is to be with the single pneumostomal appendage
found in Gadinia (Schumann, 1911), where of the more primitive pulmonates such as
it may represent a secondary simpli- Chilina is undecided. Probably the siphon
fication of the more complex condition is not homologous to the pseudobranch,
found in Siphonaria and Willamia. but together they may be homologous to
In the more advanced pulmonates there the single appendage of Chilina, the siphon
are 2 fundamentally distinct types of representing the anterior and the pseudo-
pneumostomal apparatus: The contractile branch the posterior part ofthat structure.
and the non-contractile pneumostome. (1) Observations on living Chilina would help
In the former type the mantle margin resolve this question. The single
is greatly thickened, and occludes the pneumostomal appendage in Siphonari-
hypopeplar cavity. The pulmonary cavity idae, Amphibola and Latia seems
is used only for holding air, never water. anatomically similar to that in Chilina,
The contractile type is found in the No pneumostomal appendage is present
Ellobiidae, Carychiidae, Stylommatophora in Gadinia or Lanx, the "gill" described
and Onchidiidae (and possibly in Gadinia: by Baker (1925) in the latter genus being
see Schumann, 1911: 62). (2) The non- a hemocoel sinus in the swollen mantle
contractile pneumostome of the second margin that is not homologous to
type is associated with a spacious hypo- the pseudobranch of other forms. A
peplar cavity, left unoccluded by the pseudobranch is present in Acroloxus and
mantle margin; the hypopeplar cavity the Ancylidae while a siphon is absent.
may be larger than the pulmonary cavity A siphon is present and a pseudobranch
(e.g., Ancylidae, Acroloxus). In this type is absent in the Physidae and Lymnaeidae
the pulmonary cavity is used exclusively ( except Lanx). In some Planorbidae and
for holding water, or to hold air and Bulinidae both siphon and pseudobranch
water, but never exclusively for air. occur, as noted earlier.
This type occurs in Chilina, Amphibola, The pulmonary cavity lamella is a
the Siphonariidae and in all the higher structure which has been misinterpreted
limnic Basommatophora. in the literature. In Siphonaria, Willamia,
Among the higher limnic Basommato- (but not Gadinia), Latia and Chilina there
phora, the pneumostome may have 2 is a flattened band of enlarged, ciliated
appendages associated with it. (1) The cells running along on the floor of the
pseudobranch, inaptly called gill in some pulmonary cavity from the pneumostome
literature. (2) The siphon, or pneumo- to the apical end of the pulmonary cavity
stomal lappet. The latter is probably (or to heart area, in patellate genera).
present only in those snails which use the There it is reflexed to the roof of the
pulmonary cavity for holding both water pulmonary cavity, where it contiunes,
and air. It is extensile and can be parallel to the first part, back to the
rolled into a tube, the free tip of which pneumostome. The presence of this
breaks thé surface of the water to allow lamella in Amphibola is dubious, as the
an exchange of air in the pulmonary accounts in the literature disagree (see
cavity. When these 2 appendages occur Simroth, 1908-1928: 423, and Hubendick,
toghether (only in some Planorbidae and 1945: 108).
in Bulinidae), the siphon is anterior to Plate (quoted by Haeckel, 1911: 100)
370 H. W. HARRY
long ago realized that this lamella was The posterior margin of the siphon, which
only a single fold, and Simroth (1928: they call "median rectal ridge" scarcely
451) called attention to the similarity of merits a special designation, though it
this fold in Siphonaria to the fold in is large enough in some species to be
"Planorbis" (now Planorbarius). Simroth perplexing. I think Paraense and
is probably correct in considering the Deslandes (1958) confused it with the
pulmonary lamella in those lower pulmonary lamella in Drepanotrema,
pulmonates to be homologous to the which is not present in that genus.
pulmonary fold in some Bulinidae and The pulmonary lamella bears an
Planorbidae, Some members of both intimate and varied relationship to the
families have lost it, but when it is present pseudobranch in different pulmonate
it differs from that of Chiana, etc., in groups. Whether or not it continues over
being elevated from the epithelium of the the lateral surface of the pneumostomal
floor and roof, so that its 2 moieties appendage of the lower pulmonates is often
constitute a real, if ineffective, partition. not clear in the literature, and I was not
The pulmonary lamella may be used to able to determine this point satisfactorally
keep water circulating in the pulmonary in Chilina, although Haeckel figures it as
cavity by its ciliary action. In the extending to the free margin of that organ.
Planorbidae it is generally present in Of more significance is the fact that the
the larger species (Taphius, Helisoma, rectum extends well out on the pneumo-
Planorbarius, butalsoMeneiws) and some- stomal appendage in those forms.
times absent in the smaller forms The pulmonary lamella is not present
{Gyraulus, Drepanotrema s.s,, both of in any pulmonate with a contractile
which I have studied alive). The pneumostome, or with a non-contractile
compartmentalization which this lamella pneumostome which lack a pseudobranch
affords may be an incipient division of (i.e., Physidae, Lymnaeidae and Gadinia).
the pulmonary cavity longitudinally, so However, a pseudobranch may be present

that the right side will be used chiefly in the absence of the pulmonary lamella
for air, the left only for water. (Acroloxidae, Ancylidae). In the Planor-
Stiglingh et al. (1962) noted that the bidae, the 2 structures are either both
dorsal fold of the pulmonary cavity in present or both absent, although in the
Biilinus tropicus was continuous with the latter case the anus may project on a
rectal fold at the apex of the cavity, a small papilla which simulates a reduced
fact which I have been able to confirm in pseudobranch, as I have found in living
Bulinus sp. In preserved material it is Gyraulus parvus.
often difficult to determine this continuity Where both structures are present in the
without serial sections, and live material Planorbidae (e.g., Menetus, Helisoma,
should always be studied to settle this Taphius, Planorbarius), the lamella ex-
point. I suggest that in the Planorbidae tends to the tip of the pseudobranch, and
and Bulinidae in which the dorsal lamella forms a prominent ridge on the lateral
is prominent near the pneumostome, it is surface of that organ. The rectum also
continuous apically with the rectal part extends well out onto the pseudobranch
of this lamella, and I think it is doubtful (Planorbid pseudobranch). IntheBulinidae,
whether one part is ever present without neither the rectum nor the pulmonary
the other in those 2 families. lamella extend onto the pseudobranch, and
Schutte and van Eeden (1959) did not secondary transverse lamellae have ap-
realize the continuity of this lamella peared on both sides of it, which may have
(their "lateral rectal ridge" and "inter- tertiary lamellae on them. This type may
mediate mantle ridge"). The subrenal be termed the bulinid pseudobranch.
ridge may be a new structure in the The pulmonary cecum of Chilina is a
Planorbidae and Bulinidae (larger species feature said by Haeckel to be present
only) which need not concern us here. also in Latia (1911: 101), but it seems
.
not to be present in any other genus of account. The jaw of mulitple plates found
Basommatophora or Stylommatophora. in C. fluctuosa is similar to that found
Haeckel has an interesting discussion of in some Planorbidae, Bulinidae, Ancyl-
the homology of this structure to a similar idae, Acroloxidae and some Stylommato-
one in tectibranch opisthobranchs. phora (e.g.. Punctum), but the Physidae
The osphradium of Chilina seems to and Lymnaeidae always have the dorsal
be identical in form and position with part of the jaw fused into one solid piece
that in Siphonaria and Amphibola (and in (very small in Physa).
some tectibranchs, according to Haeckel, The peculair bulbed radular sac which
1911: 129). Schumann (1911: 63) found no I found in Chilina is unlike the cylindrical
osphradium in Gadinia, and its presence projection which Haeckel figured and
in Willamia is doubtful (Hubendick, 1945: described. His interpretation shows a
93). No osphradium has been found in structure similar to the one in the higher
the EUobiidae or in Stylommatophora. limnic Basommatophora, while what I
Thus, it seems to be absent in those found agrees with the form of this structure
groups with a contractile pneumostome. in some EUobiidae. Moreover, the radular
In the higher limnic Basommatophora flexures (which the ribbon assumes when
the osphradium is on the partition between it is dissected from the tissue surrounding
the pulmonary and hypopeplar cavities, it) in Chilina are unlike those found in
near the pneumostome, but it fronts on any of the higher limnic Basommato-
the hypopeplar cavity, and is thus outside phora, but similar to those of some
the pulmonary cavity. It is conical in EUobiidae.
shape, and its major mass has sunk The radular dentition which Haeckel
below the surface of the mantle. figures (1911, Figs, 24-26), particularly
The kidney in Chilina is essentially a in the form of the central tooth, suggest
large, simple sac, lacking any true ureter similarities with the limnic Basommato-
comparable to that found in the higher phora, but I suspect he has been some-
limnic Basommatophora and Stylommato- what diagrammatic in these drawings
phora. It is thus very similar to the (see Thiele, 1931: 472). The very long
kidney in the EUobiidae, Carychiidae, dorsal, or free, plate of each tooth, with
Amphibola and the Patelliformia. Among its constricted base and expanded end,
the nigher limnic Basommatophora it most is a peculiarity of Chilina found neither
resembles that of Lymnaea, except that in the higher limnic Basommatophora nor
the latter has a very short ureter. Tra- in the EUobiidae. It recalls the condition
beculations in the kidney seem to be a in some unrelated genera of the
function of the size of a given species in Stylommatophora, e.g. Gaeotis, Anoma,
the limnic Basommatophora, being Am.phidromus , Calycia, Sasakina and
present only in the larger species, where Hyalimax, as these raduale are pictured
their nature and extent may be of by Thiele (1931). In some of those genera,
systematic importance at the generic the transverse rows appear to be chevron-
level. Whether solid granules are formed shaped, thus further enhancing the
by the kidney epithelium of Chilina can similarity. Gadinia also has radular
only be determined from living material, teeth with the free plate long and narrow
as these granules do not survive alcoholic (Schumann, 1911).
preservation. Such granules are present The arrangement of the teeth on the
in all higher limnic Basommatophora radular membrane, in angled rows which
which I have examined. They are also form a chevron-shaped pattern, is a
present in the EUobiidae, Carychiidae and condition which Chilina shares with the
Stylommatophora. Physidae (and possibly Acroloxus ?), but
The amount of fusion in the plates of in the other families of the higher limnic
the jaw probably varies in different Basommatophora the transverse rows
species in Chilina, judging from Haeck el's form almost a straight line.
372 H. W. HARRY
The lack of differentiation of the teeth that it lacks sand grains. The presence
into laterals and marginals is another of abundant sand grains in the stomach
character whereby Chilina differs from of Chilina is a feature shared with the
the limnic Basommatophora, In the latter higher limnic Basommatophora (personal
group the lateral and marginal teeth observation; see also Hubendick, 1962:
are differentiated from each other by 33 and André, 1893) and not found in
the change in shape of the tooth, and the the Ellobiidae or Stylommatophora.
increase in number of denticles on the The pattern of the intestine in Chilina,
cutting edge of the marginal teeth. This forming a lower loop apertural to the
transition is often sharp, occurring in stomach and an upper loop apical to the
the space of 2 or 3 teeth, usually about same organ, is characteristic of all limnic
1/3 of the distance between the central Basommatophora. It contrasts with the
tooth and the extreme marginal one. But pattern found in the Stylommatophora,
in Chilina, the only indication of marginals Ellobiidae, Carychiidae and Siphonaria,
is a reduction of the number of denticles in which the intestine does not pass
as the size of the tooth itself is reduced, apically of the stomach. Latia agrees
and this occurs near the extreme margin with Chilina in this feature also. Both
of each row. The Ellobiidae and Latia and Chilina have only a single
Stylommatophora have radular differ- opening of the liver into the stomach,
entiation similar to that in the higher in the pyloric end of the latter, near the
limnic Basommatophora. cecum. This condition is found in Planorb-
Haeckel noted the bilobed nature of the idae, Bulinidae and most Ancylidae
muscular stomach in Chilina. Among the (Hubendick, 1958, found 4 openings in
higher limnic Basommatophora the dis- Ferris sia tenuis), but the Acroloxidae,
continuity of the band of muscle in the Lymnaeidae and Physidae have 2 hepatic
stomach wall is most prominent
in the openings, hence 2 major liver lobes. As
Lymnaeidae; it but little developed
is far as known, all Stylommatophora,
(in some Planorbidae) or absent (the Ellobiidae and Patelliformia have 2 hepatic
muscle tissue forming a complete, un- openings.
interrupted sphincter) in other genera of Haeckel's diagram of the gonad of
this group. The general form of the Chilina is possibly incorrect in showing
stomach in Chilina, i.e., the expanded, the hermaphroditic duct as merely a
membranous anterior and posterior terminal continuation of the major
portions and a sphincter-shaped muscle collecting duct of the gonad. In the species
between which encircles the single tube, I examined the hermaphroditic duct arises
is a condition characteristic of the higher well above the lower end of the collecting
limnic Basommatophora which contrasts duct. The collecting duct of Chilina is
with the shape and structure of the stomach probably homologous to the gonad atrium
in the Ellobiidae, Carychiidae, and of the higher limnic Basommatophora,
Stylommatophora. Moreover, the pres- The gonad in that group consist of a
ence of a gastric cecum opening into the short, swollen, sacculate atrium, along
pyloric part of the stomach is a feature which the gonad follicles are crowded,
which Chilina (and Latia: see Simroth, and into which they open almost directly,
1908-1928: 339) shares with the higher having at most very short, wide ducts
limnic Basommatophora. Such a structure which are only occasionally branched. The
is absent in the Stylommatophora and sacculate atrium is as wide as it is long
Ellobiidae. Hubendick (1962: 34) indicates in patelliform genera (Ancylidae, Acrolox-
a crystalline style in this cecum in idae, and Lanx), and therefore the relation
Acroloxus. A style is not known from of the hermaphroditic duct to the atrium
any other pulmonate snail. He also is rather meaningless in them. But in
suggests that the stomach of Acroloxus those families in which the atrium is
lacks a muscle and specifically states elongate, the hermaphroditic duct arises
at the apertura! end in the Planorbidae and limnic Basommatophora, i.e., one arising
Bulinidae, and about midway the length of from the lower end of the vagina. The
the atrium in Physidae and Lymnaeidae function of the spermatheca in the
(except Lanx). It is not practical to pulmonates not well known, and this
is
formulate extensive generalities about the structure is only thus called for want of
form of the gonad in the Ellobiidae and a better name. The homology of the
Stylommatophora at present, but suffice it vesícula seminalis vaginalis of Chilina
to say that both types may be present in furnishes an interesting problem. Such
both groups. a structure seems to be present, in
The condition of the seminal vesicle, Siphonaria, Latia, and Otina, all of which
i.e., whether it forms a convoluted or have also a true spermatheca arising
outpocketed part of the hermaphroditic from the vagina near its opeining. The
duct (or both, as in Chilina) is a character Ellobiidae and Stylommatophora may have
which is variable at the generic level in only the upper of these 2 structures.
the higher limnic Basommatophora. In In Chilina the male and female openings
the Stylommatophora and Ellobiidae the are widely spaced, as in the Ellobiidae,
hermaphroditic duct may also be con- in Gadinia and in the higher limnic
voluted, though an outpocketed condition Basommatophora; but the external
is unknown in those groups. In the reproductive groove seems to be present
Siphonariidae both conditons occur only in Chilina (and Latia?) and in the
(Hubendick, 1945). Ellobiidae. The peculiar digitform
The peculair elongate tube continuing the appendages of the penis of Chilina are
hermaphroditic duct in Chilina, which I not found in any of the higher limnic
have called the talon, is not well under- Basommatophora, but such ornamentation
stood. It is lacking in the higher limnic recalls the condition in Onchidium (see
Basommatophora. Haeckel's figure of this Simroth, 1928: 507). The male genitalia
part of theanatomy is vague. The presence oí Chilina, Latia, Gadinia, the Ellobiidae
of a spermoviduct in Chilina is a feature and higher limnic Basommatophora show
shared with the Ellobiidae, Stylommato- a fundamental similarity in that there is
phora, Patelliformia, Latia and Amphi- a differentiation of a preputium and penis
bola. Only Acroloxus of the higher limnic sac, the latter containing the penis, which
Basommatophora has this condition. In has the vas deferens opening at the tip,
the other genera of that group, the male or near it. This condition contrasts
and female tubes are completely separated markedly with that in the Stylommato-
(diaulic) below the albumen gland. phora (and some Siphonaria), where there
In being incorporated within the male is no differentiation into penis sac and
part of the wall of the spermoviduct, the preputium and where the vas deferens
prostate gland of Chilina resembles that may be dissociated from the "glans",
of Lymnaea, Siphonaria, the Ellobiidae which may correspond to the penis proper
and of the Stylommatophora. The higher (including the pseudopenis or ultrapenis)
limnic Basommatophora other than of the higher limnic Basommatophora.
Lymnaeidae have a prostate gland made The calcareous granules present in
up of a few to many free follicles, clustered the lumen of the vagina have not been
along the vas deferens (or along a separate reported in any of the higher limnic
duct in some Planorbidae: see Hubendick, Basommatophora, nor previously in
1955). Chilina. Possibly these are incorporated
The uterine cecum, or vesícula in the egg mass, as a shell, in this
seminalis vaginalis, arising from the species. The presence of a spermato-
upper end of the vagina, grossly resembles phore in Chilina is a character whereby
a spermatheca. It might have been called they differ from higher limnic Basom-
that, were it not for the presence also of matophora, where such a structure is
a spermatheca typical of that in the unknown. Some Stylommatophora and
374 H. W. HARRY
Siphonariidae (see Hubendick, 1945) have Lymnaeidae (and LanxT) among the higher
a spermatophore. limnic Basommatophora. The other
The anterior or cerebral nerve ring of families which I have been able to investi-
Chilina is probably permanently situated gate (Physidae, Planorbidae, Bulinidae,
around the front end of the buccal mass Ancylidae) have granules, though only
in life. In the higher limnic Basommato- of the macrogranular type, i.e., they are
phora this ring is always behind the buccal much larger than the microgranules. The
mass, and thus behind the opening of the condition in Acroloxus, Lanx and Latia is
esophagus and salivary glands. Its position unknown. At least some Stylommato-
may vary in different Ellobiidae, and it phora have granules of the microgranular
is generally behind the buccal mass in the type.
Stylommatophora. According to my
observations, the posterior (or visceral) PHYLOGENETIC CONSIDERATIONS
nerve ring has on it only 5 ganglia.
Haeckel's description of additional minute An attempt is here made at evaluating
ganglia is not very convincing, nor does the several characters discussed above
he illustrate them. However, Hubendick relative to their appearance in the evo-
(1945: 139) agrees with Haeckel in finding lution of Chilina and the higher limnic
one further ganglion which I can not Basommatophora. The scheme is de-
prove to be such. The homology of these pendent first on discerning what
ganglia with the ganglia of other pulmo - consitutes a genetically controlled charac-
nates is still an open question, but it ter, and determining when it may have
seems probable that these are the same appeared or undergone modification in
5 to be found on the posterior nerve ring relation to comparable changes in other
of the limnic Basommatophora. In Chilina characters. We assume that evolution
this ring is large, extending well back proceeded by the addition and loss of
into the hemocoel of the lower visceral characters, and that once a character
complex. This condition prevails in the is lost, it is not likely to reappear in
Ellobiidae also. But in the Stylommato- the progeny of the group which lost it
phora and higher limnic Basommatophora (there may be exceptions). Small changes
the connectives are short, so that this of form, or of the relative position of
ring is small, and its ganglia are closely organs may have been more reversible
approximated to each other and to the than the change from presence to absence
anterior nerve ring. No chiastoneury is and back to presence of a given feature.
indicated in the higher limnic Basommato- As others have noted, the patellate
phora, but this condition is strongly form probably arose several times
suggested in Chilina. independently in the Gastropoda (Baker,
It would be of interest to know the color 1925), Sinitsin (1931: 802-807) proposed
of the blood in Chilina. In the Stylommato- an interesting theory relating the loss of
phora and the basommatophoran families trematode infection to the secondary
Ellobiidae, Carychiidae, Physidae, Ancyl- reversion to a limpet form. Apart
idae and Lymnaeidae the blood is color- from the primitive patellate form
less, or lacks sufficient hemoglobin to be thought to be ancestral to all gastropoda,
grossly detectable. This condition varies and perhaps represented today by Neo-
at the subfamily or generic level in the pillina, the patellate form resulted in
Bulinidae and Planorbidae. Thus the blood many later groups through the loss of
is red in Taphius, Helisoma, Menetus and the ability to form a large, multi-
at least some Bulinus, but it is colorless whorled visceral mass. Nearly every
in Drepanotrema, Gyraulus and Plesio- family of the group we are presently
physa. The calcareous granules of the considering has given rise to patellate
connective tissue of Chilina are of the descendants. Possibly Gadinia and
microgranular type, found only in the Siphonaria (and with the latter Willamia)
are independent patellate derivatives of probably had the following characters by

what is termed below the "Primitive the time they were recognizable as
Pulmonate Stock", before the "Urbaso- pulmonates.
mmatophoran Stock" had originated. As 1. They were probably marine.
in the case of Acroloxus, the ancestors 2. The foot was broad, oval, used for
with well-coiled visceral mass from which creeping. The visceral mass was large,
they were immediately derived may be multi-whorled, and the shell high-spired.
extinct today. The shell had an heterostrophic, punctate
apex. Both the shell and animal were
PRIMITIVE PULMONATE STOCK
dextral. An operculum was present.
i 3.The mantle cavity was constricted
URBASOMMATOPHORA
transversely, so that pulmonary and hypo-
Ellobiidae peplar cavities were differentiated, with
PROCHILINID STOCK a pneumostome between them. The
pneumostome was non-contractile, and
Chilinidae there was a single pneumostomal
Latiidae appendage projecting into the hypopeplar
cavity. A pulmonary lamella and pulmo-
PROACROLOXID STOCK nary cecum were present.
(unknown family) 4. An osphradium was present, forming
an elongate oval projection from the
Acroloxidae
surface of the mantle epithelium, with a
PROLYMNAEID STOCK longitudinal groove on its surface. It
"Lymnaeidae was situated just inside the pulmonary

cavity, at the anterior end of the pneumo-
Lancidae stome.
PROPHYSID STOCK 5. A radula was present. The number
of teeth in a transverse row was large,
^"^Physidae
and these were not differentiated into
PROPLANORBID STOCK laterals and marginals, other than by a
slight reduction in size, and loss of
Planorbidae
^ 9 denticles on teeth near the margin. The
Ancylidae rows were arranged in a pattern of
Bulinidae chevrons.
6. The anterior margin of the atrial
Diagrammatic scheme showing the cuticle of the buccal mass was thickened
FIG. 21.
possible phylogeny of the Chilinidae to form a jaw, the thickening consisting
and the higher limnic Basommato- of discrete platelets. These always
phora. included the median dorsal margin (in
contrast to many prosobranchs).
The following scheme (see also Fig. 21) 7. The intestine extended both above
attempts merely to outline a pattern of (apicad) and below (aperturad) the
evolution which relates the Chilinidae to stomach. Two liver lobes were present;
the higher limnic Basommatophora. It there was no gastric cecum, nor a crystal-
is cumulative, meaning that any feature line style,
not specifically stated as being modified 8. They were hermaphroditic, with the
or lost at a given level remains as it sperm and ova produced in the same
was before that level was attained. follicles. The sperm were of only one
type (eupyrene; in contrast to many
Primitive Pulmonate Stock. prosobranchs). The gonad was multi-
The first snails to attain a level con- foUiculate, with the follicles arranged in
sidered as "Primitive Pulmonate Stock" bunches (with branched ducts) widely
376 H. W. HARRY
spaced along a narrow collecting duct. immediately ancestral to the Ellobiidae
The hermaphroditic duct, draining the and Chilinidae, which may be termed
gonadal collecting duct had some modi- "Urbasommatophora" (following Burch,
fication for holding sperm, either by 1962) had undergone the follwing modi-
being swollen and convoluted, or out- fications:
pocketed, or both (the siminal vesicle). 1. The reproductive system had become
At the lower end of the hermaphroditic diaulic at least for the lower half of its
duct was a large, compact albumen gland, course between the albumen gland and
which had a single short duct connecting the exterior. A prostate gland was present
it to the hermaphroditic duct. Beyond the in the lower part of the spermoviduct.
junction of these 2 structures, the A vas deferens coursed along the uterus
reproductive tube became thick-walled and vagina, as a separate duct. The 2
by the addition of gland cells, and the sex openings were widely separated on
cavity was divided longitudinally into 2 the right side of the body, and an external
parts (spermoviduct). The reproductive reproductive groove connected them. The
system was monaulic for most, if not male intromittent organ contained a penis,
all, of its passage between the albumen i.e., a projection of the upper end of the
gland and the exterior. A vesicula sac, with the vas deferens running through
seminalis vaginae was present at the upper it and opening at or near its tip. The
end of that portion which served as vagina, preputium was structurally not well
and a spermatheca was present near the differentiated from the penis sac.
opening of the female part of the tract. 2. No ureter was present. The kidney
The intromittent organ of the male part contained solid secretory granules in its
of the system consisted of an expanded epithelium. This organ was restricted
part of the vas deferens (or terminal entirely to the roof of the pulmonary
part of the spermoviduct, if it was mona- cavity, not touching on the upper visceral
ulic to that extent). This organ was complex or diaphragm.
everted in copulation. No permanently 3. The teeth of the radula became differ-
external male genitalia were present. The entiated into laterals and marginals, by
eggs were encased in a gelatinous mass, addition of denticles to the marginals,
and a free swimming veliger larva was and by a change in shape of the teeth.
present. A spermatophore was present. The transition was sharp.
9. The musculature of the buccal mass 4. One pair of broad, flattened tentacles
and stomach contained a red pigment. was present, with the eyes situated at the
This pigment was not present in the outer side of their bases. The mouth
columellar retractor muscle. The latter was surrounded by a pair of labial palps.
was adnate to the body wall throughout 5. The external surface of the body had
its length, andthecephalopedal mass could become tesselated, at least in larger

not be inverted when the snail retracted. species.
10. The anterior and posterior nerve 6. The calcareous granules of the
rings had long connectives and commis- connective tissue were present, and of the
sures. The anterior nerve ring encircled microgranular type, or of both micro-
the buccal mass anteriorly. Chiastoneury and macrogranular types,
was present. 7. The operculum was lost in the adult,
but retained in the embryo.
Urbasommatophora.
Tne Ellobiidae s.s.
Bythe addition and loss of characters, This family probably evolved directly
and changes in form and position of those from the urbasommatophoran stock by
which remained, several groups of the loss of the spermatophore, pneumo-
pulmonates may have been derived from stomal appendage, pulmonary cecum
this primitive pulmonate stock, only one and lamella: the acquisition of a con-
of which need concern us here. The group tractile pneumostome; by moving the
intestinal loops entirely below the stomach 3. Loosing the anterior liver lobe.
(aperturad); by modifying the shape of 4. Ornamenting the penis and internal
the tentacles to elongate, cylindrical, and surface of the penis sac with fleshy
moving the eyes to the center of their papillae (this may, however, be a primi-
bases; and by developing a few charac- tive character which was later lost by
teristics which are entirely peculiar to the Ellobiidae and higher limnic Baso-
them, but need not detain us here. mmatophora).
5. Producing calcareous granules in the
Prochilinid Stock.
lumen of the vagina.
A Prochilinid stock evolved from the 6. Loosing apical punctation of the shell.
Urbasommatophoran stock separately
The Latiidae .
from the Ellobiidae, by the following

This family may represent a patelli-
modifications:
form derivative of the Chilinidae, after
1. The formation of a gastric cecum
the latter evolved from the prochilinid
(which probably does not have its counter-
stock. After differentation (or special-
part in the Ellobiidae, contrary to
ization) of the modern Chilinidae, the
Morton's account, 1955). The musculature
prochilinid stock became further modified
of the stomach completely surrovinded the
to produce the proacroloxid stock, as
tube ( was not merely a bulge on one side
follows.
of the tube, as in Siphonaria, Stylommato-
phora and the Ellobiidae) and was divided Proacroloxid Stock.
into right and left lobes.
1. The
2 nerve rings became con-
2. A post-tentacular sense organ was
stricted, through shortening of their
produced.
commissures and connectives, and the
3. The animals now lived in fresh
anterior nerve ring shifted to a position
water, rather than in a marine environ-
behind the buccal mass. This may have
ment. The larval stage was retained
occurred earlier, or independently in the
within the egg.
several pulmonate groups (Stylommato-
4. The preputium became differentiated
phora, some Patelliformia); only a slight
from the penis sac by an increase of
change of form is involved, i.e., the
musculature in its wall.
length of the commissures and con-
5. The albumen gland was fusiform,
nectives. Chiastoneury was obliterated.
and situated at the lower end of the
2. The osphradium shifted to a position
upper visceral complex, with its long
fronting on the hypopeplar cavity, near
axis transverse to the visceral mass
the pneumostome. At the same time it
(this may have characterized an earlier
sank into the body wall, forming a cone,
stage).
with a very small, exposed, functional
6. The operculum was lost in the embryo
surface.
as well as in the adult. This may only
3. The eyes moved to the medial side
have happened in the proacroloxid stage;
of the base of the tentacles.
the embryology of Chilina and Acroloxus
4. The external reproductive groove
is completely unknown.
was lost.
The modern Chilinidae . 5. A ureter was added to the kidney.
This family diverged from the 6. Heterostrophy of the shell's apex
prochilinid stock by the following was lost.
modifications: 7.The collecting duct of the gonad
1.Loosing the ability to differentiate became enlarged, sacculate, with the
the marginal from the lateral teeth of follicular ducts shortened, expanded, and
the radual. crowded together along one surface of
2. Modifying the r adular teeth into a the atrium.
distinctive type with the elongate, con- 8. The vesícula seminalis vaginalis was
stricted base of the free plate. lost.

378 H. W. HARRY
9. The tesselation of the body surface seem eminently suited to the presence
was lost. of a ureter.
10. The pulmonary cecum was lost. 3. The gonad became imbedded on the
columello-parietal side of the upper liver
The modern Acroloxidae .
lobe, well below the apex of the shell,
This family was derived from the
and the hermaphroditic duct emerged
proacroloxid stock by the following about midway the length of the elongated
changes: gonadal atrium. This, too, may have
1. appeared a unique configu-
There occurred in the proacroloxid stock.
ration theof ureter, which loops
completely around the kidney, unlike that The Lymnaeidae s.l.
in any other basommatophoran. This family differentiated directly from

2. A patelliform shape was assumed, the prolymnaeid stock by the following
with concomitant loss of size and loss of modifications:
coiling of the visceral mass. The pulmo- 1. Loss of the pulmonary lamella and
nary cavity became greatly reduced, and pseudobranch, but retention of the siphon.
the pulmonary lamella disappeared. From 2. Straightening of the transverse rows
Hubendick's account (1962) it is not of radular teeth (loss of the ability to

entirely clear whether the pseudobranch form a chevron pattern).
is the bulinid type (doubtful), or a
like 3. Loss of apical punctation of the shell.
simple, much folded gill with no true 4. Solidification of the jaw into a single
transverse lamellae. piece dor sally.
3. The musculature of the stomach was Most other characters of Lymnaea are
reduced, if not entirely lost. essentially retention of characters
As in all patelliform higher limnic
4. gained up to this time, and lost or modi-
Basommatophora, the gonadal atrium fied by other groups later. Thus, they
became funnel-shaped, with the her- retained the dextral body form, the 2 lobes
maphroditic duct emerging from the lower of the liver, the bilobed musculature of
end of the funnel. It is not clear from the stomach, the microgranular form of
Hubendick's account whether the prostate the connective tissue granules, the
is like that of Chilina and Lymnaea, or prostate follicles imbedded in the wall of
that usual in the higher limnic Baso- the male tube (vas deferens, the sperm-
mmatophora, and it may be transitional. oviduct having been completely divided),
5. The tentacular shape is not clear the flattened, triangular tentacles, and a
from Hubendick's account, but it may be large, sacculate kidney with a simple,
flattened, as in Chilina and Lymnaea. short ureter.
6. The apical punctation of the shell was There have been some additional
lost. The suggestion of hyperstrophy in changes at the generic and specific level,
the shell is not easily explained (see such as the extension of the mantle over
below, proplanorbid stock). the outer surface of the shell in Myxas
(a simple change in form, probably inde-
Prolymnaeid Stock. pendent of that in Physa); an expansion
of the hemocoel of the penis, with the
After the emergence of Acroloxus, the vas deferens sometimes convoluted in
proacroloxid stock evolved into the pro- this hemocoel, or the penis convoluted
lymnaeid stock by the following changes: in the penis sac; the greater infolding
1. The male and female reproductive of the wall of the prostatic part of the
tubes became completely separated as vas deferens; the reversal of symmetry
far up as the albumen gland (i.e., com- in some species, so that a sinistral snail
pletely diaulic). is produced. For details see Hubendick,
2. A
siphon was added to the margin 1951.
of pneumostome.
the This may have
happened at the proacroloxid stage, and The Lancidae.
been lost in Acroloxus s.S., for it would Lanx was probably derived directly
.
from Lymnaea after the latter had condition or one of loss is not known, but
diverged from the prolymnaeid stock, by the reflection in Physa is probably in-
becoming patelliform and loosing the dependent of that in Myxas.
siphon (the pseudobranch had been lost 3. Aglandular area appeared in the wall
by Lymnaea' s ancestors). of the preputium.
4. The kidney and ureter became con-
Prophysid Stock.
voluted into a distinctive pattern,
After the differentiation of theLymnae- simulated, however, in the planorbid genus
idae, the prolymnaeid stock evolved into Parapholyx
the prophysid stock by the following 5. The jaw was reduced to a very small,
changes: single thickening in the midline, dorsally.
1. Modification of the tentacles to a 6. The pseudobranch, and with it the
filiform shape, with a circular cross pulmonary lamella, were lost,* but the
section. siphon was retained.
2. Differentiation the prostate to
of 7. The was modified (elongated,
foot
form free attached in a line
follicles tapered), so that these snails became
along the wall of the vas deferens, but rapid crawlers. This change merely
no longer imbedded in the wall of that necessitates a change in form, and was
duct. probably independent of that in the
3. Inversion of symmetry from a dextral planorbid Drepanotrema s.s.
which probably
to a sinistral organisation, 8. The apical punctation of the shell
did not involve hyper strophy (see below). was lost.
4. Modification of the calcareous 9. The r adular sac was modified, so
granules of the connective tissue from that the end turns up posteriorally, unlike
the microgranular type to the macro- that of any other member of the limnic
granular type (or suppression of the Basommatophora. An appendage was
microgranular type, if both were present added to each radular tooth, but the
previously, as suggested above). chevron -shape of the radular rows was
5. Conversion of the bilobed muscle retained,
of the stomach to a simple sphincter.
Proplanorbid Stock.
The Physidae .
This family was differentiated at this After the differentiation of the Physidae,
stage, by the following modifications the prophysid stock evolved into the pro-
(among others; this list is far from planorbid stock by the follwing changes:
complete): 1. The pulmonary lamella was elevated
1. Originating a distinctive distribution from the floor and roof of the cavity,
of the musculature of the dorsal part so that its 2 moieties formed an in-
of the cephalopedal mass and of the effective longitudinal partition,
partition between the hypopeplar and 2. The anterior
liver lobe with its duct
pulmonary cavities, and in the roof of was so that there was only a single
lost,
the pulmonary cavity, with a loose opening from the liver into the stomach.
attachment of this muscle mass on the 3. The transverse rows of radular teeth
lip of the shell. This may be termed the were straightened.
physid musculature. It seems to have 4. Hemoglobin was developed in the
been completely ignored in the literature; blood, but independently lost in different
however, its anatomical details are too subgroups.
extensive to describe here. 5. The hermaphroditic duct shifted to
2. Reflecting the mantle margin over a terminal exit at the apertural end of
the outer surface of the shell, and forming the gonadal atrium, and the whole gonad
a digitate margin on it. Whether the shifted to a position apical to the liver.
absence of this reflection in Aplexa (and 6. The tendency for hyperstrophy
Petrophysa 7) represents a primitive appeared, Hyperstrophy is the incongruity
380 H. W. HARRY
between the symmetry of the animal and 5. The shell became planispiral inmost
the symmetry of the shell, i.e., a dextral genera, or ultradextral (i.e., a distinctly
animal in a sinistral shell, or vice versa. dextral shell with sinistral animal), as
Pelseneer (1906: 82) defined the 3 phe- in Carinifex, Parapholyx, Acrorbis and
nomena, heterostrophy, hyperstrophy and some phenotypes of Taphius andecolus.
inverted symmetry (his situs inversus The planispiral stage represents incipient
viscerum) very adequately, giving hyperstrophy, and the ultradextral shell
examples of each. Walker (1920, 1923) represents hyperstrophy completed.
discussed the phenomenon of hyperstrophy Some characters present in the pro-
in the patelliform pulmonates, but he was planorbid stock were lost independently
mistaken in 2 points: (1) in saying that in different subfamilies and genera, after
the embryonic shell of the Ancylidae is the Planorbidae s.s. had arisen, e.g.,
lost in the adults, and (2) in asserting apical punctation of the shell, the pseudo-
that none of the ancylids show the phe- branch, the hemoglobin in the blood.
nomenon of hyperstrophy. He seems to Whether any Planorbidae s.s. have
have misinterpreted the coiling of the retained the pseudobranch but have lost
shell in these snails, not visualising their the siphon is unknown. Many additional
shells as simple cones, nor being able characters, either unique to the
to project their direction of coiling Planorbidae or shared with some Bulin-
sufficiently well to understand the true idae, are present, but are limited to a
nature of coiling. It is quite possible few subfamilies and genera. These include
that hyperstrophy in the pulmonates is flagellae on the penis sac (which may
a stage leading to the reduction of the be a primitive character lost by other
visceral mass found in the patellate forms families independently), development of
(see the theory proposed by Sinitsin, 1931). dentition in the shell's aperture, a pre-
putial organ, distinctive patterns of
The Planorbidae s.s.
melanin pegmentation in the cephalopedal
This family, exclusive of the Bulinidae, mass and pulmonary cavity, and many
has evolved from the proplanorbid stock others.
by the following modifications: The Planorbidae s.s. probably repre-
1. A simple, reflexed hook was formed
sent the main stem of the phylogenetic
in the lower end of the ureter.
tree of the limnic Basommatophora. They
2. The siphon was probably always
are today the most diversified of all the
retained, but the pseudobranch, and with
families in that group, even if we exclude
it the pulmonary lamella, have been To include the latter in
the Bulinidae.
independently lost in different subfamilies
the Planorbidae as a subfamily will
and genera. ultimately necessitate the recognition of a
3. The prostate follicles retained their
taxon between the subfamily and generic
linear arrangement along the vas
level, thus needlessly compounding con-
deferens, or, in some cases, along a
fusion in systematics. That some Bulin-
separate duct entering the vas deferens idae share peculiar characters with some
(see Hubendick, 1955). They did not Planorbidae may only indicate that the
become crowded into the rosette pattern
definition of the proplanorbid and earlier
characteristic of the Bulinidae.
stages given above needs revision and
4. The albumen gland became flattened
extension, and this in turn can only be
and L-shaped, with one portion filling done after further studies on more
the lower end of the upper visceral complex
material, preferably living.
transversely, and the other passing
peripherally, either under (Drepano- The Bulinidae.
trematinae) or over (Taphiinae, Heli- This family probably evolved from the
somatinae) the lower loop of the intestine. proplanorbid stock independently of the
The peripheral portion nearly covers the Planorbidae s.S., by undergoing the
stomach. following modifications:
1. The ureter forms

2 flexures inmost, 3. Reduction in size and assumption of
if not species (the literature is far
all the patellate form, with loss of the siphon
from clear on this point: Hubendick, and pulmonary lamella, but retention of the
1955), and the extent of the ureter between pseudobranch (which is bulinid in at least
these flexures is fairly long. some genera).
2. The siphon is always retained (if 4. A
reduction in the number of chromo-
we relegate Protancylus to the Ancylidae, somes from the common number found
or to a family of its own). The pseudo- in other limnic Basommatophora (see
branch is retained (except in Campto- Burch, 1962, who, however, relates
ceras ? see Hubendick, 1959) but the pulmo- progressive evolution with increase in
nary lamella never extends onto its chromosome number).
surface, nor does the rectum. Instead,
Arguing in favor of a polyphyletic origin
transverse secondary and tertiary
is the following evidence:
lamellae developed on both sides (bulinid
1. Variation in the type of penis (true
pseudobranch).
penis or pseudopenis).
3. The albumen gland retained the fusi-
2. Flagellum present or absent on the
form shape (also in Indoplanorbis ? the
penis sac.
literature is not clear).
3. Various r adular types.
4. The prostate follicles have become
There also exists a further modification
crowded to a small part of the vas defer ens, of the ancylid form, namely the develop-
forming a rosette.
ment into the "Gundlachia" stage (see
5. The shell remained sinistral, or
Basch, 1959). Be it noted that this stage
rarely became planispiral (e.g., Indo-
is not a reversion to one with a multi-
planorbis, Amerianna; see Hubendick,
coiled visceral mass, although it does
1948), but the ultradextral shell with
result, at least in its more extreme form,
elongated spire has not been attained.
in a larger animal.
The loss of apical shell punctation, Thus, by using many characters which
of hemoglobin in the blood, etc., may have
vary within a large group of organisms,
occurred independently in different sub- such as the limnic Basommatophora, and
families and genera. Shell dentition was evaluating them in relation to each other,
never developed (or may have been lost it is possible to construct an outline of
in all), and the penis has been transformed
phylogeny as a continuum. The taxa into
into a pseudopenis (or ultrapenis) in some
which this continuum is divided, and the
bulinids, whereas none of the Planorbidae
relative ranking of same, i.e., whether a
have such modifications. particular subgroup represents a genus,
The Ancylidae. subfamily, etc., is largely arbitrary. At
This family possibly arose either from the present stage of malacology as a
several distinct subgroups from within science, endeavor would be most
the Planorbidae and Bulinidae (poly- profitably directed to defining these taxa,
phyletic origin), or they may have arisen in terms of morphological charac-
from one group (monophyletic origin), teristics, rather than be merely limited
in which case it was possibly the Bulin- to establishing identities of a subgroup as
idae. Evidence speaking for a mono- a "type" for a larger group, and to the
phyletic origin is: resulting debates over the relative ranking
1. Common egg type (see Bondesen, to be accorded individual taxa.
1950) not found in other families of the It is at present impractical to make
limnic Basommatophora. analyses of the several groups of the

2. Common pattern of the ureter not higher limnic Basommatophora touched
found in any other family of the limnic on above in greater detail. This can
Basommatophora, but more similar to only be done when much more has been
that of the Bulinidae than to that of the accomplished in the way of comparative
Planorbidae. anatomical studies, including accounts of
382 H. W. HARRY
the characters noted herein and , 1948, Studies on Bulinus.

others which will undoubtedly be Ark. Zool. 40(16): 1-63.
discovered subsequently to be of value. , 1951, Recent Lymnaeidae.
Doubtlessly much revision of the more Kungl. Svenska Vet. Handl. 3(1): 1-223.
primitive stocks defined above will have 1955, Phylogeny in the Planor-
,
to be made when we attempt to integrate bidae. Trans. Zool. Soc. London, 28:
the Patelliformia, Ellobiidae,Stylommato- 453-541.
phora and other groups into a complete 1958, On the family Ancylidae
,
scheme of pulmonate phylogeny. I have with special regard to Ferrissia tenuis

tried to integrate the interesting studies (Bourguignat) the suspected inter-
by Walter (1962) on apical shell mediate host of Schistosoma haemato-
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but have mostly neglected the valuable Trop. Med. and Malaria 2: 17-21.
data by Burch (1962) on chromosome 1959, A note on Camptoceras
,
numbers and that by Bondesen (1950) prasfiadi (Clench). Venus 20: 263-271.
on egg types. However, the outline as 1962,
, Studies on Acroloxus
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patible with their data. Mus. Zool. Avd. 133:1-68.
MORTON, J. E., 1955, The evolution
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et PARAENSE, W. L. and N. DESLANDES,
Ancylus lacustris et fluviatilis. Rev. 1958, The Brazilian species of
Suisse Zool. 1: 427-461. Drepanotrema. VI. D. kermatoides
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egg capsules of freshwater pulmonate SCHUTTE, C. H. J. and J. A. VANEEDEN,
gastropods. Natura Jutlandica3: 1-208. 1959, Contributions to the morphology
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Anatomie der Gattung Chilina. Zool. pp 214-240.
Jb. Suppl. 13, Bd. 4, Fauna Chilensis, SIMROTH, H. and H. HOFFMANN, 1908-
p 89-136. 1928, Pulmonata. in Bronn's Klassen
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HUBENDICK, ., 1945, Phylogenie und Phylogenie der TrematodenV. Revision
Tiergeographie der Siphonariidae. of Harmostominae in the light of new
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STIGLINGH, L, J. A. VAN EEDEN and 1923, Ancylidae of South Africa.
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P. A. J. RYKE, 1962, Contributions Privately printed, 82 pages,

to the morphology of Bulinus tropicus. WALTER, H, J,, 1962, Punctation of the
Malacologia, 1: 73-114. embryonic shell of Bulininae (Planorb-
THIELE, J., 1929-1935, Handbuch der idae) and some other Basommatophora
systematischen Weichtierkunde, 4 Vols. and its possible taxonomic-phylogenetic
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WALKER, B,, 1920, Hyperstrophy in
ZUSAMMENFASSUNG
ERNEUTE UNTERSUCHUNGEN AN CHILINA FLUCTUOSA UND
ENTWURF EINER PHYLOGENIE DER HÖHEREN LIMNISCHEN BASOMMATOPHOREN
Wiederuntersuchung der grî5beren Anatomie dieser Art zeigte mehrere bei den
Chiliniden noch nicht beschriebene Merkmale, wie z. B. einen post-tentakulären Lappen:
Anwesenheit von Sandkörnern im Magen; einen Spermatophor; Verwachsung des Musculus
retractor columellaris mit der Körperwand; Zotten an der Haut der cephalopedalen
Körpermasse und Anwesenheit von Kalkkörnchen im Bindegewebe sowie im Lumen
der Vagina.
Chilina wurde anatomisch mit den höheren Basommatophoren des Süsswassers
verglichen, und gelegentlich auch mit anderen Hauptgruppen der Pulmonaten. Diesen
Vergleichen schliesst sich ein vorläufiger Entwurf einer Phylogenie für Chilina und
die höheren limnischen Basommatophoren an, worin die einzelnen Stadien ihrer
Evolution definiert sind. Der Umriss beginnt willkürlich auf einer "Primitiven Pulmo-
naten-Stufe" und schlägt vor, wann welche Merkmale hinzugefügt, verändert oder
verloren wurden, in Bezug auf ähnliche Veränderungen anderer Kennzeichen. Eine
Reihe von Zügen charakterisiert jede der verschiedenen hypothetischen Entwicklungs-
stufen , aus deren jeder eine oder mehrere der heute lebenden Familien entsprossen
sind. Die Bezeichnungen für diese hypothetischen Stufen haben keine nomenklatorische
Gültigkeit.
Das hier vorgeschlageneSchema zeigt die Evolution als ein ununterbrochenes
Kontinuum. Es erschöpfende Definition der verschiedenen taxonomischen
ist nicht als
Gruppen gedacht, obwohl manche Familien weitgehend gekennzeichnet sind.
Als eines der grundlegendsten Merkmale der Pulmonata wird die Einschnürung der
Mantelhöhle angesehen, durch die sie in eine apikale Lungenhöhle und eine mit ihr durch
einen Pneumostom verbundene aperturale Hypopeplarhöhle geteilt wird. Dieser
Mechanismus hat sich nach 2 verschiedenen Richtungen hin entwickelt; nämlich: 1.
Zu einem Zweig mit kontraktilem Pneumostom, bei dem die Lungenhöhle ausschliesslich
der Luftatmung dient. 2. Zu einem Zweig mit nicht-kontraktilem Pneumostom, bei
dem die Lungenhöhle entweder nur für die Aufnahme von Wasser, oder von Wasser und
Luft, nie aber nur für Luft allein benutzt wird. Nur dieser zweite Typus kommt bei
Chilina und den höheren limnischen Basommafophoren vor. Durch die Entstehung einer
pulmonären Lamelle, sowie mehrerer pulmonärer Anhängsel wurde diese Form weiter
modifiziert. Es wird dargelegt, dass die Pseudobranchie und der Sipho der höheren
SOsswasserschnecken niemals verwachsen, sondern immer deutlich gesondert und nicht
miteinander homolog sind. Die Frage der Homologie des Syphons und der Pseudo-
branchie mit dem Pneumostomal-Lappen der niederen Pulmonaten bleibt noch ungeklärt.
Verlust und Verwandlung dieser Organe bei den höheren limnischen Basommatophoren
wird besprochen.
Andere hier erörterte Merkmale sind: Heterostrophie, Hyperstrophieund invertierte
Symmetrie: Rückbildung der Schale zur Napfform; Stellung des Zerebralnervenrings
vor dem Bukkaiapparat, sowie Verengung des Viszeral- und Zerebralnervenrings:
Stellung der Augen und Gestalt der Tentakeln; Veränderung des Typs der im Bindegewebe
befindlichen Kalkkörner; Vorhandensein von Haemoglobin; apikale Punktierung der
Schale; Zähnung des Schalenmundes; Anordnung der Intestinalschleifen; Anwesenheit
eines gastrischen Caecums: Anzahl der Leberlappen: verschiedene Merkmale der
Radula; Vorhandensein eir r äusseren Genitalrinne; das Ausmass der Trennung des
384 H. W. HARRY
männlichen und weiblichen Geschlechtsganges unterhalb der EiweissdrOse; Verlust der
vesicula seminalis vaginae; Typus des Sammelraumes der ZwitterdrOse und Ansatz
des Zwitterganges in Bezug darauf, sowie örtliche Beziehung der ZwitterdrOse zum
oberen Leberlappen.
RESUME
L'ANATOMIE DE CHJLJNA FLUCTUOSA RÉEXAMINÉE, ET EBAUCHE D'UNE
PHYLOGÉNIE DES BASOMMATOPHORES LIMNIQUES SUPÉRIEURS
(GASTROPODA: PULMONATA)
Un nouvel examen de l'anatomie grossière de cette espèce a montré un certain

nombre de caractères qui n'avaient pas encore été signalés chez les chilinidés, tels
que la présence d'un pan post-tentaculaire, de grains de sable dans l'estomac, d'un
spermatophore, de la nature adnée du muscle rétracteur columellaire, d'un tégument
tesselé de la masse céphalopédieuse et de granules calcaires dans les tissus conjonctif s
ainsi que dans la lumière du vagin.
L'anatomie de Chilina est ensuite comparée à celle des Basommatophores limniques
supérieurs et, dans une plus faible mesure aussi à celle des autres groupes majeurs
de Pulmones. Cette comparaison est suivie d'une ébauche préliminaire de Phylogenese
reliant Chilina et les Basommatophores supérieurs limniques, dans laquelle chaque
stade de leur évolution est défini. L'esquisse commence arbitrairement au niveau
"Pulmoné Primitif et suggère progressivement quels caractères furent ajoutés,
modifiés ou perdus par rapport au moment ou d'autres caractères subirent des
transformations semblables. Des traits déterminants multiples sont donnés pour
chacun des divers stades hypothétiques, dont dérivent une ou plusieures familles
survivantes. Les désignations de ces stades hypothéthiques n'ont aucune valeur de
nomenclature.
L'esquisse montre l'évolution sous forme de continuité ininterrompue. Elle ne veut
nullement définir complètement tous les groupes taxonomiques, quoique certaines
familles se trouvent amplement définies.
La constriction de la cavité du manteau, qui la divise en une cavité apicale pulmonaire
et une cavité aperturale hypopéplaire, reliées l'une à l'autre par un pneumostome, est
tenue être un des caractères les plus fondamentaux des Pulmones. Ce mécanisme
évolua ensuite en 2 directions: 1) type à pneumostome contractile, où la cavité pulmo-
naire sert exclusivement à la respiration aérienne; 2) type à pneumostome non-con-
tractile, où la cavitépulmonairesert soit à contenir exclusivement de l'eau, soit de l'eau
et de l'air, mais jamais exclusivement de l'air. Seul le second type se trouve chez Chilina
et chez les Basommatophores limniques supérieurs, où il a été modifié par l'addition
d'une lamelle pulmonaire et de quelques appendices pulmonaires. L'auteur suggère que
la pseudobranchie et le siphon des Basommatophores limniques supérieurs sont des
entités toujours distinctes, jamais fusionées, qui ne sont jamais homologues l'une à
l'autre. La question de l'homologie du siphon et de la pseudobranchie avec le pan
pneumostomal des pulmones inférieurs reste ouverte. La perte et la modification
de ces organes dans les Basommatophores limniques supérieurs est discutée.
Les autres caractères pris en considération dans la phylogènie sont: l'hété-
rostrophie, l 'hyper strophie et la Symmetrie inversée; retour de la coquille vers le
type patelliforme. Le déplacement du collier nerveux en arrière de la masse buccale
et la constriction du collier nerveux cérébral et de l'anse viscérale; la position de
l'oeil et la forme des tentacules; la modification du type des granules calcaires du
tissu conjonctif; présence d'hémoglobine; ponctuation apicale de la coquille; denticu-
lation de la coquille; répartition des boucles intestinales; présence d'un caecum
gastrique; nombre de lobes majeurs du foie; divers caractères radulaires; disparition
du sillon génital externe; degré de séparation des tubes génitaux mâle et femelle
audessous de la glande de l'albumine; perte de la vésicule séminale du vagin; le type
de l'atrium de la gonade et le point d'insertion du conduit hermaphrodite par rapport
à lui; et les relations topographiques de la gonade et du lobe supérieur du foie.
RESUMEN
REVISION ANATÓMICA DE CHILINA FLUCTUOSA (GRAY), CON PROLEGÓMENO
SOBRE LA FILOGENIA DE LOS BASOMATOFOROS DULCEACUICOLAS
SUPERIORES (GASTROPODA: PULMONATA)
Un reexamen de la anatomía de esta especie reveló algunas características hasta

ahora no indicadas para los Chilinidae, como el lóbulo post-tentacular, la presencia de
granos de arena en el estómago, el espermatóforo, la condición adnata del músculo
retractor columelar, el integumento teselado de la masa ceíalopedal,
y granulos
calcáreos en el tejido conectivo y el lumen vaginal.
Se compara la anatomía de Chilina con aquella de los Basommatophora de agua
dulce más avanzados e incidentalmente con otros grupos mayores de pulmonados. A
esta comparación sigue un esbozo preliminar de la filogenia de Chilina y Basommato-
phora dulceacuícolas superiores, en los cuales cada estado evolutivo es definido. El
esquema comienza, arbitrariamente, al nivel del "Pulmonado Primitivo", sugeriendo
después los caracteres agregados, modificados o perdidos, en relación al tiempo en
que otros caracteres experimentaron cambios similares. Múltiples caracteres se
consideran peculiares a cada uno de los varios estadios hipotéticos que dieron origen a
una o más familias sobrevivientes. Estos estadios hipotéticos están denominados de
una manera que no tiene validez nomenclátor ial.
El esquema muestra la evolución como un continuo ininterrumpido, sin intentar
una completa definición de los taxa, aunque algunas familias están ampliamente definidas.
La constricción de la cavidad del manto para formar una cavidad pulmonar apical y
una cavidad hipopeplar apertural, interconectadas por el neumóstoma, se considera
una de las características más fundamentales de los pulmonados. Este mecanismo
luego evolucionó en dos direcciones diferentes: una con neumóstoma contráctil, en la
cual la cavidad pulmonar es usada exclusivamente para la respiración aérea; otra
con neumóstoma no contráctil, en la cual la cavidad pulmonar es usada o exclusivamente
para retener agua, o agua y aire, pero nunca exclusivamente aire. Sólo el segundo
tipo ocurre en Chilina y los Basommatophora de agua dulce. El ha sido modificado
por la adición de una lámela pulmonar y varios apéndices neumostomales. Se sugiere
que la seudobranquia y el sifón de los Basommatophora superiores de agua dulce son
siempre entidades distintas, nunca fusionadas, y no homologas entre sí. Pero la
homología del sifón y de la seudobranquia con el lóbulo neumostomal de los pulmonados
inferiores es cuestión aún no decidida. Se discute, aquí, la pérdida y modificación
de éstos en los Basommatophora superiores de agua dulce.
Otos caracteres considerados en la filogenia son la heterostrofía, hiperestrofía
y simetría invertida; la reversión a formas pateloides: el movimiento del anillo
nervioso cerebral, posterior a la masa bucal, y la constricción de los anillos nerviosos
visceral y cerebral; la posición del ojo y forma de los tentáculos; la modificación
del tipo de granulos calcáreos del tejido conectivo; la presencia de hemoglobina; la
punctuación apical de la concha; la dentición de la concha; la distribución de las
vueltas intestinales; la presencia de un ciego gástrico; el número de lóbulos hepáticos
mayores; algunos caracteres radulares; la desaparición del surco reproductivo externo;
el grado de separación de los conductos reproductores masculino y femenino debajo
la glándula albuminígera; la pérdida de la vesícula seminal; el tipo de atrio gonadal
y la inserción del ducto hermaírodita en relación a aquel, y la relación topográfica
de la gonada y lóbulo superior del hígado.
.
CYTOLOGIC AL STUDIES OF PLANORBmAE (GASTROPODA: BASOMMATOPHORA).

I. THE AFRICAN SUBGENUS BULINUS S.S.
J. B. BURCH
Museum of Zoology, University of Michigan
and
The Liberian Institute
The American Foundation for Tropical Medicine
Harbel, Liberia
ABSTRACT
The gastropod genus Bulinus medical importance because various of its
is of
members are the vectors of human urinary schistosomiasis
in Africa and the
Near East. The cytology of its species is of interest because polyploidy occurs
among them. The subgenus ßuZmMSs. s (here understood to comprise species
.
of the "tropicus" and "truncatus groups") has a geographical distribution which,

in general, covers the entire African continent. Species of the more northern
"truncatus group" serve as intermediate hosts for Schistosoma haematobium;
the more southerly located "tropicus group" does not.
Bulinus truncatus truncatus, B. truncatus rohlfsi and B. coulboisi, all
belonging to the "truncatus group", have 36 pairs of chromosomes. B. sericinus,
also of the "truncatus group", has 72 pairs of chromosomes. These species and
subspecies are considered to be polyploid since they contain exact multiples of
the number of chromosomes (n=18) usually found in most other species of the
family and in the "tropicus group"'(B. tropicus tropicus, B. tropicus angolensis,
B. tropicus zanzebaricus;,B. natalensis has mostly 19 elements present at
meiotic metaphase I, but some cells have 20 and 21). Coupled with these high
polyploid chromosome numbers is a susceptibility to infection with S. haema-
tobium.
In critical geographical areas where the "truncatus "and "tropicus groups"
overlap,chromosome numbers may proveto serve as differentiating criteria for
workers trying to identify the species. Chromosome number determinations may
also help in discerning which field populations are capable of transmitting
schistosomiasis
INTRODUCTION region and the Near East, and Indo-

planorbis, restricted to the Indian region.
The Bulininae comprise a group of The genus Bulinus is of cytological sig-
freshwater snails of the Basommatophoran nificance because it contains species which
family Planorbidae. The subfamily, as undoubtedly have had polyploid origins
presently understood, comprises two (Burch, 1960a). The genus is of medical
genera: /5, occurring in the African importance because various of its
""Contribution No. 1, Intermediate Hosts of Schistosomiasis Program, Institute of Malacology.
^This investigation was supported (in part) by a Public Health Service research career program
award (number 1-K3-AI-19, 451-01) and by research grants 5 Tl AI 41-05 and E-2409, from the
National Institute of Allergy and Infectious Diseases, U. S. Public Health Service, and GB 787
from the National Science Foundation, Washington, D. C, U. S. A.
(387)
388 J. . BURCH
^.
-^
^
f
z
^ truncatus
^7 truncatus forms
transversalis
trigonus
coulboisi •
Truncatus group
^ ccr
rohlfsi
nyassanus o.
mutandaensis
*See Mandahl-Barth (1960).
FIG. 1. Geographic distribution of the "truncatus group" of Bulinus s.s. in Africa (from Mandahl-Barth,
1958, 1960). The actual ranges of most of the species and subspecies are probably much greater
than shown here.
CYTOLOGY OF BULINUS 389
members are the intermediate hosts of Africanus group {=Physopsis)

schistosomiasis haematobia in Africa and
the Near East. These snails also transmit B. africanus africanus (Krauss)
other schistosomes of the haematobium B. africanus ovoideus (Bourguignat)
group, infecting herbivores and man. B. abyssinicus (Martens)
Cytology of its species has a special B. nasutus nasutus (Martens)
interest from the standpoint of medical B. nasutus productus Mandahl-Barth
zoology since there may exist a possible B. ugandae (Mandahl-Barth)
connection between polyploidy and sus- B. globosus (Morelet)
ceptibility to infection with schistosomes B. jousseaumei (Dautzenberg)
(Burch, 1960a).
In an attempt to assess whether or not Tropicus group
any correlation exists between sus-
ceptibility and chromosome number within B. tropicus tropicus (Krauss)
a systematic group of closely related B. tropicus angolensis (Morelet)
species, I have had the opportunity to B. tropicus zanzebaricus (Clessin)
study 8 nominal species and subspecies B. tropicus toroensis Mandahl-Barth
of the bulinine subgenus Bulinus s.s. (i.e., B. tropicus alhmudi (Dautzenberg)
the "truncatus" and "tropicus species B. liratus (Tristram)
groups" of Mandahl-Barth, 1958, 1960) B. guemei (Dautzenberg)
from the African continent. The results
of these studies are presented in this Truncatus group
paper.
B, trans versalis (Martens)
SYST EM ATICS . truncatus truncatus (Audouin)
B. truncatus sericinus (Jickeli)
Bulinus is a widespread genus of many B. truncatus trigonus (Martens)
nominal species (over a hundred) dis- B. truncatus rohlfsi (Clessin)
tributed largely over the entire continent B. mutandaensis (Preston)
of Africa. Its many named "species* B. coulboisi (Bourguignat)
have resulted from the perplexing mor- ß. nyassanus (Smith)
phological variability found within and
among the various populations, a trait Forskalii group
common to all basommatophoran genera
of wide distribution. The latest mono- B. camerunensis Mandahl-Barth
graphic study of the genus, that of Mandahl- B. reticulatus Mandahl-Barth
Barth (1958, emended in 1960), recognizes B. cemicus (Morelet)
only twenty species and nine subspecies. B. forskalii (Ehrenberg)
The various species of the genus ßwiznits . scalaris (Dunker)
have traditionally been grouped into three . senegalensis Müller
taxa (lately referred to as subgenera,
formerly as genera): Bulinus s.S., Mandahl-Barth considers the "africanus
Physopsis and Pyrgophysa. This nomen- group" to correspond to subgeneric rank
clatorial system has been criticized by (subgenus Physopsis) and the other three
Mandahl-Barth (1958), who instead divides "groups" to comprise the subgenus
the genus Bulinus s.l. into four species Bulinus s.S. In spite of intensive research
"groups" based on shape and sculpture of and numerous publications "...species
the shell and on some anatomical discrimination within Bulinus is extremely
structures. His classification as modi- difficult, and it is inevitable that further
fied by him in 1960 is as follows: studies will change the number of ac-
390 J. . BURCH
tropicus
angolensis
zanzeoaricus
toroensis ' Tropicus group
allvaudi
c^
liratus
guemei
FIG. 2. Geographic distribution of the "tropicus group" of Bulinus s.s. in Africa and Madagascar (from
Mandahl-Barth. 1958, 1960). The actual ranges of most of the species and subspecies are probably
much greater than shown here.
knowledged species and subspecies" while its related, more southerly, species
(Mandahl-Barth, 1958: 59). do not normally carry S. haematobium,
H. J. Walter (personal communication) infection.
of the American Foundation for Tropical
Medicine, after an intensive conchologi- MATERIALS AND METHODS
cal-anatomical study of African bulinine
snails, concludes that the "tropicus* and The species studied in this investigation
"truncatus groups" together form a are listed below. Shells of duplicate
distinct single group in the genus that specimens, when available, have been
can be defined both conchologically and deposited in the collections of the Museum
anatomically. He considers the older of Zoology, University of Michigan.
taxonomic practice of 3 subgenera more
convenient (and more accurate bio- Truncatus Group
logically, not nomenclatorially), with
if
Bulinus s.S. containing the "tropicus" and Bulinus truncatus truncatus (Audouin).
"truncatus groups", the subgenus Pyrgo- Sardinia. From live stocks at the
physa equal to the "forskalii group", and London School of Hygiene and Tropical
the subgenus Physopsis equal to the Medicine. P. L. LeRoux, December 14,
"africanus group" (Walter, 1962). 1961. Four animals studied.
Although information on susceptibility Teheran, Iran. From live stocks at
of bulinine snails to schistosome infection the British Museum (Natural History).
is incomplete, it seems that, in general, C. A. Wright, September, 1962. Two
members of the "africanus group" animals studied.
commonly act as the hosts of Schistosoma Baghdad, Iraq. From live stocks at
haematobium and its allies south of the the London School of Hygiene and
Sahara, while members of the "truncatus Tropical Medicine. P. L. LeRoux,
group" serve as intermediate hosts in December 14, 1961. Five animals
their distribution area (Fig. 1), i.e., in studied.
northern and northeastern Africa and the Mena area, Giza province, Egypt.
adjacent Mediterranean and Near Eastern Collected by Fikry el Tawil, March 9,
regions, where the "africanus group" does 1961. Four animals studied.
not occur. Members of the "tropicus Sudan. From live stocks at the
group", which has a southern distribution London School Hygiene and Tropical
of
(Fig. 2), do not usually serve as inter- Medicine. P. L. LeRoux, December 14,
mediate hosts. Only a few members of 1961. Four animals studied.
the "forskalii group", in widely separated
Bulinus truncatus sericinus (Jickeli).
areas, transmit S. fiaematobium. The
West Aden Protectorate. From live
snails serving as intermediate hosts are
stocks at the British Museum (Natural
usually more susceptible to the regional
History). C. A. Wright, September,
schistosome strains and may be com- The
1962. Three animals studied.
pletely refractive to schistosome strains
specimens which Jickeli (1874, Nova
from a remote region. Acta Leop. Carol., 37: 194) named
If the subgenus Bulinus s.s.
bulinine
"Isodora" sericina came from Mekerka
is accepted in the traditional sense (and
at Toquor River, Ethiopia. It is possible
that of Walter, 1962) as forming a distinct
that the Aden specimens used in my
morphological unit (="tropicus" + "trun-
studies are not the same species as
catus groups"), then this taxon has a geo-
that described byJickelisince they came
graphical distribution which extends over
from a remote, isolated population,
the entire length of Africa (Fig. 3). But long separated from Africa.
only in northern Africa, including the
Siidan, does it serve as an intermediate Bulinus "sp." (truncatus "? sericinus"
host for human urinary schistosomiasis. (Jickeli)). Awasa, Ethiopia. From live
392 J. . BURCH
Truncatus group
Bulinus s.s
Tropicus group
FIG. 3. Probable generalized distribution of the "truncatus" and "tropicus groups" in Africa, Note the
large area of probable distributional overlap (cross-hatched).
stocks at the British Museum (Natural seem doubtful and its chromosome
History). C. A. Wright, September, number is closer to that found in species
1962. Two animals studied. of the "tropicus group*.
Bulinus truncatus rohlfsi (Clessin). Bulinus {"? natalensis"

(Küster)). Uni-
Vicinity of Kumasi (Ashanti), Ghana, versity College ponds, Salisbury,
West Africa. Collected by F. Wick- Southern Rhodesia. Collected by A. D.
remasinghe. May, 1961. Two animals Harrison, January 25, 1963. Two
studied, animals studied.
Bulinus coulboisi (Bourguignat). Bulinus s.S. "sp.". Kisumu, Lake Victoria,
Mwanza, Tanganyika. Collected by John Kenya. From live stocks at the London
McClelland, May, 1961. Identified by School of Hygiene and Tropical
Dr. G. Mandahl-Barth. Two animals Medicine. P. L. LeRoux, December
studied. 14, 1961. Eight animals studied.
Tropicus Group
The material examined consisted of
ovotestes in active stages of gameto-
Bulinus tropicus tropicus (Kraus s).
genesis. The tissues were fixed and
De Villiers cement reservoir, Malelane,
preserved in either Carnoy's (1887)
South Africa. Collected by H. J.
(acetic-ethanol-chloroform, or
1:6:3)
Schutte, July, 1961.
Newcomer's (1953) fixatives and stained
Small tributary of Gwebi River on
by the acetic-orcein squash technique
New England Farm, near Salisbury,
(La Cour, 1941). Observations were made
Southern Rhodesia. Collected by A. D.
with a Nippon Kogaku microscope using
Harrison, January 25, 1963. Ten
a lOOX (n.a. 1.25) oil immersion objective
animals studied.
and 10-30X oculars. The chromosomes
Bulinus tropicus angolensis (Morelet). were drawn with the aid of a camera
Northern Rhodesia. From live stocks lucida and reproduced at a table-top
at the London School of Hygiene and magnification of 3450X or 5400X. Photo-
Tropical Medicine. P. L. LeRoux, graphs were taken using a 20X ocular,
December 14, 1961. Six animals studied. an oil immersion objective, a Kodak
Wratten 57A (green) filter, and Kodak High
Bulinus tropicus zanzebaricus (Clessin).
Contrast Copy (Micro-File) film.
Mwanza, Tanganyika. From live stocks
at the London School of Hygiene and
OBSERVATIONS AND DISCUSSION
Tropical Medicine. P. L. LeRoux,
December 14, 1961. Six animals studied. Chromosome numbers of species studied
Bulinus natalensis (Küster). Lake shown in Table I.
in this investigation are
Mcllwaine, Southern Rhodesia. Col- Bulinus truncatus truncatus consistently
lected by A. D. Harrison, January 27, has the haploid chromosome number 36
1963. Four animals studied. "Physa" throughout the part of its range so far
natalensis KOster was listed as a sampled (i.e., Sardinia, Iran, Iraq, Egypt
synonym of B. tropicus tropicus by and Sudan). In addition, its subspecies
Mandahl-Barth (1958). However, B. truncatus rohlfsi from Ghana also has
Mandahl-Barth now places natalensis in this haploid number. However, the so-
the "truncatus group" (according to A. called "subspecies" sericinus^ from the
D. Harrison, personal communication). Western Aden Protectorate, although also
It is placed in the "tropicus group" in polyploid, has the haploid number 72,
the present paper because its affinities indicating that it is not a subspecies of
^See p 391.
394 J. . BURCH
B. truncatus, but is most probably repro- ("? sericinus") has not as yet been
ductively isolated from it. Its polyploid implicated in the transmission of schisto-
(octaploid) number indicates that Mandahl- somiasis; however, one specimen from
Barth (1960) correctly removed it from the sericinus sample (n=72) from the
the "tropicus group" and included it in Western Aden Protectorate shed schisto-
the "truncatus group". On the other some cercariae which produced infections
hand, the specimens from Awasa, Ethiopia with female worms in hamsters. Since
tentatively identified by C. A. Wright as Schistosoma haematobium was known in
Bulinus "sp." {truncatus "? sericinus") the particular valley where the sample
(n=18) is neither truncatus nor sericinus was collected, while bovine schisto-
(as understood above), but reproductively somiasis was not, he thinks it likely
isolated from both because of the great that his female worms were S. haema-
differences in chromosome numbers (e.g., tobium.
a hybridization between "? sericinus"
. Bulinus coulboisi from Tanganyika,
and B. t. truncatus, i.e., a diploid and a placed by Mandahl-Barth (1958) in the
tetraploid, would result in infertile tri- "truncatus group", also has 36 pairs of
ploids). Moreover, its haploid chromo- chromosomes, indicating that this number
some number 18 indicates that its affinities of chromosomes is not restricted to B.
are probably with the "tropicus group" truncatus, but occurs in at least one
rather than the "truncatus group". other species as well. It is perhaps
Anatomical andparasitological evidence significant to note that B. coulboisi has

seems to bear out these apparent relation- been infected experimentally with,Sc/2¿sío-
ships based on cytological observations. soma haematobium.
Bulinus sp. ( "?ser¿c¿nits'')(n=18) appears Bulinus tropicus tropicus from South
to be related on anatomical grounds to Africa and Southern Rhodesia, B. tropicus
B. tropicus (D. S. Brown, personal com- angolensis from Northern Rhodesia, and
munication). Further, according to C. A. B. tropicus zanzebaricus îromTasiganyika.
Wright (personal communication), B. sp. all have the haploid chromosome number
^
TABLE I. Chromosome Numbers of Bulinus s. s.
Species
396 J. . BURCH
12 » •
I
13
^ 14
18. In B. tropicus "? natalensis" from with Schistosoma haematobium, the other
Southern Rhodesia most of the Metaphase is not. Because of the apparent complex
I cells have 18 chromosomal elements, overlapping variation of morphological
but some cells have 19. On the other characters exhibited by both species
hand, in B. tropicus natalensis from groups, specialists have yet to uncover
Southern Rhodesia most Metaphase I cells clear-cut characters on which to make
have 19 chromosomal elements, but some specific determinations (regardless of the
cells have 20 and 21. These varying many publications on these snails during
chromosome numbers may be the result the past 10 years). And many populations
of hybrid origins of the two populations. are almost impossible to assign to one
Nevertheless, these anomalous numbers species group or the other. By using
are not the result of polyploidy since chromosome numbers as separating
they do not approach the 36 pairs of criteria, perhaps other more gross
chromosomes found so far in members differentiating characters can be foiind
of the "truncatus group". None of the which will prove of use for easily
members of the "tropicus group" have separating species of the two groups in
been implicated as transmitting schisto- critical areas. Chromosome number
somiasis on the continent, although B. determinations may therefore prove of
liratus appears to be the intermediate value in helping solve someof the complex
host on Madagascar. taxonomic problems of the snail vectors.
It appears therefore that the subgenus Such chromosome number determinations
Bulinus s.S. (as here understood) is indeed might also prove helpful in discerning
made up of two species "groups" (ana- which field populations of Bulinus s.s.
tomically closely related to each other, are capable of transmitting schisto-
and more distantly related to Physopsis somiasis and which populations are not.
and Pyrgophysa). Members of one group
have 18 pairs of chromosomes (excepting ACKNOWLEDGEMENTS
the 19-21 Metaphase I elements in
natalensis), the number most usually found It is a priviledge to acknowledge the
for members of its family (Burch, 1960b); generosity of the late Prof. P. L. LeRoux
the other group has species with exact of the London School of Hygiene and
multiples of this number (n=36 or 72). Tropical Medicine, who gave me the
The group with the higher chromosome specimens on which a large part of this
numbers is implicated in the transmission paper is based. Grateful acknowledgement
of human urinary schistosomiasis; the is also made to Anne Gismann for
group with the lower numbers is not. arranging to get many of the other
Polyploidy, per se, may not be directly specimens, and for helpful suggestions
responsible for susceptibility to schisto- concerning the manuscript; to Henry
some infection in the "truncatus group", van der Schalle for allowing me to study
but it is strongly suggestive that poly- snails from his laboratory stocks; to
ploidy was a key factor in the origin of C. M. Patterson for technical assistance;
this group. Coupled with this, either to G. Mandahl-Barth for identifying
originally or evolving through the group's Bulinus coulboisi; and to the following
reproductive isolation from the "tropicus colleagues for sending me cytological
group", seem to be physiological charac- material: A. D. Harrison, John Mc-
ters allowing for schistosome infection. Clelland, H. J. Schutte, Fikry elTawil,
The practical implications of these cyto- F. Wickremasinghe, and .
A. Wright.
logical findings are clear. In regions of
geographical overlap (Fig. 3), it is ex- REFERENCES
ceedingly difficult for workers to differ-
entiate the two species groups from each BURCH, J. ., 1960a, Chromosome
other, but one is capable of being infected numbers of schistosome vector snails.
398 J. . BURCH
^^ ft
23 Si.
26 ^
24
28
• -
27
#•
30
• ^
^ ir
32
3. /
FIGS. 23-32. Camera lucida drawings of chromosomes of ßu/imis s.S. ("tropicus group"). FIG. 23. Bulinus
tropicus tropicus, Malelane, o' diakinesis. FIG. 24. Bulinus sp., Lake Victoria, '
diakinesis. This is a
drawing of the chromosomes shown in Fig. 8. FIG. 25. B. tropicus tropicus, Malelane, diakinesis (note
=
heterochromatic areas). FIG. 26. ß. tropicus zanzebaricus, í diakinesis (note heterochromatic areas). This
is a drawing of the chromosomes shown in Fig. 10. FIGS. 27, 28. Bulinus sp.. Lake Victoria. Fig. 27. /
Metaphase I. This is a drawing of the chromosomes shown in Fig. 7. Fig. 28. ' Metaphase U. This is a
drawing of the chromosomes shown in Fig. 6. FIG. 29. B. natalensis, Lake Mcllwaine, spermatogonial
anaphase-telophase; 35 chromosomes are at one pole, 37 at the other. FIG. 30. B. " ? natalensis" Salisbury, ,
o" diakinesis; the upper (n=18) and lower (n=19) cells were found side by side in one individual. FIG. 31. B.
tropicus angolensis, diakinesis.
.-'
This is a drawing of the chromosomes shown in Fig. 9. FIG. 32. B.
natalensis, o' diakinesis.
Z. Tropenmed. Parasit., 11(4): 449-452. 1-89, pis. 1-40, reprinted from Bull.
, 1960b. Chromosome studies of Wld. Hlth. Org. 1957, 16: 1103-1163 and
aquatic pulmonate snails. Nucleus, 3(2): 17: 1-65.
177-208. ,
1960, Intermediate hosts of
CARNOY, H. ., 1887, Conférence donnée Schistosoma in Africa. Some recent
â la société belge de Microscopie. information. Bull. Wld. Hlth. Org.,
Appendice: I. Les Globlules polaires 22: 565-573.
de V Ascaris clavata. La Cellule, Rec. NEWCOMER, E. H., 1953, A new
Cytol. Histol. Gén., 3(2/3): 227-273. cytological and histological fixing fluid.
LA COUR, L., 1941, Acetic-orcein: A Science, 118(3058): 161.
new stain -fixative for chromosomes. WALTER, H. J., 1962, Punctation of the
Stain Techn., 16: 169-174. embryonic shell of Bulininae (Planor-
MANDAHL-BARTH, G., 1958, Inter- bidae) and some other Basommatophora
mediate hosts of Schistosoma. African and its possible taxonomi -phylogenetic
Biomphalaria and Bulinus. Wld. Hlth. implications. Malacologia, 1(1): 115-
Org. Monogr, Ser. No. 37, Geneva. 137.
ZUSAMMENFASSUNG
ZYTOLOGISCHE STUDIEN ÜBER PLANORBmAE (GASTROPODA: BASOMMATOPHORA)
I. DIE AFRIKANISCHE UNTERGATTUNG BULINUS S. S.
Die Gastropodengattung Bulinus ist von medizinischer Bedeutung weil mehrere ihrer
Mitglieder die Überträger der Blasenbilharziose in Afrika und dem Mittleren Osten
sind. Wegen der Polyploidie einiger ihrer Arten ist sie zytologisch von Interesse. Die
geographische Verbreitung der Untergattung Bulinus s.S. (hier auf die Mitglieder der
"tropicus" und "truncatus" Gruppen beschränkt) erstreckt sich im Grossen und Ganzen
über den gesamten afrikanischen Kontinent. Arten der mehr nördlichen "truncatus"
Gruppe dienen als Zwischenwirte von Schistosoma haematobium, nicht aber die mehr
südliche "tropicus" Gruppe.
Je 4 Arten beider Gruppen, aus weit voneinander entfernten Gegenden wurden
zytologisch untersucht. Bulinus truncatus truncatus, B. truncatus rohlfsi and .
coulboisi, aus der "truncatus" Gruppe, haben 36 Chromosomenpaare; B. sericinus,
auch aus dieser Gruppe, hat deren 72. Diese Arten und Unterarten werden als polyploid
angesehen, weil ihre Chromosomenzahl genau das Vielfache derjenigen ist (n=18), die
gewöhnlich bei den meisten anderen Arten der Familie, sowie in der "tropicus Gruppe"
gefunden wird, d.h. bei B. tropicus tropicus, .
tropicus angolensis und . tropicus
zanzebaricus; .
natalensis hingegen zeigt in der meiotischen Metaphase I meistens
19 Elemente, und manche Zellen sogar 20 und 21 (überzählige Chromosome). Zusammen
mit den hohen polyploiden Chromosomenzahlen findet man also Empfänglichkeit für
S. haematobium Infektionen.
Bei Unter Scheidungsschwierigkeiten in Arten aus Gebieten wo die "truncatus" und
"tropicus" Gruppen sich Ubersctineiden, wäre es möglich die Chromosomenzahlen als
Kriterium für die Artzugehörigkeit heranzuziehen, sowie auch um festzustellen welche
der Populationen im Freiland die Bilharziose zu übertragen vermögen.
RESUME
ETUDES CYTOLOGIQUES SUR LES PLANORBIDAE

(GASTROPODA: BASOMMATOPHORA) I. LESOUS-GENRE AFRICAINBÍ/LWt/S S. S.
Le genre gastéropode Bulinus est d'importance médicale parceque plusieurs de ses

membres sont vecteurs de la bilharziose vésicale en Afrique et au Moyen Orient. Le
présence de Polyploidie parmi ses espèces le rend cytologiquement intéressant. Le
sous-genre Bulinus s.S. (ici compris comme groupant les espèces des groupes "tropicus"
et "truncatus") est réparti, de manière générale, sur le continent Africain tout entier.
Les espèces du groupe "truncatus", plutôt septentrional servent d'hôtes intermédiaires
à Schistosoma haematobium, mais non le groupe "tropicus" plus méridional.
400 J- - BURCH
Quatre espèces de chacun de ces groupes, provenant de regions géographiques
éloignées, ont été examinées cytologiquement. Bulinus truncatus truncatus, B. truncatiis
rohlfsi etß. cou/bois», appartenent au groupe "truncatus", ont 36 paires de chromosomes;
B. sericinus, du même groupe, en a 72. Ces espèces et sous-espèces sont considérées
comme polloides, parceque leurs nombres de chromosomes sont des multiples exacts
de celui (n=18) trouvé chez presque toutes les espèces de la famille et aussi chez le
groupe "tropicus", c'est à dire chez B. tropicus, . .
tropicus angolensis et tropicus
zanzebaricus; . natalensis, cependant, a d'habitude 19 éléments à la métaphase
meiotique I, et dans quelques cellules même 20 et 21 (chromosomes surnuméraires).
Nous voyons donc associée aux nombres polyploides élevés une susceptibilité à l'infection
par S. haematobium.
En cas de difficultés d'identification dans certains territoires où les groles "tropicus"
et "tnmcatus" se chevauchent, une détermination du nombre chromosomique pourrait
peut-être servir de critère différentiel, et, de même, à juger quelles populations
naturelles sont capables de transmettre la bilharziose.
RESUMEN
ESTUDIOS CITOLOGICOS EN PLANORBIDAE (GASTROPODA: BASOMMATOPHORA).

I. EL SUBGÉNERO AFRICANO BULINUS S. S.
Los gastrópodos del género Bulinus tienen importancia médica por ser, varios de
sus miembros, trasmisores de la esquistosomiasis urinaria humana en África y en el
Cercano Oriente. La citología de sus especies resulta interesante por la ocurrencia de
poliploidia. El subgénero Bulinus s.s. (comprendiendo aquí las especies de los grupos
"tropicus" y "truncatus"), está en general distribuido por todo el continente africano.
Las especies del grupo más septentrional, "truncatus", sirven de huésped interme-
diario de Schistosoma haematobium, no así las del grupo "tropicus", más meridional.
BulimiS truncatiis truncatus, . truncatus rohlfst . coulboisi, pertenecientes al
primer grupo, tienen 36 pares de cromosomas. .sericimis, también del grupo
"trimcatus", tiene 72 pares. Estas especies e subespecies se consideran poliploideas
desde que contienen múltiples exactos del número de cromosomas (n=18) encontrado
habitualmente en la mayoria de las otras especies de la familia y en el grupo "tropicus"
(. tropicus tropicus, . tropicus angolensis, . tropicus zanzebaricus). .
natalensis
tiene en la mayoría 19 elementos en la metafase meiôtica I, pero algunas células tienen
20 y 21. En combinación con estos altos números poliploides de cromosomas, existe
la susceptibilidad a la infección por S. haematobium.
En áreas geográficas críticas, donde los grupos "truncatus" y "tropicus" se super-
ponen, el número de cromosomas puede servir como criterio diferencial para los
investigadores que tratan de identificar las especies. La determinación del número de
cromosomas puede también ayudar a discernir cuales poblaciones naturales son
capaces de trasmitir esquistosomiasis.
CHROMOSOMES OF SOME JAPANESE FRESHWATER SNAILS

(BASOMMATOPHORA: BRANCHIOPULMONATA) 1
J. B. Burch, J. E. Williams, Y. Hishinuma, R. Natarajan
406 Medical Laboratory

U. S. Army Medical Command, Japan^
and
Museum and Department of Zoology

University of Michigan, Ann Arbor, Michigan, U. S. A.
ABSTRACT
Chromosome numbers are reported for 10 species of Japanese freshwater
basommatophoran snails: 16 pairs of chromosomes were observed in Radix
ollula and 17 pairs in Radix japónica and R. onychia; 18 pairs of chromosomes
were found inFossaria trunca tula, Ph y sa acuta, Gyrauíus spirillus, G. perstri-
atulus, G. tokyoensis, Segmentina hemisphaerula and Camptoceras hirasei.
Two specimens of G. tokyoensis had supernumerary chromosomes, forming an
extra bivalent (n=19), or a bivalent plus a univalent (n=19+l). Since no trivalents
were found in any of the specimens of G. tokyoensis, we assume that the extra
chromosomes are not duplications of other chromosomes of the normal set. The
chromosome number of "Lymnaea" ollula is closer to that of the genus Radix
than to Fossaria, the genus to which it is usually assigned. This would seem to
corroborate Hubendick's (1951) anatomical findings.
The chromosomes of these Japanese species are monocentric and are there-
fore like those found in euthyneuran snails from other countries.
The most significant chromosome study 6 other species. It is on these
studies on freshwater basommatophoran studies that the present paper is based.

snails are those of Perrot and Perrot Grateful acknowledgement is made to
(1938), Le Calvez and Certain (1950), Dr. Tadashige Habe, National Science
Inaba (1953) and Burch (1960a, b; 1962). Museum, Tokyo; to Dr. Hiroshi Itagaki,
The only one of these authors to deal Azabu Veterinary College, Fuchinobe; to
with snails from the Japanese isles is Dr. Taizo Miura, Limnological Testing
Inaba, who describes the chromosomes of Station, Otsu; to Mr. Hikotaro Kajiyama,
4 species of freshwater Basommatophora Osaka; to Mr. Morihiko Tanaka, Nagoya;
and 2 species of marine Basommatophora. and to various members of the
We have recently had the opportunity to Department of Medical Zoology, 406
reinvestigate the 4 freshwater species Medical Laboratory, U. S. Army Medical
reported on by Inaba and, in addition, to Command, Japan, for help in securing
^This investigation was supported (in part) by a Public Health Service research career program
award (number 1-K3-AI-19, 451-01) and by research grants 5 Tl AI 41-05 from the National
Institute of Allergy and Infectious Diseases, U. S. Public Health Service, and G-21910
and GB
787 from the National Science Foundation, Washington, D. C., U. S. A. This work was
sponsored
(in part) by the Commission on Parasitic Diseases of the Armed
Forces Epidemiological Board
and was supported (in part) by the U. S. Army Medical Research and Development
Command.
Âddress: A. P. O. 343, San Francisco, California, U. S. A.
(403)
404 BURCH, WILLIAMS, HISHINUMA, NATARAJAN
the specimens used in these investigations. TABLE I. Japanese snails studied with col-
Appreciation is expressed to Dr. Akihiko lection sites.
Inaba, Mukaishima Marine Biological
Station and Dr. Tadashige Habe
Species
for reading the manuscript, and to Dr.
William J. Clench, Museum of Compara-
tive Zoology, Harvard University,
Cambridge, Massachusetts, U. S. A. for
checking our identifications of Physa
acuta. We wish to thank Professor Henry
van der Schalle, Museum of Zoology,
University of Michigan, Ann Arbor,
Michigan, U. S. A., and Major John W.
Moose, Chief, Department of Medical
Zoology, 406 Medical Laboratory, for
sponsoring these studies.
MATERIALS AND METHODS

Ten species of freshwater basommato-
phoran snails were used in this investi-
gation. All specimens were collected in
6 localities in Japan during 2 periods of
field studies, March, 1962 and August-
October, 1963. A list of species with
locality data is in Table I.
given
Duplicate specimens are in the collections
of the Museum of Zoology, University of
Michigan.
The material examined consisted of
ovotestes killed, fixed and preserved in
Newcomer's (1953) fluid. Cells of the
ovotestes were stained by the acetic-
orcein squash technique (La Cour, 1941).
All observations were on cells of
spermatogenesis and were made with
Nikon microscopes using lOOX (n.a. 1.25)
oil immersion objectives and 10, 20 and
25X oculars. All drawings of chromo-
somes were made with the aid of a camera
lucida and reproduced at a table top
magnification of 5100X, except for Figs.
6, 10, 15, 17, 25-27, and 38, which were
reproduced at a magnification of 4150X.
Photographs (Figs. 39-47) were taken
using a 20X ocular, oil immersion ob-
jective, a Kodak Wratten 57A (green)
filter, and Kodak High Contrast Copy film.
OBSERVATIONS AND DISCUSSION

Family Lymnaeidae
Previous reports which have been con-
CHROMOSOMES OF JAPANESE SNAILS 405
have 18 pairs 3. Fossaria parva and F. Radix ollula Gould. This is a very
modicella were reported to have 18 pairs common lymnaeid species in Japan and
of chromosomes and F. rustica to have certain other parts of the Orient.
19 pairs. Inaba and Tanaka reported The Japanese specimens have often been
"Lymnaea (Fossaria)" ollula pervia to referred to as Lymnaea (fiadix) ollula
have 17pairs of chromosomes and pervia (Martens), but Dr. J. P. E.
"Lymnaea (Radix)" japónica to have 18 Morrison (personal communication) of the
pairs, but recently, Dr. Inaba (personal U. S. National Museum has studied large
communication) has corrected these series of both "species" and he con-
numbers to 16 and 17 pairs respectively cludes that Limnaea ollula Gould 1859
for the 2 species. Chromosome numbers (type loc: Hong Kong) and Limnaeus
currently known for species of Radix pervius Martens 1867 (type loc: Tshi-fu,
and Fossaria are shown in Table II. Shantung, China) are one and the same
Radix japónica (Jay), This species is species and are therefore synonymous,
a very common aquatic snail in Japan; pervia not being even a subspecies of
14 specimens from 2 localities were ollula, as often cited in recent literature.
studied. Cells from 10 of the animals Most of our specimens seem to be nearly
were satisfactory for chromosome identical to the types from Yokohama,
studies. All 10 specimens had 17 pairs Japan of Lymnaea goodwini Smith 1876,
of chromosomes (Figs. 1-3, 13, 39), the
number found for most other species of
as figured by Hubendick (1951, Plate
Figs. 8-9). Hubendick (1951) considers
,
the genus (Table ). The chromosomes L. goodwini, as well as L. ollula and L.
of this species are monocentric (Fig. 3), pervia, to be synonyms oi Lymnaea viridis
as confirmed by Inaba (personal com- described by Quoy and Gaimard (1833)
munication), and not "dot-shaped" as from Guam.
previously reported. Highly contracted Some authors in the past have placed
mitotic metaphase chromosomes, such as Radix ollula with the genus Fossaria, but
reported previously in basommatophoran Hubendick (1951, p. 162) says that the shell
snails by Burch (1960a) are shown in Fig. 2. of L. viridis (= ollula?) "resembles the
Radix (Omia) onychia (Westerlund). form peregra of L. [=Radix] peregra as
This species is endemic to Lake Biwa, well as L. [=Fossaria]^ truncatula." He
Shiga Prefecture. Omiajaponica (Preston) states further that L. viridis {= R. ollula ?)
is a synonym of this species, which has "is not distinctly anatomically separated
also been placed in the genera Lithotis from L. [=Radix'\ auricularia (sensu lato).
and Erinna. Although the shape of some In L. [=Fossaria] truncatula, however, the
specimens of R. onychia might suggest male copulatory organ differs from that
that this species is only a variant of Ä. in viridis [=ollula?] by its relatively short
japónica (Jay), our observations on living and thick penis and penis sheath. The
and preserved specimens lead us to con- prostate of truncatula differs from that
clude that the two are definitely distinct of viridis [-ollula ?] by its comparatively
species. The anatomy of R. onychia has small fold." Thereiore," Lymnaea" ollula
been studied by Itagaki (1959). should perhaps be placed in the genus
We studied the ovotestes of 5 animals Radix instead of the genus Fossaria. Its
of Radix onychia, each with active stages chromosome number (n=16) is also closer
of spermatogenesis. All specimens had to Radix (n=17) than io Fossaria (n-18, 19).
17 pairs of chromosomes (Figs. 4, 14, Fifteen specimens of R. ollula were
15), which did not seem to differ signifi- studied from 2 localities, 11 of which
cantly from JR. japónica. yielded satisfactory cells (Figs. 5, 6, 16-
^Since all other species of Radix are have only 17 pairs of chromosomes, it seems de-
foxind to
sirable to confirm the report of 18 pairs of chromosomes in R. limosa. This species could have
been misidentified, or else it may have supernumerary chromosomes.
7 ^mt *
^
/
10
TABLE II. Chromosome numbers of Radix and Fossaria from different countries.
Species
^QQ
BURCH, WILLIAMS, HISHINUMA, NATARAJAN
>»nV >•-• vV^*•

15
16
9V^ 18 ^
19 ^# 20^ C»^ ^
FIGS. 13-21. Camera lucida drawings of late meiotic prophase (diakinesis) chromosomes of Japanese
Basommatophora.
FIG. 13. Diakinesis chromosomes of fíadí'x ja/)on¿ca. X3060.
FIGS. 14, 15.
FIGS. 16-18. Diakinesis chromosomes of :

Diakinesis chromosomes of Radix onychia. Fig. 14, X3060; Fig. 15, X2490.
oZ/wia. Fig. 16. Early diakinesis. Figs. 17,18. Late diakinesis.
FIG.
FIG.
19.
20.
Diakinesis oí Fossaria truncatula.
Diakinesis chromosomes of Physa acuta. X3060.
.
Fig. 46 is a photograph of the chromosomes in Fig. 17. Figs. 16, 18, X3060; Fig. 17, X2490.
chromosomes
FIG. 21. Diakinesis chromosomes of Gyraulus spirillus. X3060.

Family Physidae University. It is not known whether P.

heterostropha also occurs in Japan as
Previous reliable reports on the reported by Kuroda (1949).
chromosomes of physid snails are those
Burch and Bush Family Planorbidae
of Burch (1959, 60a, b),
(1960), Inaba (1953), and Le Calvez and Previous reports which have been con-
Certain (1950). All species (i.e., Physa sidered reliable on the chromosomes of
acuta, P. anatina, P. fontinalis, P.gyrina, planorbid snails are those of Bonham
P. heterostropha, P. sayii crassa, and (1955), Burch (1959, 60a, b, c, d, 61, 64),
Aplexa hypnorum) studiedby these authors Inaba (1953), Inaba and Tanaka (1953),
were found to have 18 pairs of chromo- Le Calvez and Certain (1950) and Natarajan
somes. An earlier study on the American (1960). These authors have studied 20
species Physa gyrina by Mahoney (1940) planorbid species, 12 of which have 18
was shown by Burch and Bush (1960) to be pairs of chromosomes (i.e.: Anisus
inaccurate. vortex, Armiger crista, Bulinus tropicus,
Physa acuta Draparnaud. This is Gyraulus deßectus, Helisoma anceps, H.
another species not native to Japan, but subscrenatum, H. trivolvis, Indoplanorbis
introduced from Europe. Eleven speci- exustus, Planorbarius comeus, Planor-
mens from 2 localities were studied, 7 bina glabrata, P. sudanica and Promenetus
of which yielded satisfactory cells. We exacuous); 2 species (ßulinus natalensis
found the chromosome numbers to be the and Planorbula crassilabris) hdive 19 pairs
same as those reported by Inaba (1953), of chromosomes; 4 species {ßulinus
i.e., 18 haploid, 36 diploid (Figs. 8, 20, coulboisi, B. truncatus, B. ugandae and
37). The chromosomes of this species Gyraulus circumstriatus) have 36 pairs;
are monocentric, as confirmed by Inaba and 1 species {ßulinus sericinus) has 72
(personal communication), and as found in pairs of chromosomes. Inaba (1953) and
other basommatophoran snails by Burch Inaba and Tanaka (1953) reported GyrawZtts
(1960a). hiemantium {=spirillus fide T. Habe) to
Burch (1960a, b) and Burch and Bush have 16 pairs of chromosomes, but
(1960) doubted Inaba's (1953) identification recently Dr. Inaba (personal communi-
of this species and thought instead that cation) has corrected his earlier report
perhaps he might have had P. integra, to 18 pairs of chromosomes. The report
since he stated that his species was of Azevedo and Gonçalves (1956) of 17
imported from North America. The pairs of chromosomes in Planorbarius
European P. acuta does not occur in comeus for the Portuguese population
North America, as has been pointed out he examined is not considered reliable
by Burch (1960b). P. acuta, however, until it has been confirmed (see Burch,
is found in Japan according to our own 1961). Chromosome numbers currently
identifications, which have been confirmed known for species of Gyraulus are shown
by Dr. William J. Clench of Harvard in Table III.
TABLE . Chromosome numbers of Gyraulus from different countries.
Species
FIGS. 22-27. Camera lucida drawings of meiotic chromosomes of Japanese Basommatophora.
FIGS. 22-24. Chromosomes of Gyraulus tokyoensis. Figs. 22, 23 are from the same individual, Figs. 24a, b
from another individual. FIG. 22. Metaphase I cell with 18 bivalents. FIG. 23. Metaphase I
cell with 19 bivalents. Fig. 41 is a photograph of these chromosomes. FIG. 24. Anaphase-
Telophase I chromosomes of a dividing cell. At one pole (Fig. 24a) t.here are 19+1 chromo-
somes (the arrow indicates the unmatched univalent); at the other pole (Fig. 24b) there are
19 chromosomes. Figs. 45a, b are photographs of these chromosomes. Figs. 22-24, X2490
FIGS. 25-27. Chromosomes of Segmentina hemispliaenila. FIG. 25. Pachytene chromosomes. FIG. 26.
Diakinesis chromosomes. FIG. 27. Metaphase I chromosomes. Figs. 25-27, X3060.
Gyraulus spirillus (Gould), This is a (ie in addition to the n=18, 2n=36 numbers)
.
.
,
rather common small planorbid in Japan, are supernumeraries and are not dupli-
and has been referred toas G hiemantium
. cations of other chromosomes of the
(Westerlund) by many authors. Dr. T. normal set. However, the supernumerary
Habe (personal communication) considers chromosomes of this species do not exhibit
G. hiemantium to be a synonym of G. differential staining and thereby differ
spirillus. Six specimens were studied from the heterochromatic elements found
from 2 localities; 4 specimens gave by Burch (1960b) in Gyraulus deflectus..
satisfactory results. Eighteen bivalents Segmentina (polypylis) hemisphaerula
were found in cells of the first meiotic (Benson). This species is common in
division (Figs. 21, 33, 34). many areas of the Orient, and has often
Gyraulus perstriatulus (Preston). This been refered to as "Polypylis nitidella
species has often been called G.biwaensis (Martens)" which is a synonym. Five
(Preston), a name considered a synonym specimens from one locality were studied,
of G. perstriatulus by Dr. Kuroda (1963). all with active gametogenesis. Eighteen
Twenty specimens were studied from 1 bivalents were found in cells of the first
locality;7 specimens gave satisfactory meiotic division of spermatogenesis (Figs.
results. Eighteen bivalents were found 25-27) and 36 chromosomes in mitotic
in cells of the first meiotic division of divisions (Fig. 11).
spermatogenesis and 36 chromosomes in Camptoceras hirasei Walker. This isa
mitotic divisions (Figs. 9, 35, 36). very rare species, now almost extinct in
Gyraulus tokyoensis Mori. Five speci- Japan. Nine specimens from 1 locality
mens of this species from the same popu- were examined; only 5 gave satisfactory
lation were studied, all with some cells results. No cells in meiotic divisions
undergoing gametogenesis. In 3 of the were observed, but prometaphase
specimens the meiotic and mitotic spermatogonial cells had 36 chromosomes
cells studied had 18 pairs of chromo- (Fig. 12).
somes (Figs. 10, 38, 42). However, the
other 2 specimens showed some variation LITERATURE CITED
inchromosome numbers. In 1 specimen,
some meiotic cells had 18 bivalents (Fig. AZEVEDO, J. F. and M. M. GONÇALVES,
22) and the only mitotic cell on which an 1956, Ensaios sobre estudo da
accurate count could be made had 36 numeraçao cromosómica de algumas
chromosomes, while other meiotic cells especies de m.oluscos de aqua doce.
clearly had 19 bivalents (Figs. 23, 41). Anais Inst. Med. trop., 13(4): 569-577.
In the other specimen showing variation BONHAM, ., 1955, Sensitivity to X-rays
in chromosome number, no meiotic cells of the early cleavage stages of the snail
were found with 18 bivalents, but they all Helisoma subcrenatum. Growth, 19(1):
had 19 bivalents or 19 bivalents plus a 9-18.

univalent. In the latter case, the uni- BURCH, J. ., 1959, Chromosomes of
valent was often displaced to one side aquatic pulmonate snails (Basommato-
of the figure (Figs. 43, 44), which might phora). Amer, malacol. Union ann.
be expected, due to pressure of the Reps., 1958, 25: 9-10. Dissert. Abstr.,
squash on its unequal attachment to the 20(4): 1487-1488.
spindle or due to the absence of a centro- 1960a,
, Chromosome mor-
mere. At Anaphase-Telophase I, such phology of aquatic pulmonate snails
cells show 19 chromosomes at one pole (Mollusca: Gastropoda). Trans. Amer,
and 20 at the other (Figs. 24, 45). Since micros. Soc, 79(4): 451-461.
we have not seen any meiotic cells in 1960b,
,
Chromosome studies
any of the 5 individuals studied with of aquatic pulmonate snails. Nucleus,
chromosomes that appear tobetrivalents, 3(2): 177-208.
we assume that the extra chromosomes 1960c,

,
Chromosomes of
412 BURCH, WILLIAMS, HISHINUMA, NATARA J AN
A /'J I
I
0m f mi
|39>
^
•âii»Vî
44
/
i?
FIGS. 39-47. Photographs of chromosomes of Japanese Basommatophora.
FIG.
FIG.
39.
40.
Spermatogonial late prophase cnromosomes of
Meiotic Metaphase I chromosomes of
; ja/)on¿ca.
Fossaria truncatula. This

X1900.
is a photograph of the chromo-

somes in Fig. 32. X2255.
FIGS. 41-45. Meiotic chromosomes of Gyraulus tokyoensis. Figs. 41-44 are of Metaphase I chromosomes;
Figs. 45a, b are of Anaphase-Telophase I chromosomes of one dividing cell. FIG. 41. Chromo-
somes of a cell with 19 bivalents. This is a photograph of Fig. 23. FIG. 42. Chromosomes of
a cell with 18 bivalents (side view). This is a photograph of Fig. 38. FIGS. 43, 44. Chromo-
somes of cells with 19 bivalents and a univalent (side view). Arrows indicate the univalents.
FIG. 45a. Pole with 20 chromosomes (n=19 + 1). The arrow indicates the unmatched uni-
valent. FIG. 45b. Pole with 19 chromosomes. Figs. 45a, b are photographs of Figs. 24a, b.
Fig. 41, X2280; Fig. 42, X2105; Figs. 43, 44, X1950; Fig. 45a, X2075; Fig. 45b, X2475.
FIGS. 46, 47. Chromosomes of Radix ollula. FIG. 46. Diakinesis chromosomes. This is a photograph of
Fig. 17. FIG. 47. Spermatogonial late prophase chromosomes. This is a photograph of Fig.
47. Fig. 46, X2000; Fig. 47, X2485.
Gyraulus circumstriatus, a freshwater Japan with the comparison with the

snail. Nature, 186(4723): 497-498. European material. Venus, Jap. J.
,
1960d, Chromosome numbers Malacol., 19(1): 25-42.
of schistosome vector snails. Z. , 1959, Anatomy of two forms
Trop. -Med. Parasit., 11(4): 449-452. of Lymnaea onychia Westerlund and
, 1961, The chromosomes of their ecological notes. Ibid., 20(3): 274-
Planorbarius comeus (Linnaeus), with 288.
a discussion on the value of chromosome LA COUR, L., 1941, Acetic-orcein: A
numbers in snail systematics. Basteria, new stain fixative for chromosomes.
25(4/5): 45-52. Stain Techn., 16: 169-174.
1962,
, Cytotaxonomic studies LE CALVEZ, J. and CERTAIN, P., 1950,
of freshwater limpets (Gastropoda: Données caryologiques sur quelques
Basommatophora). I. The European pulmones basommatophores. R. .
Lake Limpet, Acroloxus lacustris. Acad. Sei., Paris, 231: 794-795.
Malacologia, 55-72. 1(1): KURODA, T., 1949, In: Illustrated en-
, Cytological studies of
1964, cyclopedia of the fauna of Japan. Ed.:
Planorbidae (Gastropoda: Basommato-
phora). I. The African subgenus
s.S.. Ibid., 1(3): 387-400.
/5 Seionsuke Uchida. TheHokuryukanCo.,
Ltd., Tokyo, p 1-1898.
1963, Mollusks in Ogura-ike,
,
BURCH, J. B. and BUSH, L. L., 1960, (Survey on Ogura-ike; Ogura-ike Land

Chromosomes of Physa gyrina Say Improvement Area). Venus, Jap. J.
(Mollusca: Pulmonata). J. Conchyl., malacol., 22(3): 295-297.
100: 49-54. MAHONEY, F. J., 1940, Spermatogenesis
HUBENDICK, ., 1951, Recent Lymnae- with special reference to certain extra-
idae; their variation, morphology, tax- nuclear structures in the pulmonate
onomy, nomenclature, and distribution. Physa gyrina Say. Univ. Colorado
Kungl. svenska Vet.-akad. Handl., ser. Studies, 26(3): 81-83.
4, 3(1): 1-223. MARTENS, E. v., 1867, Ueber die
GOULD, A. A., 1859, Lymnaea ollula. ostasiatischen Limnaeaceen. Mala-
Proc. Best. Soc. Nat. Hist., 7: 40-45. kozool. Bl., 1867: 211-227.
INABA, A., 1953, Cytological studies in NATARAJAN, R., 1960, Further cyto-
moUusks. I. Chromosomes in logical studies in Pulmonata (Mollusca:
basommatophoric Pulmonata. J. Sei. Gastropoda). J. zool. Soc. India, 12(1):
Hirosh. Univ., Ser. ., Div. 1, 14: 69-79.
221-228. NEWCOMER, E.H., 1953, A new cyto-
1961,
,
Cytotaxonomy of the logical and histological fixing fluid.
Euthyneuran Gastropods. Venus, 21(4): Science, 118(3058): 161.
402-413. PERROT, J.-L., 1930, Chromosomes
INABA, A. andTANAKA, H., 1953, Studies et hétérochromosomes chez lesgastér-
on the chromosome numbers of some podes pulmones. Rev. suisse Zool.,
freshwater gastropods. J. Sei. Hirosh. 37: 397-434.
Univ., Ser. ., Div. 1, 14: 213-220. , 1934, A propos du nombre des
ITAGAKI, H., 1956, Anatomy of Lymnaea chromosomes dans les deux lignées
iFossaria) truncatula (Müller) from germinales du gastéropode herma-
nos. 35, 36. Chromosomes oi Gyraulus perstriatulus . FIG. 35. Metaphase , polar view. FIG.
36. Metaphase I, side (equatorial) view. Figs. 35, 36, X3060.
FIG. 37. Metaphase I chromosomes oí Physa acuta, polar view. X3060.
FIG. 38. Metaphase I chromosomes of G jrau/us iofeyoensis, n=18, side (equatorial) view.
Fig. 42 is a photograph of these chromosomes. X2490.
r^
32
35 36
37
%'^
38
FIGS. 28-38. Camera lucida drawings of meiotic metaphase chromosomes of Japanese Basommatophora.
FIGS. 28-31. Chromosomes of Radix ollula. are in polar view; Figs. 30, 31 are in side
Figs. 28, 29
(equatorial) view. FIG. 28. Metaphase I chromosomes. FIG. 29. Metaphase II chromosomes.
FIG. 30. Metaphase I chromosomes. FIG. 31. Anaphase I chromosomes. Figs. 28-31, X3060.
FIG. 32. Metaphase I chromosomes of Fossaria truncatula. Fig. 40 is a photograph of these chromo-
somes. X3060.
FIGS. , 34. Metaphase I chromosomes of Gyraulus spirilliis. FIG. 33. Polar view. FIG. 34. Side
(equatorial) view. Figs. 33, 34, X3060.
phrodite Limnea stagnalis (variété Zoologie. Tomes 2-3. Atlas.

rhodani). Ibid., 41: 693-697. SMITH, E. A., 1876, Descriptions of
PERROT, J.-L. and PERROT, M., 1938, some new species of land and fresh-
Note sur les chromosomes de cinq water shells, and remarks on other
espèces de limnées. C. R. Soc. Phys. species found in Japan. Quart. J.
Hist, nat, Genève (Suppl. Arch. Sei. Conch., 1: 118-127.

phys. nat.), 53: 92-93. TAYLOR, D. W., WALTER, H. J. and
QUOY, M. and GAIMARD, P., 1832-35, BURCH, J. ., 1963, The subgenus
Voyage de découvertes de l'Astrolabe Hinkleyia Lymnaeidae) in
{Stagnicola:
execute par ordre du Roi pendant les the western United States. Malacologia,
années 1826-1829 sous le commande- 1(2): 237-281.
ment de M. J. Dumont d'U^ville.
ZUSAMMENFASSUNG
CHROMOSOMENZAHLEN EINIGER JAPANISCHER SÜSSWASSERSCHNECKEN
(BASOMMATOPHORA: BRANCHIOPULMONATA)
Die Chromosomenzahlen für 10 japanische SOsswasserbasommatophoren sind: 16

Paare für Radix ollula, jedoch 17 Paare R. japónica und R. onychia; und 18 Paare
für Fossaria truncatula, Physa acuta,Gyraulus spirillus,G. perstriatulus, G. tokyoensis,
Segmentina hemisphaerula und Camptoceras hirasei. Zwei Exemplare von G. tokyoensis
hatten 2-3 überzählige Chromosomen, die entweder ein zusatzliches zweiwertiges
Element darstellten (n=19) oder ein zwei- und ein einwertiges (n=19+l). Da jedoch
trivalente Chromosomen nicht vorkamen, wird angenommen, dass die Oberzähligen
Chromosomen nicht Duplikationen von normalerweise vorhandenen sind. Dass die
Chromosomenzahl von "Lymnaea' ollula derjenigen der Gattung Radix näher liegt als
der vonFossana.zuwelcher letzteren sie gewöhnlich gerechnet wird, scheint Hubendicks
(1951) anatomische Befunde zu bestätigen.
Die Chromosomen dieser japanischen Arten sind monozentrisch und gleichen in
dieser Hinsicht denen euthyneurer Schnecken aus anderen Ländern.
RESUMEN
CROMOSOMAS DE ALGUNOS CARACOLES DULCEACUICOLAS JAPONESES

Se informa el número de cromosomas para 10 especies de basomatóforos japoneses.

Se observaron 16 pares de cromosomas en Radix ollula y 17 pares en .
ja/xmica y
R. onychia; l8 pares en Fossaria truncatula, Physa acuta, Gyraulvs spirillus, G.
perstriatulus, G. tokyoensis, Segmentina hemisphaerula y Camptoceras hirasei. Dos
ejemplares deG. tokyoensis tenían cromosomas supernumerarios, formando un bivalente
extra (n=19), o un bivalente más un univalente (n=19+l). Desde que no se encontraron
trivalentes en ningún espécimen de G. tokyoensis, se que los cromosomas
adicionales no son duplicaciones de otros cromosomas del juego normal. El número
de cromosomas de 'Lymnaea' ollula es más cercano al del género Radix que al de
Fossaria, al cual es corrientemente asignada; esto parece corroborar las observaciones
anatómicas de Hubendick (1951).
Los cromosomas de estas especies japonesas son monocéntricos y por lo tanto
semejantes a los que se encuentran en los eutineuros de otros países.
ç
THE SPECIES OF VENTRIDENS

(STYLOMMATOPHORA:
Leslie Hubricht^
ABSTRACT
)
The bidentate species (section Ventridens s. s. ) of the genus Ventridens are
characterized by the presence of 2 apertural lamellae in the neanic stage. In
this paper 12 species are recognized and divided into 2 groups, the V. gularis,
group (4 species), in which the animal is dark, and the V. pilsbryi group (8
species), in which it is pale. Two species are described as new: V. pilsbryi
which is separated from V. gularis, but more closely related to V. collisella,
and V. monodon, which is most closely related to V. theloides.
The species of each group do not normally live with each other, but will live
with species of the other group. Their distribution in the U. S. A. has been
mapped. The disjunct distribution shown is, in some cases, real and not due to
lack of data and cannot be related to ecological factors.
The species treated in this paper belong V. lasmodon which has been found with
to the Section Ventridens s.s. of Pilsbry other species at 3 localities. Not only
(1946). They are characterized by the do the different species occur in
presence of 2 lamellae within the aperture separate colonies, but they are often
in the neanic stage, one on the columella geographically exclusive. One area will
and one on the base. One or both of these have only one species, and an adjoining
lamellae may be reduced or wanting in area another; so that the species occur
mature shells. In addition there may be a in a crazy-quilt pattern that seems to
small tubercule or lamella above the have no relation to the underlying geo-
columellar lamella, and in very young logical formations, exposure, or other
shells one or more fine lamellae within readily observable ecological factors.
the outer wall. The disjunct distributions of some of the
The bidentate species of Ventridens can species as shown on the accompanying
be divided readily into 2 groups by the maps is real in some cases and not due
color of the animal. In the V gularis . to the absence of collecting in the
group the animal is dark, olive or bluish intervening areas.
gray. In the V. pilsbryi group the animal
is pale, yellow with perhaps some grayish Family ZONITIDAE
flecking along the back. In the V. gularis
group the shell rarely has more than 6.5 Subfamily GASTRODONTINAE
whorls, and the aperture is proportionally Genus Ventridens W. G. Binney
larger. In the V. pilsbryi group the
diameter of the whorls usually increase Ventridens pilsbryi Group
more slowly, the whorls are more
flattened above, and the shell may have Key to the species
as many as 9 whorls.
la. Striation of upper surface sharp and dis-
Species of the V. gularis group are
tinct 2
frequently found with species of the V.
lb. Striation weaker, the striae rounded,
pilsbryi group. Species of the V. gularis
glossy 3
group have not been found living together.
Species of the V. pilsbryi group have not 2a, Umbilicus a narrow perforation
been found together, with the exception of V. collisella
^3235-23rd Ave. , Meridian, Miss. , U. S. A.
(417)
.
418 L. HUBRICHT
2b. Umbilicus more open, 10-12 times in dia- and acute; within the aperture there are
meter of shell V. decussatus 2 lamellae;the columellar lamella in
young shells is long, high, and leans
3a. Umbilicus in adults a narrow perforation
4
strongly toward the basal lamella, in
or closed
adults this lamella is greatly reduced;
3b. Umbilicus at least 0. 4 mm in diameter in
adults 6
in young shells the basal lamella is long,
stout, usually erect, in adults it
and
4a. Adults with lamellae 5
becomes shorter, higher, and thinner, the
4b. Adults without lamellae V. theloides
apex often curving toward the columellar
5a. Adults with 2 lamellae V pilsbryi
.
lamella; in some immature shells there
5b. Adults with basal lamella only is a small nodule on the columellar callus
V. monodon above the columellar lamella. Animal
pale yellowish, with some gray along the
6a. Adults less than 7 mm in diameter
V. coelaxis
back.
6b. Adults larger 7
Diam. 9.6 mm, height 6.8 mm; 7.8
whorls. Holotype.
7a. Shell elevated, umbilicus less than 0.5 Diam. 8.7 mm, height 7.1 mm; 8.0
mm in diameter 8
whorls. Paratype.
7b. Shell more depressed, umbilicus larger.
Diam, 9.1 mm, height 5.1 mm; 7.4
9
whorls. Paratype.
8a. Aperture without lamella in the adult Typelocality: Tennessee; Monroe
V. theloides CoT^ near Citico Creek, at mouth of
8b. Aperture with a basal lamella in the adult. Hell Hole Branch, Cherokee National
V monodon
.
Forest. Holotype 127735, figured para-
9a. Umbilicus contained 5 to times in dia-
6 types 127736, other paratypes 127737,
meter V. lawae Chicago Natural History Museum, other
9b. Umbilicus 3 to 4 times in diameter paratypes 27051, collection of the author.
V lasmodon
.
Ventridens pilsbryi, although a
common species, has remained unde-
scribed because of confusion with V.
Ventridens pilsbryi, new species gularis. The latter has a smaller, more
globose shell with fewer whorls, and the
PL I A-F. Map I A.
animal is nearly black. It is most closely
Ventridens gida ris (Say). Pilsbry, related to V. coZ/¿se/ía, which has a larger,
1946, Acad. Nat. Sei. Philadelphia, more elevated shell, with very sharp
Monogr. 3, vol. ,
p 443 (in part). sculpture above.
Ventridens pilsbryi ranges from south-
Shell pale brownish-yellow, translucent ern Kentucky and southwestern Virginia
except near aperture where it is made south to southern Alabama and eastern
opaque by an internal callus; spire low Louisiana.
dome-shaped, composed of 7.5 to 8 narrow Named in honor of the late Dr. Henry
tightly coiled whorls; periphery somewhat A. Pilsbry.
angular in immature shells, but becoming
well-rounded in adults; umbilicus about
Ventridens collisella (Pilsbry)
0.4 mm
in diameter in shells with up to
4 whorls, becoming rapidly smaller, not Map I
over 0.2 mm inshellsof 6or more whorls,
sometimes completely closed; surface Gastrodonta collisella Pilsbry.
glossy, with numerous distinct growth- 1896. Nautilus, 9: 123.
wrinkles, which are somewnat Tveaker on Ventridens collisella (Pilsbry).

the base; base with indistinct spiral Pilsbry. 1946, Acad. Nat. Sei. Phila-
striae; aperture lunate, peristome thin delphia Monogr. 3, vol. II, p 450.
BIDENTATE SPECIES OF VENTRIDENS 419
Plate I
FIG. A-C. Ventridens pilsbry? Hubricht, FIG. G-I. Ventridens monodon Hubricht,
holotype. holotype.
FIG. D-F. Ventridens pilsbryi Hiibricht, FIG. J-L. Ventridens monodon Hubricht,
paratypes. paratypes.
420 L. HUBRICHT
Ventridens collisella is the largest of Ventridens theloides is distinguished

the bidentate s'geciesoi Ventridens. Large by moderately high, dome-shaped
its
specimens may exceed 11 mm in diameter spire, and by the absence of teeth in the
and 8 mm
in height. It can be distinguished mature shell. In young shells the
from all other species except V. umbilicus is 0.5 to 0.8 mm in diameter.
decussatus by its sharp sculpture of the In most mature shells it remains about
upper surface. V. decussatus has a the same, but in some lots it becomes
smaller shell with a more open umbilicus. much smaller, being nearly closed, as in
V collisella ranges from southwestern
. form nodus. It can be separated from
Virginia through eastern Tennessee to V. lawae by its higher spire and smaller
northeastern Alabama. umbilicus.
V. theloides ranges from southeastern
Ventridens decussatus (Walker Kentucky, southern West Virginia, and
and Pilsbry) southwestern Virginia, south to southern
Alabama.
Map I D.
Ventridens monodon, new species

Gastrodonta gularis decttssaia "Pils-
bry & Vanatta. " Walker & Pilsbry.
1902, Proc. Acad. Nat. Sei. Phila-
PL I G-L. Map I E.
delphia, p 436.
decussatus Shell pale brownish-yellow, sub-trans-
Ventridens gularis
(Walker & Pilsbry). 1946, Acad. Nat. lucent except near aperture where it is
Sei. Philadelphia, Monogr. 3, vol. II,
made opaque by its internal callus; spire
p 448. moderately raised, dome-shaped, com-
posed of 7.5 to 8 narrow, tightly coiled
Ventridens decussatus can be recog- whorls; periphery somewhat angular in
nized by the sharp sculpture of the upper immature shells, but becoming well-
surface, the angulate periphery, the rounded in umbilicus about 0.8
adults;
distinctly open umbilicus, the presence of mm in diameter in shells with up to 5
lamellae in the adult shell, and the pale whorls, becoming somewhat smaller, to
animal. nearly closed, in adults; surface glossy,
V. decussatus is usually found at higher with numerous distinct growth-wrinkles,
elevations in the southern Appalachians, which are much weaker on the base; base
ranging from southern West Virginia, with weak spiral striae; aperture lunate,
western North Carolina and eastern peristome thin and acute, the basal lip
Tennessee to northern Georgia. straight, the outer lip well rounded; with-
in the aperture there are 2 lamellae;
Ventridens theloides (Walker the columellar lamella in immature shells
and Pilsbry) is long, high, and leans strongly toward
the basal lamella, in adults this lamella
Map I B.
is absent; in immature shells the basal
lamella is long, thin, and leans toward
Gastrodonta gularis theloides A. D.
Brown. Walker & Pilsbry. 1902 Proc. the columellar lamella, in adults it
Acad, Nat. Sei. Philadelphia, p 434. becomes shorter, higher and the apex
Ventridens gularis theloides (\Ma\ker curves toward the columellar lamella.
& Pilsbry). Pilsbry, 1946, Acad. Nat. Animal pale yellowish, with some gray
Sei. Philadelphia, Monogr. 3, vol. II, along the back.
p 447. Diameter 8.3 mm. height 5.4 mm; 8
Ventridens gularis form nodus Pils- whorls. Holotype.
bry. 1946, Acad. Nat. Sei. Phila- Diameter 9.1 mm, height 6.2 mm; 7.8
delphia, Monogr. 3, vol. ,
p 447. whorls. Paratype.
BIDENTATE SPECIES OF VENTRIDENS 421
Map I
Distributions of species of the Ventridens pilsbryi Group.
A. Ventridens pilsbryi Hubricht. E. Ventridens monodon Hubricht.

B. Ventridens theloides (Walker & Pilsbry). F. Ventridens coelaxis (Pilsbry).
Ventridens collisella (Pilsbry). G. Ventridens lawae (SN. G. Binney).
D. Ventridens decussatus (Walker & Pilsbry). H. Ventridens lasmodon (Phillips).
422 L. HUBPICHT
Diameter 8.1 mm, height 5,7 mm*, 8.2 Nautilus, 12: 140.
whorls. Paratype. Ventridens coelaxis (Pilsbry) Pils- .
Type locality; Alabama; Butler Co.; bry, 1946, Acad. Nat. Sei. Phila-
ravine, 3 miles northwest of McKenzie. delphia, Monogr. 3, vol. p 456.,
Holotype 127738, figured paratypes
127739, other paratypes 127740, N. H. Ventridens coelaxis is the smallest
M., other paratypes 23293, collection of species of the V. pilsbryi group, rarely
the author. exceeding 6.5 mm
in diameter and with
Ventridens monodon is most closely never more than 7 whorls. Immature
related to V. theloides, differing in the shells have an angular periphery and a
presence of the basal lamella in adult third, smaller, lamella above the
shells. columellar lamella, by which it can be
V. monodon is known only from a distinguished readily from immature V.
small area in southern Alabama. It has lawae.
been collected in Crenshaw, Lowndes, V. coelaxis is usually found in the
Butler, Conecuh, and Clarke Counties. mountains at high elevations. It ranges
from southwestern Virginia, to north-
Ventridens lawae (W. G. Binney) eastern Tennessee, and northwestern
North Carolina.
Map I G.
Zonites lawae Binney. 1892, Bull.
Ventridens lasmodon (Phillips)
Mus. Comp. Zool. , 4th Suppl. 22: 167.
Ventridens lawae (W, G. Binney). Map I H.

Pilsbry. 1946, Acad. Nat. Sei. Phila- Heitx lasmodon Phillips. 1841,
delphia Monogr. 3, vol. p 453 (in , Proc. Acad. Nat. Sei. Philadelphia, 1:
part). 28.
Ventridens lawae cumberlandicus Zonites macilenta Shuttleworth.
Pilsbry. 1946, Acad. Nat. Sei. Phila- 1852, Mittheil. naturforsch. Ges. Bern.
delphia Monogr. 3, vol. p 455. , Nr. 248, p 195 (nomen oblitum).
Ventndens lasmodon (Phillips).
Ventridens lawae is characterized by Pilsbry, 1946, Acad. Nat. Sei. Phila-
its low spire, and large umbilicus. The delphia, Monogr. 3, vol. p 457. ,
umbilicus in very immature shells is about
1 mm in diameter, in mature shells it Ventridens lasmodon can be confused
may vary from 0.4 to 2 mm, mm only with V. lawae, from which it differs
approaching V. theloides on one hand and by its larger umbilicus In V. lawae the
.
V. lasmodon on the other. In some lots umbilicus is usually well-like, with the
the adults are all edentate and in others later whorls of about the same diameter.
they are usually laminate, but in large In V. lasm,odon the umbilicus is usually
lots there are at least a few adult shells funnel-shaped, with the later whorls
which are edentate. V. lawae is an increasing in diameter.
extremely variable species, but there is
Ventridens lasmodon ranges from
not sufficient pattern to the variation for
southwestern Virginia through eastern
the erection of satisfactory subspecies.
Tennessee to St. Clair Co., Alabama.
V. lawae ranges from southern Kentucky
The type locality for this species is
and southwestern Virginia south to north-
"Alabama", and is known only from St.
western Georgia and northeastern
Clair County. I found it abundant on a
Alabama.
hillside, 3 miles west of Pell City. Since
Ventridens coelaxis (Pilsbry) this is on the main Birmingham -Atlanta
highway and was readily accessible, it
Map I F.
could very well be the original type
Gastrodonta toxis Pilsbry. 1899, locality.
BroENTATE SPECIES OF VENTRIDENS 423
Map
Distributions of species of the Ventridens gularis Group.
A. Ventridens gularis (Say). C. Ventridens suppressus (Say).

B. Ventridens cerinoideus (Anthony). D. Ventridens virginicus (Vanatta).
424 L. HUBRICHT
Ventridens gularis Group Amer. J. Conch. 1- 351.

Ventridens cerinoideus (Anthony).
Key to the species
Pilsbry, 1946, Acad. Nat. Sei. Phila-
Umbilicus a narrow perforation or closed
la.
2
delphia, Monogr. 3, vol. , p 451.
lb. Umbilicus wider, more than l/lO of the

diameter 3
Ventridens cerinoideus differs from V.
gularis by the absence of lamellae in the
2a. Adults with 2 lamellae, west of the Coastal adult shell. Rarely they are present
Plain V. gularis There are 2
although reduced in size.
2b. Adults with lamellae greatly reduced or
forms of this species: one with a distinct
absent Coastal Plain
,
basal callus is found in upland habitats,
V. cerinoideus
and the other with the basal callus greatly
3a. Columellar lamella simple in immature reduced or wanting is found in swamps.
shells V. suppressus For the most part these 2 forms are
3b. Columellar lamella double in immature distinct, but there is some intergradation.
shells V. virginicus This species could, with some propriety,
Ventridens gularis (Say) be considered to be a subspecies of V.
gularis. Breeding experiments to de-
Map II A. termine whether the differences are
Helix gularis Say. 1822, J. Acad. genetic or due to ecological factors are
Nat. Sei. Philadelphia, 2: 156. desirable. V. gularis ranges from the
Helix bicostata Pfeilier. 1840, Piedmont physiographic province to the
Symbolae ad Hist. Hel. 3: 69. (nomen edge of the fall line, and V. cerinoideus
oblitum). occurs on the Coastal Plain below the
{Zonites cerinoidea) var. cuspidata fall line.
Lewis. 1875, Proc. Acad. Nat. Sei. Ventridens cerinoideus is a species of
Philadelphia, p 335. the Coastal Plain. It ranges from eastern
Ventridens gularis {Say). Pilsbry, Maryland south to northern Florida and
1946. Acad. Nat. Sei. Philadelphia, west to southern Alabama.
Monogr. 3, vol. ,
p 443 (in part).
Ventridens suppressus (Say)
Ventridens gularis form cuspidatus
(Lewis). Pilsbry, 1946, Acad. Nat. Map II C.
Sei. Philadelphia, Monogr. 3, vol. , Henx suppressa Say. 1829, The
p 446.
Ventridens suppressus magnidens Disseminator of Useful Knowledge. 2:
Pilsbry. 1946, Acad. Nat. Sei. Phila- 229.
Ventridens suppressus (Say). Pils-
delphia, Monogr. 3, vol. p 442., bry, 1946, Acad. Nat. Sei. Phila-
Ventridens gularis can be distinguished delphia, Monogr. 3, Vol. ,
p 438.
from all other species by the nearly Ventridens suppressus divisidens
black animal, the very small umbilicus, Pilsbry, 1946, Acad. Nat. Sei. Phila-
and the presence of lamellae in adult delphia, Monogr. 3, vol. ,
p 443.
shells.
Ventridens gularis ranges from Ventridens suppressus may be recog-
southern Indiana to southwestern Pennsyl- nized by its small size, its distinctly open
vania and south to southern Alabama, umbilicus, and simple columellar lamella
It is the most widely distributed species
in the immature shell. The amount of
in the typical section of the genus,

reduction in the lamellae at maturity
varies greatly. They may be fully de-
Ventridens cerinoideus (Anthony) veloped to completely absent. In some
lots from the southwestern part of the
Map II B.
range the basal lamella moves up onto
Helix cerinoidea Anthony. 1865, the outer wall at maturity. Although there
SPECIES OF VENTRIDENS 425
seems to be a trend towards larger Zool., Harvard, 22(4): 163-204, pis.

lamellae in adults in the southwestern part 1-4.
of the range of the species, the variation LEWIS, J., 1875, Descriptions of new
is too great in any area for the erection species of American land and fresh-
of satisfactory subspecies. water shells. Proc. Acad. Nat. Sei.
V. suppressus ranges from southeastern Philadelphia, 1875, 334-337.
Michigan to southern New York, south to PFEIFFER, L., 1846, Symbolae ad
north-central North Carolina. Historiam Heliceorum. 3: 69-91.
PHILLIPS, J. S., 1841, Descriptions of
Ventridens virginicus (Vannatta) two new American species of the genus
Helix. Proc. Acad. Nat. Sei. Phila-
Map D.
delphia, 1: 27-28.
Zonitoides suppressus virginicus PILSBRY, H. A., 1896, Description of a
Vanatta. 1936, Nautilus, 49: 99. new Gastrodonta. Nautilus, 9(11): 123-
Ventridens suppressus virginicus 124.
(Vanatta). Pilsbry, 1946, Acad. Nat. ,
1899, Descriptions of new
Sei. Philadelphia, Monogr. 3, vol. II, American land shells. Ibid., 12(12):
p440. 140-142.
, 1946, Land Mollusca of North
Ventridens virginicus can be readily America (North Mexico). Acad., Nat.
of
distinguished from all other species by the Sei. Philadelphia, Monogr. 3, 2(1): 438-
double columellar lamella. Fully adult 458.
shells usually show enough of the double SAY, T., Descriptions of univalve
1822,
lamella to be distinguishable from those of shells J. Acad.
of the United States.
V. suppressus. Nat. Sei. Philadelphia, 2: 149-179.
V. virginicus ranges from southeastern 1829,
,
Descriptions of some
Michigan to northwestern New York south new terrestrial and fluviatile shells of
to eastern West Virginia and western North America. New Harmony Dissem.
Virginia. Useful Knowledge. 2(15): 229-230.
SHUTTLEWORTH, R. J., 1852, Diagnosen
LITERATURE CITED neuer Mollusken. Mittheil, naturforsch.
Ges. Bern., Nr. 248: 194-207.
ANTHONY, J. 1865, Description of
G., VANATTA, E. G., 1936, A new subspecies
new species of shells. Amer, J. Conch., of Zonitoides (Ventridens) suppressa
1(4): 351. (Say). Nautilus, 49(3): 99-100.
BINNEY, W. G., 1892, A fourth supple- WALKER, ., and PILSBRY, H. A., 1902,
ment to the fifth volume of the The Mollusca of the Mt. Mitchell region,
Terrestrial Air-Breathing Mollusks of North Carolina. Proc. Acad. Nat. Sei.
the United States. Bull. Mus. Comp. Philadelphia, 54: 413-442.
ZUSAMMENFASSUNG
DIE ZWEIGEZÄHNTEN ARTEN VON VENTRIDENS
(STYLOMMATOPHORA: ZONITmAE)
Die zweigezähnten Arten der Gattung Ventridens (Sectio Ventridens s.S.) sind im
jugendlichen Stadium durch die Anwesenheit von 2 aperturalen Lamellen gekennzeichnet.
Hier werden nun 12 Arten anerkannt und in 2 Gruppen eingeteilt: die V. gîlaris Gruppe
(mit 4 Arten) in der das Tier dunkel ist, und die V. pilsbryi Gruppe (mit 8 Arten) in
der das Tier blass ist. Zwei Arten sind neu beschrieben: V. pilsbryi sp. ., die bis
jetzt mit V.gularis verwechselt wurde, und V. monodon, sp. ., die von V. theloides
unterschieden wird.
Die Arten jeder dieser Gruppen kommen normalerweise nicht gemeinsam vor, leben
jedoch mir Arten der anderen Gruppe zusammen. Ihre Verbreitung in den V.S.A. (im
426 L. HUBRICHT
Osten und Südosten) Karten gezeigt. Das zersplitterte Vorkommen einiger Arten
is auf
ist in manchen Faellen echt undlässt sich weder durch mangelnde Aufzeichnungen
noch durch Ökologische Bedingungen erklären.
RESUME
LES ESPÈCES BroENTÉES DE VENTRIDENS
(STYLOMMATOPHORA: ZONITIDAE)
Les espèces bidentées du genre Kenindens (Section Kenindens s. s.) sont caractérisées
par la présence de 2 lamelles aperturales dans le stade juvenile. Ici, 12 espèces sont
reconnues et divisées en 2 groupes, le groupe V. gularis (4 espèces) où l'animal est
foncé, et le groupe V. pilsbryi (8 espèces) où il est pâle. Deux espèces nouvelles sont
décrites: V. pilsbryi sp. ., qui avait été confondu avec V. gularis; et V. monodon,
sp. ., qui est distingué de V. theloides.
Les espèces de chaque groupe ne vivent pas normalement l'ime avec l'autre, mais
vivent ensemble avec des espèces de l'autre groupe. Leur distribution dans les É.U.A.
(l'est et le sud-est) a été marquée sur des cartes géographiques. La distribution
discontinue de certaines espèces est, en certains cas, réelle est n'est pas due à l'absence
de données; aussi, elle ne s'explique pas par les données écologiques.
RESUMEN
LAS ESPECIES BIDENTADAS DE VENTRIDENS
(STYLOMMATOPHORA: ZONITIDAE)
Las especies bidentadas (sección Ventridens s. s.) del género Ventridens están
caracterizadas por la presencia de dos lámelas aperturales en el estado neánico. En
este trabajo se reconocen 12 especies, divididas en dos grupos, el de V. gularis (con
4 especies), en el cual el animal es obscuro, y el de V. pilsbryi (con 8 especies),
con animal pálido. Dos especies se describen como nuevas: V. pilsbryt, segregada
de V. gularis pero más cercana a V. collisella, y V. monodon, más estrechamente
relacionada con V. theloides.
Las especies normalmente no viven juntas con las del mismo grupo, pero pueden
vivir con especies del otro grupo. Se da un mapa de distribución en U. S. A. En
algunos casos la distribución discontinua es real y no debida a falta de datos, y no se
puede relacionar con factores ecológicos.
THE MUSSEL FAUNA OF THE UPPER CUMBERLAND BASIN

BEFORE ITS IMPOUNDMENT 1
Joe Kendall Neel^ and William Ray Allen^
ABSTRACT
During 1947- 1949, shortly before impoundment of Lake Cumberland by closure
of Wolf Creek Dam, a study was made of the Naiad fauna of the upper Cumber-
land River basin (Kentucky, U. S. A.). Results are compared with an earlier
survey of the same area in 1911. In the survey here reported, 59 distinct fresh-
water mussels or clams were found of which 16 belong to the "Cumberlandian"
group unique to the Cumberland and Tennessee Rivers. Only 4 species (including
a single Cumberlandian form) had surmounted Cumberland falls and none of
these were restricted to that area. The earlier survey had disclosed 60 species,
of which only 56 coincided: 3 species recovered in the later survey had not been
found previously and 4 species then collected were not found later. In the 36
years since the earlier survey various mussels have suffered decimation by coal
mine acids in the uppermost main stem and in the major tributaries to this area
of the river. The genus Dysnomia has been practically eliminated from its
former region of greatest variety. A number of other mussels have gained in
prevalence.
Only some of the marked changes observed in prevalence, relative abundance
and distribution may be explained by the decline in pearling and the presence of
mine acids. The upper Cumberland was not significantly exploited by the pearl
button industry. Incertain instances the causes of the variation are still obscure.
The Cumberland and Tennessee Rivers the lower 50 miles of their channels (in
are inhabited by many of the common Kentucky and Tennessee) are generally
mussels of the Mississippi Basin; living within 10 miles of each other. At one
there also are a number of species that locality (about 20 river miles up the
have been reported from no other drainage Tennessee from its mouth) the Cumberland
areas. Students of mussels, particularly loops to within a mile of its sister river.
A. E. Ortmann, have referred to these The Cumberland originates in south-
endemic forms as the Cumberlandian eastern Kentucky and northeastern
Group (see Table 4), although more of Tennessee. Its main stem flows west and
them occur in the Tennessee River and south from Kentucky into Tennessee,
none are unique to the Cumberland. Studies assumes a westerly course in Tennessee,
of the mussel fauna of the Tennessee and then bends northwest to reenter
River have been reported in a number of Kentucky and join the Ohio. The main
articles, but the mussels of the Cumber- stem of the Tennessee is largely formed
land River are known mainly from a single by rivers that originate in western Virginia
publication of Wilson and Clark (1914) and North Carolina and proceed toward a
based upon a 1911 survey. Ortmann col- rendezvous in the general region of Knox-
lected there but used his data to supple- ville, Tennessee. They are later joined
ment observations on the Tennessee. by a tributary that drains a small area
The Cumberland and Tennessee both in northern Georgia. The main stem
enter the Ohio River from western moves southwest across Tennessee and
Kentucky. Their mouths are separated by enters Alabama just west of the Alabama-
only a 12 1/2 mile reach of the Ohio, and Georgia line. Its course in Alabama
^Contribution from the Department of Zoology, University of Kentucky and the Potamological
Institute University of Louisville.
,
^Director, The Potamological Institute, University of Louisville, Louisville, Kentucky, U.S.A.
'^Late Professor of Zoology, University of Kentucky.
(427)
—
428 NEEL AND ALLEN
(southwest, then northwest) requires about STREAM CHARACTERISTICS

200 channel miles. It then bends almost AND HABITATS
due north through Tennessee and finally
takes a nornorthwest to northwest path in The main stem of the Cumberland River
Kentucky. begins at the union of Poor, Clover, and
It is assumed that the close proximity Martin's Forks near Harlan, Kentucky
(or possible earlier confluence) of the (Fig. 1). Its course is generally westward
mouths and lower reaches of these two until it leaves the State from Monroe
rivers has facilitated dispersion of County.
Cumberlandian naiads throughout, each This report concerns the eastern reach
from individual focal points in either river. of the river from Harlan County, Kentucky,
Wilson and Clark (1914) and Ortmann to the Tennessee line. This stretch is
(1918, 1924, 1925) believed they located separated into two distinct mussel realms
focal areas for a number of forms. by Cumberland Falls (between Stations
Information for this article was obtained 9 and 10, Fig, 1). The Laurel and
shortly before closure of Wolf Creek Dam Rockcastle Rivers enter shortly below the
across the Cumberland River just below falls and Big South Fork flows in at Burn-
Rowena Ferry in Russell County, Kentucky side. About 50 years ago the U. S. Army
(between Stations 21 and 22, Fig. 1). Field Corps of Engineers constructed a system
studies covered the period October 1947 of locks and dams that was intended to
to December 1949. Impoundment has maintain year round towboat navigation up
since formed "Lake Cumberland" that to the region of the falls. At the time
extends upstream to Cumberland Falls and of our study (1947-49) the system had
into the lower reaches of most tributaries largely fallen into disuse, although the
that are represented in our collections. dam at Lock 21 (Station 18) formed a pool
Additional distributional records — that extended above Burnside.
lacking since Ortmann'sworkofthe 1920's The Upper Cumberland drainage basin
led to revision of interspecific relations contains many coal deposits that are rather
of the Cumberlandian and other groups extensively mined. Iron sulfide that occurs
had long been sought, and time to secure with the coal is usually exposed to water,
them in the upper Cumberland under oxygen, and bacterial action in these mines
relatively natural conditions was growing and the following reactions take place:
short in 1947. This situation also offered
FeS2 + 202 FeS04 + S
one of the few remaining opportunities to FeS2 + 702 + 2H2O 2FeS04 + 2H2SO4
study big river mussels before and after
impoundment of their normal habitat. Sulfuric acid (H2SO4) created by the
The authors were aided financially by second reaction is responsible for the
the Department of Zoology, University of acidity of drainage from the mines. The
Kentucky, and by a grant from the Uni- presence of this mine water has un-
versity Research Fund. Dr. Henry van doubtedly restricted occurrence of naiads
der Schalle, Curator of Molluscs, Museum in some locations, particularly in Big
of Zoology, University of Michigan, Drs. South Fork. A pH of 5.4 was found at
Aurèle La Rocque, J. B. Owen, B. L. Station 16 on this stream in November,
Ridley, Harold L. Harry, and Keith V. 1949. A few badly eroded shell fragments
Slack, all contributed to the collections were the only evidences of a former
used in the study. Dr. van der Schalle mussel population there.
loaned reference specimens and literature The upper stream bed is rocky, varying
from the Museum of Zoology. The majority from large areas of bedrock, such as those
of specimens taken during the study are just above the falls, to bars of small
now in the collections of the Museum of stones and gravel. Some sand occurs in
Zoology, University of Michigan. protected pockets and intermixed with
MUSSELS OF THE UPPER CUMBERLAND BASIN 429
3
Ô
to
cd
'
-)
S
§•
^
430 NEEL AND ALLEN
stones, but no extensive sand bars were dale

noted. In the area just below Cumberland 13. Cumberland River at Artemus
Falls sand pockets among large stones and 14. Cumberland River above
slabs contained great numbers of mussels, Pineville
many of which were quite large and very 15. Cumberland River at Wallins
old. Shingle bars are important naiad 16. Big South Fork at Yamacraw
habitats in the river below the falls, and 17. Big South Fork above Burn-
mussels were usually abundant in areas of side
shingle and gravel up to the vicinity of 18. Cumberland River below Lock
Pineville where they appeared to be deci- 21
mated by mine Rockcastle River
acids. 19. Cumberland River at Horse-
has a rocky bottom with scattered small shoe Bottom
shingle or shingle and gravel bars up to 20. Beaver Creek near mouth
the region of Laurel Fork. Sand and gravel 21. Cumberland River at Rowena
accumulations in pools and shingle and Ferry
gravel riffles yielded many mussels in 22. Cumberland River at Long
Laurel Fork; Laurel River had numerous Bottom (just below Wolf Creek
bed-rock areas and mussels occurred Dam)
among rather large stones (6-12in.diame- 23. Cumberland River at Neeleys
ter); and Beaver Creek contained numer- Ford
ous shingle and gravel bars. The lower
two miles of Big South Fork were included Figs. 67, 68 show some mussel bars in
in the Lock 21 pool; all workable bottoms the Cumberland below Lock 21, Fig. 2
were shingle or larger stones and long portrays the lower Rockcastle River, and
rocky rapids occupied about 2 1/2 miles Fig. 3 is a fairly typical stretch of the
of stream bed above the pool. Acid main stem above the falls. Views offered
coal mine waters have destroyed all by these photos are reasonably repre-
mussels in this stream from the region sentative of areas sampled.
of Yamacraw (Station 16) to the area just
above the mouth. Very few mussels were METHODS
found in the vicinity of the mouth. Col-
lections were made at the following 23 Mussels were collected by hand: probing
locations shown in Fig. 1. into various loose substrates with the
fingers, turning over large stones, digging
Station 1. Laurel Fork of the Rock- out sand pockets, robbing muskrat piles,
castle near McGee locating siphons, tracks or displaced in-
2. Laurel Fork of the Rockcastle dividuals. In the main stem below the
River falls collection by these methods was
3. Rockcastle River near Ardery possible only when and where water was
4. Rockcastle River above shallow enough to permit wading, usually
Livingston on shingle bars. These areas had dense
5. Rockcastle River at Billows mussel populations that were most readily
6. Rockcastle River near mouth sampled by probing through stones with the
7. Laurel River at Lily fingers. It was necessary to dig deeper
8. Laurel River at Keavey in colder weather, and one species,

9. Cumberland River below Lastena lata, always seemed to avoid the
Cumberland Falls uppermost layers. Cumberlandta mono-
10. Cumberland River above donta occurred only on those bars where
Cumberland Falls it could find the shelter of large stones.
11. Cumberland River at Suttons Muskrat piles were most common in

Mill Beaver Creek, on main stem bars at
12. Cumberland River at Gaus- Stations 19, 21 and 23, and in the lower
m^
'^
* --^^^^^î^rs ~-r
FIG. 2. Rockcastle River shortly above mouth near Mt. Victory, Kentucky. Station 6.
432 NEEL AND ALLEN
Rockcastle River. These rodents place of effort formerly expended along this line.
mussels on rocks and logs above the Wilson and Clark (1914) noted as many
water line and later return to dine on the as 200 pearlers working a single bed and
soft parts after the shells have opened. were shown some valuable pearls. Interest
Many living specimens and empty shells in this activity died shortly after their
fall into the water and are gradually trip down the river and pearling was
moved downstream by currents. Such unheard of in the late 1940' s. Conditions
areas often yield fine specimens of small in both the Rockcastle and Cumberland
and/or rare species. Rivers appear favorable for development
Quantitative aspects of a mussel popu- of pearls (most of our specimens con-
lation are difficult to evaluate with any tained slug pearls or baroques) but a
collection methods. It is usually im- great amount of time must be expended
possible to cover all regions of a sampled in location of a valuable specimen.
reach and the effectiveness of methods Pearling was probably abandoned after its
used is often difficult to ascertain. Col- unprofitable nature became evident. Local
lectors are therefore generally obliged to fishermen advised that Lampsilis ovata
use a system that is somewhat subjective. and Elliptio crassidens were considered
In this account clams are termed abundant the most valuable pearl mussels. We
if they were generally spaced within 12in. found slug pearls in almost all species
(20.5 cm) of each other, common if their collected (Fig. 69).
number per square meter was at least 4, Wilson and Clark (1914) predicted a
and rare if more poorly represented. great future for the mussel fishery of the
Species were considered abundant if they Cumberland, but little materialized in
formed 25% or more of an abundant popu- the upper reaches.
lation or 40% or more of a common
population, common if they comprised as SYSTEMATIC ACCOUNT OF NAIAD
much as 5% of an abundant population or OCCURRENCE
20% of a common population, and rare if FAMILY MARGARITANIDAE
less numerous. Larger species were
The species recovered in the present
seldom as closely grouped as were smaller
survey of the upper Cumberland are listed
ones and they were considered abundant
in Table l(p 429). In the following, details
or common if they formed slightly smaller
are given on each of the 59 species or
percentages than those listed.
forms found. A species that is probably
now extinct, Pegias fabula, is also
COMMERCIAL VALUE mentioned. All but 4 of these forms are
illustrated in Figures 4 to 66.
The Cumberland River has
upper
received little from the button
attention
Cumberlandia mondonta (Say) 1829. Figs.
industry. A number of indigenous species
4 and 8.
reputedly have nacre that is too hard for
button manufacture, and the more im- Confined to the main river below the
portant shell beds have been relatively falls; livesunder large stones and maybe
inaccessible. Areas mapped for fishing collected only when or where water is
by button companies in the early 1900's shallow enough to permit wading and
proved unprofitable and lay virtually un- moving of such stones; very poorly repre-
touched for 30 years prior to 1947. sented in muskrat piles. Living specimens
Until about 1915, local residents were collected only at Horseshoe Bottom
practiced pearling to a considerable extent (Station 19).
during slack agricultural seasons. This mussel is uncommon in the lower
Mussels stripped in search of pearlswere Cumberland or Tennessee, and is regarded
thrown out upon the banks and great piles as rare in the Ozarks and other regions
of crumbling shells offer mute evidence of the Mississippi Basin. Ortmann (1918)
CZ3
Ü
3
>
•iH
Pi
T3
cd
I—i
Ü
434 NEEL AND ALLEN
found it focally abundant in the upper vicinity of Laurel Fork.It occurred also
Tennessee, Clinch, and Holston Rivers, inBeaver Creek. Wilson and Clark (1914)
and reported many dead shells at Mussel reported it only from smaller streams.
Shoals in 1925. Wilson and Clark (1914) Common names: Three -ridge; Blue point.
did not find it in the upper Cumberland,
and encountered it only occasionally in the Megalonaias gigantea (Barnes) 1823.
lower river. Fig. 7.
Common name: Spectacle Case.
Abundant in the main stem below Burn-
side but absent in all tributaries and the
FAMILY UNIONIDA E area immediately below the falls. Wilson
and Clark (1914) failed to find it in the
SUBFAMILY UNIONINAE upper Cumberland.
Fusconaia flava (Rafinesque) 1820. Fig. Common name: Washboard.

9.
Quadrula cylindrica (Say) 1817. Fig.
Small flattened specimens were found 11.
in Beaver Creek (Station 20) but it was
not encountered elsewhere. Wilson and A rare shell in the upper Cumberland
Clark (1914) reported it only from the that exhibited no marked predilection for
East Fork of Stones River at Water hill, any type of environment, occurring in
Tennessee. Ortmann (1926) considered it both the main stem and smaller streams
native to smaller headwaters only. (Table 1).
Common name: Rabbits- Foot.

Fusconaia undata (Barnes) 1823.
This species was also taken only in Quadrula mentanevra Rafinesque 1820.
Beaver Creek; 3 small, flattened speci- Fig. 14.
mens were found. Wilson and Clark
A typical big river form, but found
(1914) rarely uncovered it and also
also in Beaver Creek and the lower Rock-
obtained only small shells.
castle River; missing, however, from the
Common name: Pigtoe. area just below the falls. It is a very
abundant mussel on the main stem bars,
Fusconaia subrotunda (Lea) 1831. Fig. but considered rare by Ortmann (1925)
6. and Wilson and Clark (1914). These
collectors could hardly have overlooked
A typical mussel of the large bars in such numbers as existed in 1947-49.
the main stem below the falls, but not a
particularly abundant one. Very large Common name: Monkey-face.
flattened specimens occurred in the Rock-
castle River below Mt. Victory. Reported Quadrula pustulosa (Lea) 1831. Figs.
from the Ohio, Green, Kentucky, and other 12 and 15,
streams of the Ohio Basin. Ortmann
Very abundant on big river bars and
(1925) found it only in the main stem of
also in lower reaches of the Rockcastle
the Tennessee.
River and Beaver Creek. The only
Quadrula present just below the falls.
Amblema costata Rafinesque 1820. Fig.
Cumberland River specimens are typical
10.
of the species, exhibiting the broad green
This species is characteristic of umbonal stripe, and losing pustules toward
smaller streams and was found in the the headwaters. Some specimens from
Cumberland only at Station 22. It was just below the falls were considerably
very common in the Rockcastle up to the flattened (Fig. 12).
FIG.
436 NEEL AND ALLEN
Common name: Warty back. Reported as rare in the Tennessee by

Ortmann (1918, 1925). May be
confused
Tritigonia verrucosa Raiinesque 1820. with Quadrula
pustulosa but lacks the
Fig. 23. green umbonal band.
This mussel is one of the most wide- Common name: Cumberland Pigtoe.
spread of the upper Cumberland system.
It was most numerous on the big river Pleurobema cordatum (Rafinesque) 1820.«
bars, but was also quite abundant in the Fig. 5.
Rockcastle. Many large specimens
showing marked sexual dimorphism were Four forms of this species were common
taken just below the falls. Wilson and in the upper Cumberland drainage. The
Clark (1914) noted a similar distribution. parent form, cordatum, was restricted to
the main stem where it was numerous
Common names: Pistol grip; buckhorn. on big shoals. It was very abundant in
the collections of Wilson and Clark (1914)
Cyclonaias tuberculata (Rafinesque) 1820. who considered it the most important
Fig. 13. commercial species in the river.
The specimens of this species
finest
The three distinct variants of P.
occurred in lower Rockcastle.
the It
cordatum are usually listed as trinomials
was present on some of the main stem by students of North American naiads,
although they often occur side by side
bars but was largely replaced there by
with the parent form. P. c. coccineum is
the variety granifera.
normally the only member of the complex
Common name: Purple warty -back. found in headwaters, but it often lives in
downstream areas frequented by P.
Cyclonaias tuberculata granifera (Lea) cordatum. The other two forms (P. c.
1838. Fig. 16. plenum, and P. c. pyramidatum) seemingly
The inflated big river form of have the same habitat preferences as the
tuber-
culata was found only on big river bars. parent form. The trinomial system is a
convenience, and this complex has long
Common name: Purple warty-back. been a part of our mussel lore, but no
claims are made for the validity of the
Plethobasu^ cyphyus (Rafinesque) 1820. subspecific rank.
Fig. 19.
Common name: Ohio River Pigtoe.
A rare species in the upper Cumberland
taken only at Stations 19 and 22. It was
also rare at the time of Wilson and Clark
Pleurobema cordatum plenum (Lea) 1840.
Fig. 22.
(1914) who reported it had very hard
nacre. Although called a rare form of P.
Common names: Bullhead; Sheepnose. cordatum by Wilson and Clark (1914) and
by Ortmann (1925) P. c. plenum was a
Plethobasus cooperianus (Lea) 1834. Fig.
common shell in the Cumberland at the
time of our collections, and was found
17.
in goodly numbers on all main stem bars.
Also rare in the upper Cumberland in This variety appears intermediate between
the late 1940's but reported as common by the typical P. cordatum and P. c. pyrami-
Wilson and Clark in 1914. A very at- datum. It is slightly more rounded than
tractive species with pale pink diffused P. pyramidatum with a less distinct
through the white nacre. The perio- sulcus.
stracum is pale brown, usually less
pustulate than the specimen illustrated. Pleurobema cordatum pyramidatum (Lea)
438 NEEL AND ALLEN
1831. Fig.21. Elliptio crassidens (Lamarck) 1819, Fig.
25.
This, the highest, shortest form of the
cordatum group, occurred only on the big Primarily a main stem species that was
river bars. Its extreme form is un- not present in the Rockcastle River above
mistakable, but some specimens are Mt. Victory (Station 6), in Beaver Creek,
difficult to separate from P. c. plenum. or the Lower Laurel River, and rarely
It is tempting to group both forms under occurred in Big South Fork; however, 2
P. pyramidatum, which has priority, specimens were picked up in the Laurel
but they are generally distinct and perhaps River at Lily under marked headwater
deserve to retain separate identities at conditions. This was a puzzling oc-
present. currence and may reasonably be regarded
as accidental and as an aftermath of some
Pleurobema cordatum coccineum{Conra.d) unusual condition. It shows, however, that
1836. Fig.20. no general pattern may be taken for
granted. E. crass ¿dens was very abundant
This variety occurred in the main river
in the main river below the falls and
but attained its maximum concentrations
formerly was highly regarded as a pearl
in the Rockcastle, clearly demonstrating
species. Larger examples had thick shells
its general relationship to headwaters.
and were usually much eroded. Nacre
was usually dark pink but several white
Pleurobema oviforme (Conrad) 1834.
nacred specimens were noted.
Fig. 18.
Common name: Elephant ear.
Ortmann (1918, 1924, 1925) assumed this
species had replaced the Ohio valley form
Elliptio dilatatvs (Rafinesque) 1820. Fig.
P. clava in the Cumberland and Tennessee.
27.
We found it only in Beaver Creek and
in the extreme headwaters of the Rock- This mussel occurred at every station
castle River in Laurel Fork. Ortmann except No. 7, and was the most widely
described 2 forms in the Tennessee, a distributed and abundant mussel in the
flattened headwater P. o. argenteum, and an upper Cumberland in the late 1940's.
inflated big river P. o. holstonense . Its Wilson and Clark (1914) reported it much
appearance and occurrence in the upper less abundant than E. crassidens and
Cumberland suggest it has definite af- poorly represented in a great number of
P. c. /
finities to P. cordatum, coccineum. In the
lower and middle Rockcastle River PZewr-
obema is represented only by the variety
and the P. speci-
mens found in Laurel Fork appear to be
an extreme headwater form of cocineum.
Wilson and Clark (1914) did not find P.
their collections.
This
Falls.
species
Its predominance at
the time of our study was one evidence of
considerable change in the mussel fauna
over the preceding 36 years.
clearly demonstrates
isolation of mussels above Cumberland
Everywhere below the falls its
oviforme below Burnside and thought it nacre was uniformly purple, but speci-
a small stream type. Ortmann's dis- mens from above, were partly white or
coveries in the Tennessee suggest wholly white inside.
different relationships, and indicate that
additional collections should be carried
Common names: Lady finger; spike.
out there before any attempt is made to

Lastena lata (Rafinesque) 1820. Figs.
revise the position of P. oviforme. If it
24 and 28.
were known only from our collections in
the Cumberland it should certainly be A rare Mississippian form that was
considered a form of P. cordatum that found in fair numbers on main stem shoals
replaces P. coccineum in small upper below the falls. It burrowed deeply among
reaches. the small stones and gravel and was
FIG. 17. Plethobasus cooperianus (Lea)

FIG. 18. Pleurobema oviforme (Conrad)
FIG. 19. Plethobasus cyphus (Rafinesque)
FIG. 20. Pleurobema cordatum. coccineum. (Conrad)
FIG. 21. Pleurobema cordatum pyramidatum (Lea)
FIG. 22. Pleurobema cordatum plenum (Lea)
440 NEEL AND ALLEN
captured because of digging down with here will stimulate additional search as
the hands. Beautifully rayed specimens opportunity permits.
were taken (Fig. 24). Ortmann (1918,
1924, 1925) rarely found it in the Strophitus rvgosus (Swainson) 1822. Fig.
Tennessee System, and Wilson and Clark 26.
(1914) seldom collected it in the Cumber-

A common mussel on the main stem
land. They suggested that its rarity in
bars, but also present in the Laurel and
their collections may have simply repre- Rockcastle Rivers and Beaver Creek. It
sented the ineffectiveness of crowfoot bar s was most abundant at Station 19. S.
for this species. rvgosus and Elliptio crassidens were the
only mussels found in the upper Laurel
SUBFAMILY ANODONTINAE River. Our specimens had a dark brown
or black periostracum and salmon and
Lasmigona costata (Rafinesque) 1820. bluish nacre. It was seldom seen by
Fig. 307 Wilson and Clark (1914) but they did find
preferred headwater it above the falls. It now appears to be
This species
streams and occurred main stem
in the restricted to the region below Cumberland
only at Station 19. It was most abundant Falls. Ortmann found it abundantly in the
in the Rockcastle River. Ortmann (1918, upper and rarely in the lower Tennessee
1924, 1925) reported it as a small stream River.
type in the Tennessee drainage. Wilson Common name: Squaw foot.
and Clark (1914) rarely found it in the
upper Cumberland and not at all in the SUBFAMILY LAMPSILINAE
lower river. Our Rockcastle River speci-
mens contained numerous baroque pearls. Ptychobranchus fasciolaris (Rafinesque)
1820. Fig. 32.
Common name: Fluted shell.
This mussel was almost as widespread
Alasmidonta marginata {Sdiy) 1819. Fig. as Elliptio dilatus (Table 1). Specimens
29. taken just below the falls (Station 9) were
very large and of great age. Shells had
A rather uncommon mussel in the a white thick nacre and would apparently
Cumberland. (Table 1). Our specimens have some commercial value. Wilson
were small and attractively rayed. Wilson and Clark (1914) found it commonly, but
and Clark (1914) considered it rare, and never noted it to be abundant. In the late
Ortmann (1924, 1925) reported it as rare 1940's it was abundant on many large
in theTennessee. All three authors thought river bars and in the Rockcastle River.
it a small stream form, but we found it
to be more abundant in the main stem. Common name: Kidney shell.
Common name: Elk toe. Obliquaria reflexa Rafinesque 1820. Fig.

31.
Pegias fabula (Lea) 1836.
Considered abundant by Wilson and
This species was found by Wilson and Clark (1914) but rare in our collections.
Clark (1914) in the Rockcastle River near We found it at all Cumberland River
Livingston, but extensive searches of the stations below the falls, in Beaver Creek,
same area during 1948 and 1949 failed to and the lower Rockcastle River. Ortmann
uncover it. It was reported as rare in reported it from Big South Fork (1924)
the Tennessee by Ortmann (1918, 1925). but it now appears to be extinct in lower
This is apparently a headwater form and reaches of that stream.
may now be extinct in the Cumberland
system. It is hoped that mention of it Common name: Three-horned warty back.
FIG. 23. Tritigonia verrucosa (Rafinesque)

FIG. 24. Lastena Zaía(Raíinesque)
FIG. 25. Elliptio crassidens (Lamarck)
FIG. 26. Strophitus rugosus (Swainson)
FIG. 27. Elliptio dilatatus (Rafinesque)
FIG. 28. Lastena lata (Rafinesque)
FIG. 29. Alasmidonta marginata (Say)
442 NEEL AND ALLEN
Cyprogenia irrorata (Lea) 1828. Fig. them (Station 9). It also occurred in the
33. lower Rockcastle and in Beaver Creek.

Those from the tributaries were flatter
This species was not commonly found than the main stem specimens. Flattened
and then only on the main stem bars. headwater forms are often referred to the
Wilson and Clark (1914) also found it variety O. s.lens (Lea).
infrequently. Ortmann considered it an
interior basin form that was widespread Actinonaias carinata gibba (Simpson) 1900.
but not abundant in the Cumberland and Fig. 38.
Tennessee.
This shortened, compressed form of the
Dromus dromas (Lea) 1834. Fig. mucket has been found only in the
35. Tennessee and Cumberland systems. Ort-
mann (1925) reported typical A. carinata
This isa true Cumberlandian form which
from Duck River and A. c. gibba in the
we found only on the main stem bars main Tennessee. Examples of A. carinata
below the falls, but in considerable resembling A, gibba have been reported
numbers. The shell is very hard, with from the Green and Ohio Rivers, but it
some pinkish but mostly white nacre, and seems generally agreed that the true
is reputedly unfit for button manufacture.
southern mucket (gibba) is strictly a
The flattened headwater form, D. caper- Cumberlandian form. This was a very
atus (Lea) 1845 formerly occurred in
common mussel in the main stem below
Big South Fork (Wilson and Clark, 1914)
the falls. It was found up to Mt. Victory
but we believe it is now extinct in the
in the Rockcastle River, and rarely in
upper Cumberland. D. dromas is an
Big South Fork above Burnside. Wilson
attractive naiad with neatly rayed
and Clark (1914) considered it the most
epidermis. Young individuals may have
valuable button shell in the river.
a hump near the umbones, but this area
is usually eroded in older specimens. It Common name: Southern mucket.
is apparently more abundant in the
Cumberland than in the Tennessee (Ort- Actinonaias pecterosa (Conrad) 1834.
mann 1918, 1925). Figs. 39 and 42.
Common name: Camel shell. This Actinonaias is also a true Cumber-

landian form, but it is a headwater rather
Obovaria retusa (Lamarck) 1819. Fig. than a big river species. It occurred very
34. rarely above the falls (one specimen at
Station 11), was very abundant just below
This species was fairly common on the
the falls (Station 9) and in the Rockcastle
big stream bars below the falls but it
River up to Laurel Fork; it was also
did not enter any of the tributaries. Some
present in Big South Fork and Beaver
unusually large specimens were taken
Creek. This species is longer and lower
from Station 23. Nacre is dark salmon
than A. carinata gibba (Figs. 39, 42) and has
or purple in beak cavities. Reported as
distinctive blotched or chevroned rays. It
rare by Wilson and Clark (1914) and as
occurred with A. c. gibba at Stations 9,
rare in the Tennessee by Ortmann (1918,
6, and and shows definite affinities to
16,
1924, 1925).
the latterform. An exhaustive search
was made for intergrades, and although
Obovaria subrotunda (Rafinesque) 1820.
some specimens from Station 9 nearly
Fig. 37.
qualified, the hunt was unsuccessful. Ort-
This species was very common in the mann (1918) noted that A. pecterosa was
Cumberland below the falls, although it most abundant where A. c. gibba began to
did not go up to the region just below disappear in the upper Tennessee system.
31
34 36
FIG. 30. Lasmigona costata (Rafinesque)

FIG. 31. Obliquaria reflexa Rafinesque
FIG. 32. Ptychobranchus fasciolaris (Rafinesque)
FIG. 33. Cyprogenia irrorata (Lea)
FIG. 34. Obovaria retusa (Lamarck)
FIG. 35. Dromus dromas (Lea)
FIG. 36. Truncilla donaciformis (Lea)
444 NEEL AND ALLEN
It was abundantin Cumberland headwater s Common name: Paper shell.

at the time of Wilson and Clark's (1914)
study. Leptodea laevissima (Lea) 1830. Fig.
43.
Truncilla donaciformis (Lea) 1828. Fig.
Reported as rare by Wilson and Clark
36.
(1914) and as rare in the Tennessee by
A
rare species in the Cumberland that Ortmann (1925). We found it at Stations
was found only at Stations 18 and 19. 6, and 20 in 1948 and 1949. It is not
19,
Wilson and Clark (1914) found it limited exclusively a big river species but it is
in distribution and it was rarely noted in rare in the upper Cumberland.
the Tennessee system by Ortmann (1924,
Common name: Paper shell.
1925). Our specimens were small and
quite eroded.
Leptodea leptodon (Rafinesque) 1820.
Fig. 41.
Truncilla truncata Rafinesque 1820. Fig.
40. A very rare shell throughout the interior
basin, and found only at Horseshoe Bottom
Rather uncommon, but more numerous
(Station 19) in 1948. Rarely found by
and widespread than T. donaciformis
Wilson and Clark (1914) and seldom taken
(Table 1). It was most abundant in the
by Ortmann in the Tennessee (1918, 1924,
main stem below the falls but was also
1925).
present in Beaver Creek. Reported un-
common by Wilson and Clark (1914) and Proptera alata (Say) 1817. Fig.50.
as rare in the Tennessee and Cumberland
by Ortmann (1925). This species occurred in all regions
below the falls except in the Laurel River.
Common name: Deer toe.
It was very common on the main stem
bars where it attained great size. It was

Plagióla lineolata Rafinesque 1820. Fig.
not thought especially common by Wilson
45.
and Clark (1914), but Ortmann found it
Common in the Cumberland below the to be common and abundant in the
falls,but not just below the falls (Station Tennessee drainage.
9). Very abundant on big bars below
Common name: Purple heel-splitter.
Lock 21 and in the mouths of Beaver Creek
and Rockcastle River. It attained very
Carunculina moesia (Lea) 1841.
large size at Stations 19 and 23. It was
considered common by Wilson and Clark This is a Cumberlandian form that is
(1914) was reported as rare in the
but closely allied to glans of other parts
Tennessee by Ortman (1918, 1924, 1925). of the interior basin. It was found only
It was once thought a good button shell in at Stations 19 and 21 on the main stem,
the Cumberland. but Wilson and Clark (1914) reported it
only from tributaries. This small species
Common name: Butterfly.
is easily overlooked, and it is not always
possible to exercise the particular
Leptodea fragilis (Rafinesque) 1820. Fig.
screening techniques its collection
48.
demands.
This mussel was fairly common on
big stream bars and occurred in Beaver Medionidus conradicus (Lea) 1834. Fig.
Creek. Wilson and Clark (1914) and Ort- 46.
mann (1918, 1924, 1925) thought it a big
river species not found in smaller The type locality of this species, the
streams. region just below the falls (Station 9),
37
39
\,
42
FIG. 37. Obovaria subrotunda (Rafinesque)

FIG. 38. Actinonaias carinata gibba (Simpson)
FIG. 39. Actinonaias pecterosa (Conrad)
FIG. 40. Truncilla trúncala Rafinesque
FIG. 41. Leptodea leptodon (Rafinesque)
FIG. 42. Actinonaias pecterosa (Conrad)
446 NEEL AND ALLEN
contained it in great numbers in September Laurel River. It ascended the Rockcastle
1948. It did not occur at any other up into Laurel Fork. Wilson and Clark
main stem station, but was abundant in (1914) found it only in the upper Cumber-
Beaver Creek and the Rockcastle River, land. Ortmann (1918, 1925) thought it
ascending the latter far up into Laurel rare in the Tennessee. The periostracal
Fork. It was found mainly in small color of this species is dark green. A
streams by Wilson and Clark (1914) and characteristic ray pattern appears on the
in small tributaries of the Tennessee by specimen shown in Fig. 52.
Ortmann (1918,1924, 1925), who con-
sidered it one of the most characteristic Micromya vanuxemensis (Lea) 1838. Fig.
Cumberlandian "types". This genus has 44.
not been reported from other parts of the
Mississippi Basin. We found this species alive only in
Beaver Creek. It shows affinities to M.
Micromya nebulosa (Conrad) 1834. Fig. lienosa of the Alabama River system, but
49. is distinct from that species. It does
not develop the pointed posterior dorsal

This species was found only in the shell margin that characterizes M. lienosa
Rockcastle River and in the main stem and it apparently attains a smaller size.
at Station 9. It is a very variable mussel
Our specimens had a dark brown perio-
and specimens found in the upper Cumber- stracum and red nacre. Empty shells
land represent a rather unusual form. It
were found in Big South Fork.
was not reported by Wilson and Clark
(1914). Ortmann (1924, 1925) considered
it a speciesmost characteristic of smaller Ligumia recta latissima (Rafinesque)
streams. Its type locality is the Black 1820. Fig. 51.
Warrior River, Alabama, but Ortmann This species occurred at all stations
(1925) preferred to call it a Cumberlandian below the falls except those in the Laurel
"type" although aware of its occurrence in
River. It was very abundant and almost
the Alabama River system.
reached Laurel Fork in the Rockcastle,
.
but was not considered abundant by Wilson

Micromya picta (Conrad) 1834. Fig. and Clark (1914). Ortmann reported it
47. common in the Tennessee. The nacre of
our specimens was very attractive, fading
Found only in Beaver Creek and Rock- from salmon pink in the beak cavities to
castle River. It attained very large size
white or bluish white near margins.
in the Rockcastle and went up into Laurel
Fork. Wilson and Clark (1914) did not Common names: Long John; Black sand
find it in the upper Cumberland. This shell.
is a true Cumberlandian form also
reported from the Tennessee (Ortmann Lampsilis fasciola Rafinesque 1820. Fig.
1924, 1925). 54.
Common name: Painted shell. One of the few species occurring above
the falls, this mussel was also common
below the falls, particularly in the Rock-
Micromya trabalis (Conrad) 1834. Fig.
castle River. However, it was not present
52.
at all stations below the falls (Table 1).
Another Cumberlandian form and one It is a common interior basin form that
that resembles M. fabalis (Lea) of othe: reportedly increases in number near the
parts of the interior basin. We found i headwaters of all streams it inhabits.
to be common in the upper Cumberland, It was very abundant in the Cumberland
absent only from Big South Fork and the above the falls.
FIG. 47. Micromya picta (Conrad)

FIG. 43. Leptodea laevissima (Lea)
FIG. 44. Micromya vanuxemensis (Lea) FIG. 48. Leptodea fragilis (Rafinesque)
FIG. 45. Plagióla lineolata Rafinesque FIG. 49. Micromya nebulosa (Conrad)
FIG. 46. Medionidus conradicus (Lea) FIG. 50. Proptera alata (Say)
448 NEEL AND ALLEIN
Lampsilis orbiculata (Hildreth) 1828. the falls in 1948 and 1949.
Fig. 5J.
Common name: Pocketbook.
This is another Mississippian species
that was still common in the main stem Dysnomia brevidens (Lea) 1831. Figs.
below the falls but apparently less abundant 59 and 62.
than at the time of the Wilson and Clark
A very common big river form below
(1914) survey. It slightly resembles A.
Cumberland Falls and present also in the
carinata gibba and is a very distinctive
lower Rockcastle River and Beaver Creek.
species of Lampsilis. Ortmann (1918,
Wilson and Clark failed to find it in the
1925) found it relatively uncommon in the
main river and reported it as very common
Tennessee.
in Big South Fork, where it was extinct
in the late 1940' s. Wilson and Clark
Lampsilis ovata (Say) 1817. Fig. 55.
(1914) collected only dead specimens.
This typical big river species of Lamp- Ortmann (1918, 1924, 1925) foundit rarely
silis occurred only below the falls and in in the Tennessee. Prior to closure of
the mouth of Beaver Creek. It has an at- Wolf Creek Dam it was abundant at Station
tractive light yellow epidermis, a sharp 19 and common at other main stem col-
posterior ridge, and green rays. Wilson lecting sites below the falls.
and Clark (1914) encountered it more
commonly in the upper than in the lower Dysnomia capsaeformis (Lea) 1834.
river; Ortmann (1918, 1924, 1925) found it
This small thin shelled mussel was
preferred larger rivers in the Tennessee
found at scattered localities including the
basin. We found it a good source of slug
main stem below the falls, Rockcastle
pearls (baroques), and learned it was once
River, Big South Fork, and Beaver Creek.
a favorite of pearlers.
It resembles D. florentina but may be
Common name: Pocketbook. distinguished by the solid green color on
the posterior expansion of the female shell.
Lampsilis ovata ventricosa (Barnes) 1823.
Also a member of the Cumberlandian
Fig. 58.
group, considered by Ortmann (1925) to
A
headwater form that replaced L.ot>aia be more numerous in the Tennessee River.
in theCumberland above the falls, the
Rockcastle and Laurel Rivers, and Big Dysnomia florentina (Lea) 1857. Figs..
South Fork. It is quite similar to L. ovata 56 and 57.
but lacks a sharp posterior ridge and,
May be separated from the foregoing by
typically, is slightly more elongate in
the absence of the solid green color on
outline. Specimens from smaller head-
the posterior expansion of the female
waters, Laurel River and Laurel Fork of
shell, and more numerous denticulations
the Rockcastel, were considerably short-
on the margin of the expansion. Found on
ened and stunted. It was associated with L.
most main stem bars, abundantly at Station
ovata only at Station 19, and this co-
19. Reported as rare by Wilson and Clark
existence appeared unusual. Ortmann
(1914) and as rare in the Tennessee,
(1918, 1924, 1925) reported its occurrence
except at Mussel Shoals, by Ortmann (1918,
with L. ovata in larger rivers, and stated
1925).
that it went upstream beyond L. ovata and
became a pure race. With the one exception
Dysnomia florentina walkeri (Wilson and
at Station 19 L. ovata ventricosa was a
Clark) 1914. Figs. 64 and 65.
pure race in the Cumberland. Wilson
and Clark (1914) called this form L. ventri- A larger compressed type that replaces
cosa, apparently unaware of its relation D. florentina in headwaters. The epi-
to L. ovata. It was very abundant above dermis is yellower than that of the typical
449
MUSSELS OF THE UPPER CUMBERLAND BASIN
55
FIG. 51. bigamia recta latissima (Raiinesque)

FIG. 52. Micromya trabalis (Conrad)
FIG. 53. Lampsilis orbiculata (Hildreth)
54. Lampsilis fasciola
Raiinesque
FIG.
FIG. 55. Lampsilis ovata (Say)
FIG. 56. Dysnomia florentina (Lea) female
FIG. 57. Dysnomia florentina (Lea) male
450 NEEL AND ALLEN
D. florentina First found by Wilson
. itsrange extended down into Tennessee,
Clark (1914) in small tributaries of the but they did not find it in the lower
Tennessee loop of the Cumberland and Cumberland.
later found in the Tennessee River by
Common name: Snuffbox.
Ortmann (1918, 1924) who decided it was
a variety of D. florentina. It was found
DISCUSSION
by us only inBeaver Creek, but it formerly
occurred in Big South Fork.
Our collections in the upper Cumberland
system netted 30 genera with 59 species
Dysnomia haysiana (Lea) 1833. Fig.
or forms, of which 16 belong to Ortmann' s
60.
typical "Cumberlandian* forms (see Table
This species is one of the most at- 4). These mussels are listed in Table 1,
tractive members of the genus, being small which shows their distribution and relative
and solid with a polished, brown-rayed abundance, in the order adopted in the
periostracum. We found one specimen in Systematic Account.
Big South Fork where it was formerly The mussels collected had been sub-
locally abundant. Its type locality is the jected to considerable abrasion and some
Cumberland River. acid action and only exceptional shells
escaped erosion of the umbonal areas.
Common name: Acorn.
Many old specimens had very thick shells
and were worn down to more elongated
Dysnomia lewisi (Walker) 1910. Figs.
shapes. Numerous individuals had sand
61 and 66.
blisters and many had lost almost all the
Ortmann (1918) considered this species periostracum.
the rarest Dysnomia. Wilson and Clark Comparison of our findings with those
did not mention it in 1914. We secured a of Wilson and Clark (1914) in the same
female at Station 21 and a male at Station area, in 1911, indicated that rather marked
22. The species is unmistakable and can changes in occurrence, distribution and
be easily recognized by characters shown abundance had occurred in the 36-38 years
in the Figs. It is a Cumberlandian type following their survey. These authors
and shows some affinities to Dysnomia reported 60 forms, but the actual dis-
foliataof the Ohio River, which is now crepancy involves 7 species, since we
assumed extinct. found 3 species they did not report:
Cumberlandia monodonta, Micromya
Common name: Leaf shell.
nebulosa and Dysnomia lewisi, while they
reported 4 species: Alasmidonta minor,
Dysnomia sulcata (Lea) 1829.
Dysnomia arcaeformis, Pegias fabula and
This Dysnomia is rare in all parts of Quadrula intermedia that we failed to
the interior basin where its distribution collect.
appears to center in the Ohio and Wabash Up to the Cumberland Falls the usual up-
Rivers (Ortmann, 1925). We secured stream decline of number of species was
only one specimen, at Neeleys Ford. evident (down to 15 at Station 9 from more
than 40 at Station 21). The falls were
Common name: Cats claw.
an effective barrier to most species and
a total of only 6 have been recorded above
Dysnomia triquetra (Rafinesque) 1820.
them. In the present survey only 4 forms
Fig. 63.
occurred above the idlls-.Lampsilis fas ci-
This abundant species of the Mississippi óla, L. ovata ventricosa, Elliptic dilatatus
basin was common along the main stem and Actinonaias pecterosa. The first 3
below Lock 21, and occurred in Beaver had apparently been in residence there
Creek. Wilson and Clark (1914) noted that for many generations while the last, A.
FIG. 58. Lamps ¿lis ovata ventricosa (Barnes)

FIG. 59. Dysnomia brevidens (Lea) male
FIG. 60. Dysnomia Imysiana (Lea)
FIG. 61. Dysnomia lewisi (Walker) female
FIG. 62. Dysnomia brevidens (Lea) female
FIG. 63. Dysnomia triquetra (Rafinesque)
FIG. 64. Dysnomia florentina walkeri (Wilson and Clark) male
FIG. 65. Dysnomia florentina walkeri (Wilson and Clark) female
FIG. 66. Dysnomia lewisi (Walker) male
452 NEEL AND ALLEN
TABLE 1. Distribution and abundance^ of 59 naiad species found at 23 stations in the upper Cumberland
system.
Species Station 5
10
Cumberlandia monodonta
Fusconaia flava
Fusconaia undata
Fusconaia subrotunda
Amblema costata
Megalonaias gigantea
Quadrula cylindrica
Quadrula metanevra
Quadrula pustulosa
Tritigonia verrucosa
Cyclonaias tuberculata
Cyclonaias t. granifera
Plethobasus cyphyus
Plethobasus cooperianus
Pleurobema cordatum
Pleurobema cordatum plenum
Pleurobema c. pyramidatum
Pleurobema c. coccineum
Pleurobema oviforme^
Elliptio crassidens
Elliptio dilatatus
Las tena lata
Lasmigona costata
Alasmidonta margina ta
Strophitus rugosus
Ptychobranchus fasciolaris
Obliquaria reflexa
yprogenia irr ora ta
Dromus dromas
Obovaria retusa
Obovaria subrotunda
Actinonaias carinata gibba
Actinonaias pecterosa^
Truncilla donaciformis
Truncilla trúncala
Plagióla lineolata
Leptodea fragilis
Leptodea laevissima
Leptodea leptodon
Proptera alala
Carunculina moesta
Medionidus conradricus^
Microw.ya nebulosa
Micromya picta
Microm.ya trabalis^
Micromya vanuxemensis^
Ligumia recta latissima
Lampsilis fasciola
Lampsilis orbiculata
Lampsilis ovata
Lampsilia o. venlricosa
Dysnomia brevidens^
Dysnomia capsaeformis"
Dysnomia florentina^
Dysnomia /. walkeri
Dysnomia lewisi^
Dysnomia liaysiana^
Dysuorr.iu sulcata
Dysnomia trique tra
'â, abundant; c, common; r, rare.

5See
cc Fig.
IB- '•
J. 1.
6" Cuniberlandian" typ
Table 1. (continued)
S tatio n
11 12 13 14 15 16 17 18 19 20 21 22 23
454 NEEL AND ALLEN
pecterosa, which is the only true "Cumber- years before, but the changes listed appear
landian" form, was taken in only one clear cut from conditions described by
specimen at Station 11 and appeared to Wilson and Clark and differences in
be a recent immigrant. Wilson and Clark prevailing groups listed below at least
(1914) listed Alasmidonta minor andSiro- seem valid. Circumstances responsible
phitus rugosus in addition to the 3 well for such changes are difficult to ascertain.
established species we found above. the Decline in mussel and pearl fishing may
falls. All species recorded above the have been influential in some cases for
barrier were also common or abundant the increase of some species, and pol-
inmany areas below it. lution, particularly by coal mine acids,
Mussels were rare only in Big South has had demonstrable effects in decline
Fork and the Laurel River. They were of others. Fluidity in dynamics of the
abundant on big bars in the main stem mussel population may not be ignored, and
below Lock 21 (Station 18), not quite as some variations observed may perhaps
numerous above the falls and in the Rock- illustrate a natural trend over 36 years.
castle River and only common in Beaver Ortmann's records and studies led him
Creek. The region just below the falls to conclude that the Cumberlandian naiad
TABLE 2. Comparison of the 4 most prevalent naiad species of the 2 surveys of the
upper Cumberland, in descending order of abundance.
Wilson and Clark

FIG. 67. Shingle bar at Neeleys Ford. Station 23.
FIG. 68. Collecting at Horseshoe Bottom. Station 19.

456 NEEL AND ALLEN
succumbed to the effects of acid mine the disappearance of Alasmidonta minor
waters in that river. probable that
It is and Strophitus rugosus from above the
some of these dysnomias are now extinct. falls.
Mine drainage may also be involved in
TABLE 3. Comparison of the different naiad species most common in the 2 surveys
of the upper Cumberland.
Wilson and Clark

FIG. 69. Baroque or slug pearls from Cumberland and Rockcastle River mussels.
TABLE 4. Cumberlandian naiad fauna as defined by Ortmann.
Actinonaias carinata gibba (Simpson)" Fusconaia bamesiana (Lea)^

Actinonaias pecterosa (Conrad)'^ Fusconaia bamesiana bigbyensis (Lea)^
Alasmidonta minor (Lea)° Fusconaia bamesiana tumescens (Lea)^
Carunculina moesta (Lea)'^ Fusconaia cuneolus (Lea)^
Conradilla caelata (Conrad)^ Fusconaia cuneolus appressa (Lea)^
Dromus dromas (Lea)'^ Fusconaia edgariana (Lea)^
Dromus dromas caperatus (Lea) Fusconaia edgariana análoga (Ortmann)^
Dysnomia arcaeformis (Lea) Lasmigona holstonia (Lea)9
Dysnomia biemarginata (Lea) Lexingtonia dolabelloides (Lea)^
DysnomÍ2 brevidens {Lea.) Lexingtonia dolabelloides conradi
Dysnomia capsaeformis (Lea) '
(Vanatta)9
Dysnomia florentina (Lea)'^ Medionidus conradicus (Lea)"^
Dysnom.ia florentina walkeri (Wilson and Micromya nebulosa (Conrad) '»^^^
lark) '7 Micromya picta (Conrad) '
Dysnom.ia haysiana (Lea) '

Micromya trabalis (Conrad) '
Dysnom,ia lewisi (Walker)'

Dysnomia lenior (Lea)
Dysnomia torulosa propinqva. (Lea)^
Dysnomia turgidula (Lea)
'^Found in present sèrvey of the upper Cumberland.

^Found also in Kentucky River system.
9 Reported only from the Tennessee River.
,
Micromya vanuxemensis
Pegias fabula (Lea)
(Lea)"^»^^
Pleurobemxi oviforme (Conrad)"^

Pleurobema oviforme argenteum (Lea)^
oviforme holstonense (Lea)^
1^ Found also in Alabama River system.

458 NEEL AND ALLEN
ORTMANN, A. E. and BRYANT WALKER, WILSON, C. B. and H. W. CLARK, 1912.
1922. Onthe nomenclature of certain Mussel beds of the Cumberland River
North American naiades. Occ. Pap. in 1911. Dept. Comm. Bur. Fish., Econ.
Mus. ZooL, Univ. Mich. No. 112: 1-75. Circ. No. 1: 1-4.
WALKER, BRYANT, 1911. Notes on the , 1914. The mussels of the
distribution of Margaritana monodonta Cumberland River and its tributaries.
Say. Nautilus 25: 57-58. Bur. Fish. Doc. 781: 1-63.
ZUSAMMENFASSUNG
DIE NAIAD ENFAUNA DES OBEREN CUMBERLANDBECKENS VOR SEINER STAUUNG
In den Jahren 1947-1949, kurz vor der Entstehung des Cumberland-Stausees durch
die Errichtung des Wolf-Creek-Dammes, stellten die Autoren Untersuchungen Ober die
Muscheln des oberen Cumberlandflussystems (in Kentucky, V. S.A.) an, deren Ergebnisse
hier mit denjenigen einer vorherigen, aus dem Jahre 1911, verglichen werden. In
unserer Studie fanden wir 59 verschiedene Arten, von denen 16 der den Cumberland-
und Tennessee- Flüssen eigenen, sogenannten "cumberländischen» Gruppe, angehörten.
Oberhalb des Cumberlandfalles waren nur 4 Arten (einschliesslich einer einzigen
cumberländischen) vertreten, doch waren diese keineswegs auf jene Gegend beschränkt.
In der früheren Studie waren zwar 60 Arten zitiert worden, doch stimmten davon nur
56 mit den unsrigen überein, da 3 der später anwesenden vorher nicht gefunden worden
waren während 4 der früher angegebenen später fehlten. In den 36 Jahren zwischen den
beiden Untersuchungen sind offensichtlich im oberen Hauptfluss und in seinen dortigen
wichtigeren Nebenflüssen verschiedene Muschelarten durch saure Abwässer aus
Kohlengruben dezimiert worden. So wurde z.B. die Gattung Dysnomia aus dem Gebiete
ihres ehemals vielfältigsten Vorkommens praktisch vertilgt. Andere Arten an anderen
Standorten hingegen waren häufiger vertreten als zuvor.
Die beobachteten ausgesprochenen Veränderung en in der Verbreitung und relativen
Häufigkeit mehrerer Arten lassen sich nur teilweise durch den Niedergang der Perl-
fischerei oder durch säurehaltige Abwässer erklären. Auch wurde das obere Cumber-
landgebiet weitgehend von der Perlmutterknopfindustrie verschont, so dass die Gründe
für manche der auffälligen Schwankungen bis heute unklar sind,
RESUME
LA FAUNE NAÏADE DU BASSIN SUPERIEUR DU FLEUVE CUMBERLAND
AVANT SON ENDIGUEMENT
Pendant les années 1947-1949, juste avant la formation du lac Cumberland par
l'érection du barrage de Wolf Creek, les auteurs ont fait une étude de la population
naiade du bassin supérieur du Cumberland (Kentucky, E. U. A.) dont les résultats sont
ici comparés à ceux d'une étude antérieure, en 1911, de la même région. Au courant
de l'investigation récente nous avons trouvé 59 formes naiades distinctes, dont 16
appartenaient au groupe "cumberlandais" spécial qui n'existe que dans les fleuves
Cumberland et Tennessee. "Quatre espèces seulement (comprenant une seule cumber-
landaise) avaient surmonté les chutes Cumberland, mais aucune d'entre elles n'était
restreinte à cette région seulement. Dans l'étude précédente le nombre d'espèces
citées était de 60, mais il n'y avait que 56 qui coincidaient, car 3 des espèces trouvées
dans la seconde étude manquaient alors, tandis que 4 espèces trouvées alors étaient
absentes plus tard. Dans les 36 années depuis la première investigation, certaines
naiades ont été décimées dans le tronc principal supérieur du neuve et dans ses branches
majeures régionales par les eaux acides provenant des mines à charbon. Ainsi, le
genre Dysnomia a été presque totalement éliminé de l'aire de sa plus grande variété
antérieure. D'autre part nombre d'espèces ont gagné en prévalence ailleurs.
Ces changements prononcés en distribution et abondance relative de certaines
espèces ne peuvent être expliqués qu'en partie par le charbonnage ou par le déclin de
la pêche aux perles. Notons que l'industrie de boutons en nacre n'a touché la région
que très légèrement et que les cause de certaines variations observées restent encore
obscures.
RESUMEN
LA FAUNA DE NAIADES DE LA CUENCA DEL ALTO CUMBERLAND ANTES DEL
EMBALSE DE LA RESERVA DEL LAGO CUMBERLAND
Durante 1947-1949, poco antes del embalse del lago Cumberland por cierre del
dique de Wolf Creek, se hizo un estudio de la fauna de almejas de la cuenca del alto
río Cumberland (Kentucky, U. S. A.). Los resultados son comparados con un recono-
cimiento anterior de la misma área en 1911. En el reconocimiento que aquí se informa,
se encontraron 59 distintas almejas de agua dulce, de las cuales 16 pertenecen al
grupo "Cumberlandiano", peculiar de los ríos Cumberland y Tennessee. Sólo 4 especies
(incluyendo una única forma Cumber lan di ana)han remontado los saltos de Cumberland
y ninguna de ellas estaba restricta a aquella área. El primer reconocimiento había
revelado 60 especies, de las cuales sólo 56 coincidían: 3 especies recobradas en el
último reconocimiento no habían sido encontradas previamente y 4 coletadas en el
primero no se encontraron más tarde. En los 36 años desde el primer reconocimiento
varias especies han sido diezmadas por ácidos de las minas de carbón en la parte
más alta de la corriente principal y en sus tributarios mayores. El género Dysnomia
ha sido prácticamente eliminado de su antigua región de mayor variedade, al paso que
cierto número de otras almejas han ganado en abundancia.
-. ,
Sólo algimos de los notables cambios en preponderancia, abundancia relativa y
distribución pueden ser explicados por la declinación de la industria del nácar y la
presencia de ácidos de las minas. El alto Cumberland no fué significativamente ex-
plotado por la industria del botón de nácar. En ciertos casos las causas de la
variación son todavía obscuras.
,(, -
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1947 - 1949 -
1911
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59
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36
Dysnomia
. -
. .
study of mollusks. It aims to provide a common as the following are favored: numbers should not
physiology and taxonomy. Ihe journal will try to %; abbreviations of measures (after a number): mm,
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BURCH, Museum of Zoology, University of Michigan,
-- -
Ann Arbor, Michigan,
DUNCAN, Department
U. A., or to Dr. C, J.
S.
Zoology, University of
of
Liverpool, Liverpool, England, U. K., or to Prof. E,
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. Naturelle, 55 rue de Buífon, Paris V^ France.
,
s
461
1, m 3 , 1964 ,
. , , Tresus
..
nuttalli (Bivalvia:
.
Lymnaça stagnalis
Mactridae)
-
321
331
. ..
,
Taphius glabratus
- 339
X.
Gray
(Gastropoda: Pulmonata)
-
Chilina fluctuosa
355
..
.. ,
I.
.. ,
Planorbidae (Gastropoda:
.
Basommatophora)
Bulinus
(Basommatophora: Branchiopulmonata)
s.
.
387
403
.
Ventridens
(Stylommatophora: Zonitidae) 417
.. ..
427
462 MALACOLOGIA
, ,
. , , TRESUS NUTTALLI
, -
(BIVALVIA: MACTRIDAE)
.: .
, ,.
,
,-
Tresus nuttalli -
-
-
.
60
,, .
,, , . - 60
/ (-
--
,
1958)
. , . ,
,
- ^1
-
.
. ,-
-
--
TaJCHM
.
.. .
,
,., , LYMNAEA STAGNALIS
. . . .
. , , ,, -
12 - 30
,
40 Lymnaea stagnalis
L. stagnalis ,
15
. . , -
,.
MALACOLOGIA 463
,
(Sandoz),.
.
. ,
, 0.08^ (resp. 0.1^)
27° . (resp, 20°)
3
10
. 222
--
-
-
-
5 5
. 27° .;
, -
222 {0.01% 0.3^
, ., , . ,
). - .).
0.1^ 20°
(,
20 2
TAPHIUS GLABRATUS
.
bina
115 M
) ,). 64,
.
,.
Taphius glabratus (/íustralorbis
65,
Planor-
-
(-
,
, , - -
, 6
10 200 . -
.,,,,,.
6 72
. .-
, - , 42,5^ --
. - -
, -
-
- 18,1^.
, , ,, .,.
pacTBopaj 10
-
.
10
4
. 7
-
.
464
-
,
. ,. ,, -
MALACOLOGIA
Planorbidae (Gastropoda: Basommatophora)

BULINUS S.S.
,
1.
. .
, , . Bulinus
-
trwncatusZ
,
" . . "- ". tropicus"
Bulinus s. s.
".
", . .
trimcatus" Schistosoma haema-
tobium; tropicus"
Bulinus truncatus truncatus, , truncatus rohlfsi . coulboisi,

"truncatus" 36 . sericinus, ,
"
truncatus", 72
( = 18) ,
tropicus" (. tropicus tropicus, . tropicus angolensis,
.
. zanzebaricus) . natalensis
--
tropicus . 19
I 20 21
,
,
S. haematobium..
"truncatus" "tropicus"
, . aJfce
.
.., .. ,.
.
R. onychia;
spirillus,
: 18
G. perstriatulus,
16
-
- Radix ollula 17 -
Fossaria truncatula, Physa acuta,
G. tokyoensis,
Radix japónica
Gyraulus
Segmentina hemisphaerula
-
Camptoceras hirasei. G. tokyoensis,

,- , MALACOLOGIA
= .
. (=19)
465
-
-
,. ,-
( 19+1)
G. tokyoensis ,
,
"Lymnaea"
Fossaria,
ollula
euthyneuran
Radix
(1951).
-
-
study of mollusks. It aims to provide a common as the following are favored: numbers should not
physiology and taxonomy. The journal will try to %; abbreviations of measures (after a number): mm,
All manuscripts will be reviewed by at least two Bibliography. See current number of Malacologia
French, German, Russian or Spanish. They must copies may be obtained at cost price if ordered at
be typewritten and double-spaced and are to be Correspondence. Address manuscripts to an ap-
then mainly in places of possible ambiguity or where Michigan, U. S. A., to Dr. C.J. Duncan, Department
economy of space may be achieved. Contributors in of Zoology, University of Liverpool, England, U. K.,
the English language are advised to follow the or to Professor E. Fischer-Piette, Museum National
recommendations of the "Style Manual for Biological d'Histoire Naturelle, 55 rue de Buffon, Paris V^,
ANWEISUNGEN
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DIRECTIVES
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INDEX TO SCIENTIFIC NAMES 467
abrupta, Turbonilla, 170-172 agria, Peristichia, 166, 167

Abyssochrysidae, 10 Aglajidae, 11
Acavidae, 11, 19 ahenum, Drepanotrema, 39
Acella Aillya, 19
haldemani, 256 Aillyídae, 11, 19
Achatina Akeratidae, 12, 17
fúlica, 305 alata, Proptera, 444, 452
Achatinacea, 11, 20 alatus, Paxyodon, 201
Achatinidae, 11 Alaba
Achatinellacea, 11, 19 incerta, 166, 167
Achatinellidae, 11. 19 Alasmidonta
Aciculidae, 1, 14 calceolus, 231
Acleioprocta, 13, 21 marginata, 228-232, 440, 452
Aclididae, 10 minor, 450
Acmaea sp., 166, 167 albescens, Planorbis, 42
Acmaeidae, 9 Taphius, 42
Acmeidae, 14 albicans, Planorbis, 44
Acochlidiacea, 18 Taphius, 44
Acochlidiidae, 11, 18 algicola, Cerithium, 166, 167
Acochlidioidea, 8, 11, 17, 18 alluaudi, Bulinus tropicus, 117
Acroloxacea, 11, 18 Albania
Acroloxidae, 11, 18, 19, 61, 63, 67 auberiana, 167, 169-172
Acroloxinae, 19 Amaltheidae, 10
Acroloxus, 18, 19 Amastridae, 11, 19
lacustris, 55, 316 Amberleyacea, 9
Acrorbis, 35, 41 Amberleyidae, 9
petricola, 41 ambiguus, Velesunio, 192
Acroreidae, 11 Amblema
Acteonacea, 11 costata, 434, 452
Acteonellidae, 11 Ammonitellidae, 12, 20
Acteonellinae, 17 amnicum, Pisidium, 146, 147
Acteonidae, 11 Amphibolacea, 11, 15, 18
Actinonaias Amphlbolidae, 11, 65-67
carinata gibba, 442, 452 Amphibulimidae, 12, 19
carinata, 228 amphiglyptus, Australorbis , 48
gibba, carinata, 442, 452 Amphigyra, 18
pecterosa, 442, 450, 452 Amphiperatidae, 10, 16,
acuta, Physa, 404, 409 Amphithalamus
adamsi, Seila, 166, 172 vallei, 165, 166, 169-172
Tricolia, 166, 167, 171 Ampullariidae, 9, 13
adamsi sargenti, Pisidium, 141, 153 Anachis
adamsi, Pisidium, 141, 147, 152, 153, m.agelioides, 170
155 pulchella, 165, 166, 169-171
Adeorbidae, 9, 14 sp., 166, 167
aequilaterale , Pisidium., 142 Anadara
Aeolidiidae, 13, 21, 22 notabilis, 167, 173
Aeolidiinae, 22 Anadromidae, 12
aeruginosus, Planorbis, 46 Anaplocamidae, 15, 16
affinis. He lis orna, 36 Anaplocamus, 16

Planorbis, 36 Anaspidea, 8, 12, 16, 17
africanas, Bulinus, 87, 89, 117, 118 anatinum, Drepanotrema, 38

Agathirses, 15 anatinus, Planorbis, 38
468 MALACOLOGIA
Ancy lacea, 11, 18 Aplysiacea, 12, 16, 66
Ancylastridae, 64 Aplysiidae, 12
Ancylastrum, 64 Aporrhaidae, 10
Ancylastruminae, 18, 64 applanatus, Helisoma, 36
AncyUdae, 11, 18, 19, 61, 63, 64, 66,67 Planorbis tenuis, 36
Ancylinae, 19, 61, 64 aracasensis, Drepanotrem,a, 38
ancylostomus, Helisoma, 36 Planorbis, 38
Planorbis, 36 arakanensis, Planorbis, 44
Ancy lus, 64 Taphius, 44
fluviatilis, 61 Aranucidae, 12
andecolus, Planorbis,
Taphius, 41
41
angolensis, Bulinus tropicus,
Angaillospira, 15
117, 393,
395
,re a
imbricata, 167, 173, 174
167, 173, 174
arcaeformis, Dysnomia,
Archaeogastropoda,
450
7, 8, 13, 15
angalatus, Drepanotrema, 40 Archaeopulmonata, 67
Planorbis, 40 Architectonicacea, 10, 14
Anisancylus, 64 Architectonicidae, 10, 14, 15
Anisus Arene
lenzi, 39 tricarinata, 170-172
anitensis, Drepanotrema, 39 argentinensis Planorbis,
, 43
Planorbis, 39 Taphius, 43
Arionidae, 12
grandis, 230, 231 Ariophantacea, 12
anodonta, Planorbula obstructa, 45 Ariophantidae, 12
Taphius, 45 armatum, Pisidium punctiferum., 151,
Anodontinae, 203 153
Anodontites, 189, 206 Arm¿¿§ênís, 35
ensiformis, 186 Arminidae, 12, 21
forbesianus, trapesialis, 181-183 Arminoidea, 12, 21
patagónica, 180 arrosum, Pisidium com.pressum, 142
puelchana, 186 Arthessa, 17
tenebricosus, 190 Arthessidae, 12, 17
trapesialis forbesianus, 181-183 Assim.inea
wymanni, 1 80 succinea, 165, 169-172
Anodontitinae, 189, 203, 206 Assimineidae, 9, 13
Anodontoides Asterophilidae, 10
ferrussacianus, 230 Astraea sp., 166, 167
Anomphalacea, 9 atacamensis, Taphius, 45
Anomphalidae, 9 Tropic or bis, 45
Anopsiidae, 12 Ataphridae, 9
Antediplodon, 1 98 Athoracophoracea, 11
Anthracopupa, 13 Athoracophoridae, 11, 19
antiguensis, Planorbis, 92 Atlantacea, 10
Taphius, 92 Atlantidae, 100
antillarum, Caecum, 171 Atyidae, 11
antillarum, Oxynoe, 165, 166, 169, 171 Atys
antillarum, Planorbis, 42 riiseana, 166
Taphius, 42 sp., 166
Antiopellidae, 12, 21 auheriana, Albania, 167, 169-172
Aulacopoda, 12, 66
hypnorum, 270, 271 auricularia. Radix, 256, 407
INDEX, VOL. 1 469
Australorbis , 35 boucardi, Helisoma, 36

amphiglyptus , 48 Planorbis tenuis, 36
christophorensis , glabratus, 42 boucardianus, Planorbis, 46
inßexus, 45 bourguignati, Mutela, 181
megas, bahiensis, 43
auriculatus, Helisoma, 36 eATMSÍMS, 167, 173-175
Planorbis, 36 Brachytremidae, 10
Austroselenitinae, 20 Bradybaenidae, 12
avena, Hyaline, 172 Branchiopiilmonata, 66, 67, 68
babylonia, Cingulina, 170 brazilianus, Planorbis, 47
bahiensis, Planorbis, 43 brevidens, Dysnomia, 448, 452
Taphius, 43 Br onde Ha, 64
barbadensis, Drepanotrema, 39 bryerea, Rissoina, 170-172
Planorbis, 39 Buccinacea, 10
Buccinidae, 10, 16
cancellaria, 167, 175 Buccinulidae, 10, 16
cancellata, 174 Bulimidae, 9, 13
Barleeia, 14
Barleeinae, 14 megas oma, 256
Bartlettia, 190 bulimoides techella, Stagnicola, 239, 254
stefanensis, 190, 206 Bulimulacea, 12
Bartlettiinae, 190, 206 Bulimulidae/ 12
Basommatophora, 11, 18, 19, 62, 66, 67 Bulininae, 63
Bathydorididae, 12 Bulinus, 116, 389, 391
bavayi, Drepanotrema,
, 40 iPhysopsis), 116, 38^, 391
Planorbis, 40 iPyrgophysa), 116, 389, 391
beauii, Cylindrobulla, 171 "sp.", 391, 393
becki, Planorbis, 42 africanus, 87, 89, 117, 118
Taphius, 42 alluaudi, tropicus, 117
belizensis, Helisoma, 36 angolensis, tropicus, 117, 393, 395
Planorbis, 36 canescens, forskalii, 118
,
öeZ/a, Tricolia,
Bellerophontacea,
Bellerophontidae,
166, 172
Planorbis,
Taphius,
Bertheliniacea,
Bertheliniidae,
44
20
20
8
8
44
coulboisi, 117, 393, 395
depressus, hemprichii, 75
depressus, tropicus tropicus form,
dybowskii, truncatus truncabas form,,
forskalii, 118
117
117
biangulatus, Planorbis, 43 globosus, 117, 118

Taphius, 43 hem.prichii depressus, 75
Bithyniidae, 9, 13 innesi, trancatus truncatus form, 117
karongensis, 117
varium, 165, 169-172 mutandaensis tropicus,
,
117
biwaensis, Gyraulus, 411 nasutus, 118
blauneri, Planorbis, 42 natalensis,. 393, 395, 397
Taphius, 42 « ?natalensis'' , 393, 395, 397
boetzkesi, Planorbis, 46 reticulatus, 118
bolivianas j Planorbis, 42 rohlfsi, truncatus, 117, 393, 395
Taphius, 42 scalaris, 118
bonariensis, Drepanotrema, 39 sericinus, 117, 393, 395
Planorbis kermatoides, 39 sericinas, truncatus, 391
Bornellidae, 12 trigonus, truncatus, 117
470 MALACOLOGIA
tropicus, 73 caperata, Stagnicola, 256, 261

tropicus tropicus, 117, 393, 395 capillaris, Planorbis, 47
tropicus tropicus , iorm depressus, 117 Caporbis, 14
truncatus truncatus, 391, 393, 395 capsaeformis, Dysnomia, 448, 452
truncatus truncatus, form innesi, 117 Capulidae, 10
dybowskii, 117 Carbonispira, 13
ugandae, 117, 118 caribaea, Rissoella, 165, 169-172
zanzebaricus tropicus,
, 393, 396 caribaeus, Helisoma, 36
zanzibaricus , tropicus, 117 Planorbis, 36
Bulla Carinariidae, 10
striata, 170, 172 carinata, Actinonaias, 228
Bullacea, 11 carinata gibba, Actinonaias, 442, 452
Bullata
otmliformis, 167, 170, 172 moesta, AAA, 452
BuUidae, 11 casertanum, Pisidium, 142, 147, 152,
Bulloideus, 201, 205 153, 155, 269, 270
Bumupia, 61, 64 casertanum roperi, Pisidium, 153
burroughianus Diplodon,
, 197 Casimiria, 15
Bursidae, 10 Cassididae, 10
Burtinella, 15 Castalia, 201, 204
Caecidae, 10 Castaliella, 201, 205
Caecum Castaliini, 191, 201, 203, 204
antillarum, 171 Castalina, 201, 204
coronellus, 166, 171 psammoica, 200
ßoridanum, 170, 171 castaneonitens, Drepanotrema, 40
nitidum, 165, 167, 169, 172 Planorbis, 40
pulchellum, 165, 169-172 Catantostomatidae, 9
cahawbensis, Rhodacmea, 60, 61 catascopium, Stagnicola, 256
Calceolata, 14 catillus, Drepanotrema, 39
calceolus, Alasmidonta, 231 Planorbis, 39
Caliphyllidae, 12 Cavolinidae, 11
Callomphala, 14 centimetralis, Planorbis, 43
Callonaia, 200, 201, 205 Taphius, 43
Calmidae, 13, 21 Cephalaspidea, 7, 11, 17, 66,
caloderma, Planorbis, 41 Ceriidae, 11, 19

Taphius, 41 cerina, Gouldia, 167, 174
Calyptraeacea, 10, 15, 16 cerinoideus, Ventridens, 424
Calyptraeidae, 10 Cerionidae, 19
Camaenidae, 12, 20 Ceritellidae, 11
Cambodgia, 14 Cerithiacea, 10, 13, 14, 15, 16
Cam/)ioceras Cerithiidae, 10, 14, 15
hirasei, 404, 411 Cerithiopsidae, 10
canaliculata, Odostomia,
cancellaria, Barbatia,
Cancellariidae, 10
166, 167
167, 175 7/
Cerithiopsis sp,,
algie ola, 166, 167

167
cancellata, Barbatia, 174 ebumeum, 170-172

Chime, 167, 173 litteratum, 169-172
canescens, Bulinus forskalii form, 118 variabile, 165, 170
cannarum, Planorbis, 44 chapalensis, Helisoma, 36
Taphius, 44 Planorbis tenuis, 36
canonicus, Planorbis, 43 Charcotia, 21
Taphius, 43 Charcotiidae, 12, 21
INDEX, VOL. 1 471
charruanus, Diplodon, 197, 201 Cochlicopa, 19

chemnitziana, Planorbis tenagophilus , 43 Cochlicopidae, 11, 19
Taphius, 43 Cochlostomatidae, 9
chiapasensis Helisoma,
, 36 Codakia sp., 167, 173
Planorbis ancylostomus, 36 Codonochilidae, 9
chilensis, Diplodon, 191 Iaxis, Ventridens, 422
chilensis, Planorbis, 43 Coelostylinidae, 10
Taphius, 43 collisella, Ventridens, 418
Chilina Colombellinidae, 10
fluctuosa, 356-367 colon, Helisoma, 36
Chilinidae, 11, 18, 63, 65 Planorbis, 36
Chione Columbella
cancellata, 167, 173 mercatoria, 165, 169-172
Drepanotrema,
chittyi, 40 Columbellidae, 10, 15
Chlamydephoridae, 11 columella, Pseudosuccinea, 256
Chondrinidae, 11 complanatus, Elliptio, 230-232
Chondropomidae, 9 compressa, Lasmigona, 231
Choristidae, 9 compressum, Pisidium, 142, 147, 152,
christophorensis , Australorbis glabratus, 153
42 compressum, arrosum, Pisidium., 142
Taphius, 42 compressum. conferium, Pisidium, 142
Chromodorididae, 12, 21 compressum laevigatum, Pisidium, 142
Chrysias, 20 com,pressum pellucidum, Pisidium, 142
cimex, Drepanotrenla, 40 compressum. rostratum, Pisidium, 142
Planorbis, 40 Conacea, 10, 16
cingalata, Schismope, 165, 166, 169-172 concavospira, Planorbis, 42
Cin^lina Taphius, 42
babylonia, 170 concentratus, Planorbis andecolus, 41
Cingulopsidae, 9, 14 Taphius, 41
Cingulopsis, 14 Condylocardia
Cionella, 19 smithii, 167, 173, 174
Cionellacea, 11, 19 confertum., Pisidium. compressum., 142
Cionellidae, 11, 19 confasus, Planorbis, 42
circularis, Planorbis, 46 Taphius, 42
circumlineatus Planorbis,
, 40 Conidae, 10
circumstriatus, Gyraulus, 269-271, 409 conradicus, Medionidus, 444, 452
Cirridae, 9 contortum, Pisidium nitidum, 142, 147,
Cithna, 14 153
Clausiliacea, 11, 19 contrerasi, Helisoma, 36
Clausiliidae, 11, 19 Planorbis, 36
Cleioprocta, 13 conventus, Pisidium, 140, 147, 155
clevei, Planorbis, 43 cooperianus, Plethobasus, 436, 452
Taphius, 43 Coralliophilidae, 10
Clionidae, 12 Corambidae, 12, 21
Cliopsidae, 12 cordatum., Pleurobema, 436, 452
Clisospiracea, 9 cordatum coccineum, Pleurobema, 438,
Clisospiridae, 9 452
coccinea, Pseudostomatella, 166, 172 cordatum plenum, Pleurobema, 436,452
coccineum, Pleurobema cordatum, 438, cordatum. pyramidatum, Pleurobema,
452 436, 452
Cocculinacea, 9 Coreospiridae, 8
Cocculinidae^ 9 Corillacea, 11, 19
472 MALACOLOGIA
Corillidae, 11 cylindrica. Quadrilla, 434, 452
comeum, Sphaerium, 146, 147 Cylindrinae, 16
comeus, Planorbarius , 48, 119 Cylindritella, 17
coronellus, Caecum, '
166, 171 Cylindrobulla, 17
Coryphellidae, 12, 21, 22 beauii, 171
costaricensis, Helisoma, 36 Cylindrobvillacea, 11, 18
Planorbis, 36 Cylindrobullidae, 11, 17
costata, Amblema, 434, 452 Cymatiidae, 10
costata, Lasmigona, 230-232, 440, 452 Cymbuliidae, 11
costabas, Planorbis, 47 cyphyus, Plethobasus, 436, 452
Taphius, 41 Cypraeacea, 10, 16
costulatus costulatus, Gyraulus, 119 Cypraeidae, 10
coton, Helisoma, 36 yprogenia
Planorbis, 36 irrorata, 442, 452
coulboisi, Bulinus, 117, 393, 395 Cyrtolitidae, 8
Craspedopominae, 13 dalli,Fossaria, 269, 270
Craspedostomatacea, 9 Dawsonellidae, 9
r aspe dostomatidae , 9 decipiens, Planorbis, 45
crassidens, Elliptio, 438, 452 Taphius, 45
Crassispira declivis, Drepanotrema, 39
fuscescens, 166 Planorbis, 39, 44
sp., 167 Taphius, 44
Crepidula sp., 166, 167 decussatus, Ventridens, 420
cristatum, Pisidium lilljeborgi, 142, deßectus, Gyraulus, 409
147, 153-155 Deianiridae, 9
cronkhitei. Discus, 270 delodontus delodontus, Diplodon, 194
Crossostomatidae, 9 Dendrodorididae, 12, 21
cruciatum, Pisidium, 140, 147, 151-154 Dendrodoridinae, 21
Cryptobia, 15 Dendronotidae, 12
ryptobranchia, 12, 21 Dendronotoidea, 12, 21
Ctenosculidae, 10 dentatus, Planorbis, 44
Cucumerunionina'e, 205 Taphius, 44
cultratus, Drepanotrema, 39 dentiens, Planorbis, 44
Planorbis, 39 Taphius, 44
Cumberlandia dentifer, Planorbis, 42, 46
monodonta, 432, 450, 452 Taphius, 42
cumingianus, Planorbis, 42 dentiferus, Planorbis, 44
Taphius, 42 Taphius, 44
cumingii, Planorbis, 47 depressissimus, Drepanotrema, 40
Cuthonidae, 13, 21 Planorbis, 40
Cyclomya, 201, 205 depressus, Bulinus hemprichii, 75
Cyclonaias depressus, Bulinus tropicus tropicus form,
granifera, tuberculata, 436, 452 117
tuberculata, 436, 452 desidiosa, Stagnicola palustris, 256
tuberculata granifera, 436,452 Desmopteridae, 11
Cyclophoracea, 9, 13, 15 Diaphanacea, 11
Cyclophoridae, 9 Diaphanidae, 11, 17
Cyclostrema, 13 Diastomidae, 10
Cyclostrematidae, 13, 14 Didianema, 14
Cyclostrem.iscus Dihelice, 15
omatus, 165, 169-172 dilatatus, Elliptio, 230, 232, 438, 450,
pulchellus, 170, 171 452
INDEX, VOL. 1 473
,
DIPLODON, 191, 201, 202, 205 kermatoides, 39
burroughiarms , 197 labrosum, cultratum, 40
charruanas, 197, 201 lanerianus, 39
chilensis, 191 lenzi, 39
delodontus delodontus, 194 39
parallelipipedon, 199 lue idus, 39
parodizi, 199 macnabianus, 40
rhuacoicus, 201 melleus, 39
rotundus, 200 meniscus, 39
soUdulus, 191, 200 nigellus, 38
variabilis, 199 noronhensis, 40
Diplodonta sp., 167,173 panuco, cultratum, 40
Dlplodontini, 199, 203, 205 paropseides, 39
Diplodontites, 205 peninsularis , 40
Dironidae, 12 pistiae, cimex, 40
DzscMS poeyanus, 40
cronkhitei, 270 pulchellus, 40
donaciformis Truncilla,
, 444, 452 puras, 39
donbilli, Segmentina, 44 redfieldi, 39
Taphius, 44 schubarti, 39
Dorcasiidae, 11, 19 sim,m.onsi, 41
Dorididae, 12, 21 SMC ¿news, 39
Doridoidea, 12, 21 sumichrasti, 39
Doridoididae, 12, 21 surinamensis, 39
Doridoxidae, 12 tancredii, 40
Dotonidae, 12 taeniatus, 39
Drepanotrema, 35, 38 tenuissimus, 40
ahenum, 39 ungulatus, 40
anatinum, 38 ysabelensis, 38
angulatus, 40 Drepanotrematinae, 38
anitensis, 39 dromas, Dromus, 442, 452
aracasensis, 38 Dromws
barbadensis, 39 dromas, 442, 452
bavayi, 40 dubium, Pisidium, 146, 147, 152, 153
bonariensis, 39 duenasianus, Drepanotrema, 40
castaneonitens, 40 Planorbis, 40
catillus, 39 dunkeri, Taphius, 45
chittyi, 40 Tropicorbis orbiculus, 45
cemex, 40 dunkerianus, Planorbis, 44
cultratus, 39 Taphius, 44
declivis, 39 Duvaucelia, 21
depressissimus, 40 Duvauceliidae, 21
duenasianus, 40 dybowskü, Bulinus^ truncatus truncatus
esperanzensis, 38 form, 117
fausti, 39 Dysnomi'a
haldemani, 38 arc aeformis , 450
harryi, 40 brevidens, 448, 452
heloicus, 40 capsaeformis, 448, 452
hoffmani, 39 florentina, 448, 452
involutus, 38 florentina,walkeri, 448, 452
isabel, 38 haysiana, 450, 452
jamaicensis, 39 /ew¿s¿, 450, 452
474 MALACOLOGIA
, sulcata,
triquetra,
Ecuadorea,
450, 452
450, 452
walkeri, florentina,
Cerithium,
201, 205
448, 452
170, 172
echinaticostum, Epitonium, 172
edentatus, Planorbis dentiferus, 44

guttarosea,
Euconulidae,
Taphius,
166
12
eucosmius, Planorbis,
42
eudiscus, Helisom,a duryi,
Eudoridacea,
Euglandininae,
21
20
34, 42
37
Taphius, 44 Eulima sp., 166, 167

edentula, Planorbula dentiens, 44 Eulimacea, 10, 16
Taphius Eulimidae, 10, 16
edentulus, Planorbis, 44 Euomphalacea, 8
Taphius, 44 Euomphalidae, 8, 13
Ekadantinae, 13 Euomphalopteridae, 9
Elachorbis, 14 Eupisidium, 141
Elasmonematidae, 9 Eustomidae, 10
electrina, Nesovitrea, 270 Euthyneura, 7, 8, 10, 14, 15, 16, 17, 62,
Elliptio 68
complanatus, 230-232 exaggeratus, Helisoma, 37
crassidens, 438, 452 Planorbis tenuis, 37
dilatatus, 230, 232, 438, 450, 452 excávala, Lampsilis, 215
Ellobiacea, 11 exilis, Stagnicola, 256
Ellobiidae, 11, 13, 66, 67 exustus, Brachidontes, 167, 173-175
elodes, Stagnicola palustris, 256 exustus, Indoplanorbis 118
,
Stagnicola, 269-271 eyerdami, Helisom.a, 38

Elysiacea, 12 fabale, Sphaerium, 146, 147, 153, 154
Elysiidae, 12 fabula, Pegias, 440, 450
emarginata serrata, Stagnicola, 256 Facelinidae, 13, 22
Emarginula sp., 166 fallax, Pisidium, 142, 147, 152-154
Endodontacea, 12, 20 fasciata, Tegula, 166, 171, 172
Endodontidae, 12 fasciola, Lampsilis, 446, 450, 452
Enidae, 11 Fasciolariidae, 10
ensiformis, Anodontites, 186 fasciolaris, Ptychobranchus, 440, 452
Enteroxenidae, 11, 16 fausti, Drepanotrema, 39
Entoconchidae, 11, 16 Favorinidae, 13, 22
Entomotaeniata, 8, 10, 16, 17 Ferrissia, 64
Eolidoidea, 12, 21, 22 Ferrissia
Eotomariidae, 8 paralíela, 61
Epitoniacea, 10, 15 tarda, 61
Epitoniidae, 10, 15 sp., 119
Epitonium, 15 Ferrissiidae, 19
Epitonium Ferrissiinae, 61, 64
echinaticostum, 172 ferrugineum, Pisidium, 142, 147, 152,
sp., 166, 167 153, 155
equatorius, Helisoma, 37 ferrugineum medianum, Pisidium, 142,
Planorbis, 37 153
Eratoidae, 10, 16 ferrugineus, Planorbis, 42
esperanzensis, Drepanotrema, 38 Taphius, 42
Planorbis, 38 Ferrussaciidae, 11
Etheriidae, 190 ferrussacianus Anodontoides,
, 230
Euarminoidea, 21 Ferussininae, 13
Eubranchidae, 13, 21, 22 Ficidae, 10
Eue he lus fie Ida, Planorbis, 46
INDEX, VOL. 1 475
Filholiidae, 11, 19 gigantea, Megalonaias, 434, 452

filocinctus, Planorbis, 46 Girtyspira, 14
Fimbriidae, 12 Glabarinae, 189
Fionidae, 13, 21 glabrata, Planorbina, 119
Fissurella sp., 166, 167 glabratus, Planorbis, 34, 42
Fissurellacea, 9 Taphius, 42, 283, 339-351
Fissurellidae, 9 Glaucidae, 13
Flabellinidae, 13, 21, 22 Glaucinae, 22
flava, Fusconaia, 434, 452 Glauconia, 14, 15
florentina, Dysnomia, 448, 452 Glauconiidae, 10, 14, 15
florentina walkeri, Dysnomia, 448, 452 globosus, Bulinus, 117, 118
floridanum. Caecum, 170, 171 Gnathodoridacea, 21
fluctuosa, Chilina, 356-367 Gnathodoridoidea, 12, 21
fluviatilis,Ancylus, 61 Goniaeolididae, 21
forbesianus, Anodontites trapesialis, Gonieolididae, 12
181-183 Gonieolis, 21
forskalii, Bulinus, 118 Goniodorididae, 12, 21
F os s aria Gonostomopsis , 12
dalli, 269, 270 Gosseletinidae, 8
modicella, 256, 407 GouZdm
m.odicella rustica, 256 cerina, 167, 174
obrussa, 269, 270 gracilentus, Planorbis, 46
ollula per via, 256 grandis, Anodonta, 230, 231
parva, 256, 407 Granger ellidae, 9, 13
per via, ollula, 256 granifera, Cyclonaias tuberculata, 436,

rustica, modicella, 256 452
rustica, 405, 407 guadaloupensis, Planorbis, 42
truncatula, 404, 407 Taphius, 42
Fossaridae, 10, 14 guatem.alensis, Helisoma, 37
Fossula, 189, 206 Planorbis, 37
Fossulorbis, 35 gularis, Ventridens, 424
foveale, Helisoma, 36, 37 gundlachi, Planorbis, 46
fovealis, Planorbis, 37 Gundlachia, 64
fragilis, Helisoma, 37 guttarosea, Euchelus, 166
Leptodea, 444, 452 Gymnentome, 14, 15
Planorbis, 37 Gymnodorididae, 12
fúlica, Achatina, 305 Gymnosomata, 8, 12, 20
fuscescens, Crassispira, 166 Gyrmt/MS
FMSconaèa biwaensis, 411
/Zaua, 434, 452 circumstriatus, 269-271, 409
subrotunda, 434, 452 costulatus costulaius, 119
undata, 434, 452 deflectus, 409
fuscus, Laevapex, 60, 61 hiem,antium, 409, 411
fuscus, Planorbis, 46 perstriatulus, 404, 409, 411
Fusinidae, 10 spirillus, 404, 409, 411
Galeodes, 16 tokyoensis, 404, 409, 411
Galeodidae, 10, 16 gyrina, Physa, 269, 270
Gastropteridae, 11 Haasica, 189, 206
gczudi, Planorbina pfeifferi, 118 Hainesiinae, 13
geoscopus, Segm.entina obstructa, 44 haldemani, Ace lia, 256
Taphius, 44 haldemani, Drepanotrema, 38
gibba, Actinonaias carinata, 442, 452 Planorbis, 38, 45
476 MALACOLOGIA
Taphius, 45 paacispiratas, 37
Halgerdidae, 12, 21 pertenais, 37
Haliotidae, 9 peravianas, 37
Haminoea sp.,166, 167 salvini, 37
Hancockiidae, 12 sinaosus, 37
hanleyi, Pusia, 166 solidus, 37
Haplotrematidae, 11, 20 strebe liana, 37
Harpidae, 10 tenuis, 37
harryi, Drepanotrema, 40 trivolvis, 291
havanensis, Planorbis, 45 tumens, 38
Taphius, 45 tumidus, 38
haysiana, Dysnomia, 450, 452 uhdei, 38
Hebetancylus, 64 wyldi, 38
Hedylopsidae, 11 Helisomatinae, 36
Helcionellacea, 8 Helminthoglyptidae, 12
Helcionellidae, 8 heloicus, Drepanotrema, 40
Helicacea, 12 Planorbis, 40
Helicarionidae, 12 helophilus, Planorbis, 45
Helicidae, 12 Taphius, 45
Helicinidae, 9, 13 hemisphaerula, Segmentina, 404, 411
Helic odiseas hemprichii depressus, Bulinus, 75
sonorensis, lineatus, 47 henslowanum, Pisidium, 142, 147
Helicotomidae, 8 Heroidae, 12, 21
Helisoma, 35, 36 Heterodorididae, 12, 21
affinis, 36 heteropleuras, Planorbis, 41
ancylostomus, 36 Taphius, 41
applanatus, 36 Heterurethra, 11, 66
auriculatus, 36 Hexabranchidae, 12, 21
belizensis, 36 hiemantium, Gyraulus, 409, 411
boucardi, 36 hinds ianus, Planorbis, 46
caribaeus, 36 Hinkleyia, 239
chapalensis, 36 Hipponicacea, 10
chiapasensis, 36 Hipponicidae, 10
colon, 36 Hipponix sp,, 167
contrerasi, 36 hirasei, Camptoceras, 404, 411
costaricensis, 36 hoffmani, Drepanotrema, 39
coton, 36 Holopeidae, 9
equatorius, 37 Holopoda, 12, 66
eudiscus, duryi, 37 Holopodopes, 11, 19, 20
exaggeratus, 37 honduras ensis, Planorbis, 46
eyerdami, 38 humilis, Helisoma, 37
foveale,
fragilis,
36, 37
37
guatemalensis, 37
// Planorbis,
avena, 172
37
humilis, 37 sp., 166, 167

intermedius, 37 Hydatinidae, 11
juvenilis, 37 Hydrobiidae, 9, 13
lentas, 37 Hydrocenidae, 9
mexicanas,
minor,
mysaurus,
37
araguanus
37
37
, 37
,
Hydrococcidae,
hypnorum, Aplexa,
Hyridella,
191, 204
191
9
270, 271
66
INDEX, VOL. 1 477
menziesi, 200 Julia, 20

Hyridellinae, 203, 205 Juliacea, 12, 20
Hyriidae, 198, 203, 204 Juliidae, 12, 20
Hyriinae, 199, 203, 204 juvenilis, Helisoma, 37
idahoense, Pisidium, 146, 147, 152, 154, Planorbis tenuis, 37
155 karongensis, Bulinus, 117
Idioraphe, 14 kermatoides, Drepanotrema, 39
imbricata, Area, 167, 173, 174 Planorbis, 40
immunis, Planorbis, 42 Kittlidiscidae, 9
Taphius, 42 kuhnerianus, Planorbis, 46
incerta^ Alaba, 166, 167 kuhniana, Planorbis, 46
incertulus, Planorbis, 44 labrosum, Drepanotrema cultratum, 40
Taphius, 44 láctea, Marginella, 166, 167
incertus, Planorbis, 44 Lacunidae, 9
Taphius, 44 lacustre, Sphaerium, 146, 147, 152, 153
integra, Physa, 119 lacustre jayense, 146, 147, 153, 154
intermedia, Quadrula, 450 lacustre ryckholti, 146, 147,153, 154
Indoplanorbis lacustris, Acroloxus, 55, 316
exustus, 118 lacustris, Lym.naea stagnalis, 256
inßexus, Australorbis, 45 Laevapecinae, 61, 64
Taphius, 45 Laevapex, 64
innesi, Bulinus truncatus truncatus fornij ßiscus, 60, 61
117 laevigata, Nitidella, 166
insigne, Pisidium, 140, 147, 152-155 Odostomia, 166, 172
insularum., Taphius, 44 laevigatum, Pisidium compressum, 142
Tropicorbis havanensis, 44 laevissim.a, Leptodea, 444, 452
intermedius, Helisoma, 37 Laginiopsidae, 12
Planorbis, 37 Lamellariacea, 10, 16
involutus, Drepanotrema, 38 Lamellariidaè, 10
Planorbis, 38 Lamproscapha, 206
Iravadiidae, 9 Lampsilinae, 203
irrorata, Cyprogenia, 442,. 452 Lampsilis
isabel, Drepanotrema, 38 excávala, 215
Planorbis, 38 fasciola, 446, 450, 452
/sc/znoc/z¿ícwi orbiculata, 448, 452
papillosus, 168 ouaia, 215, 448, 452
sp., 173-175 ouaia ventricosa, 448, 450, 452
isthmicus, Planorbis, 46 siliquoidea, 230, 231
Taphius, 46 ventricosa, 215, 230
Itieria, 1 ventricosa, ovata, 448, 450, 452
Itieriidae, 11 lancéala, Stagnicola, 256
Itruvia, 1 Lancidae, 11, 18, 19, 63
Jacobeanus, Planorbis, 46 Lancinae, 19, 63
jamaicensis, Drepanotrema, 39 lanerianus, Drepanotrema, 39
Planorbis, 39, 48 Planorbis, 39
janeirensis, Planorbis, 44
Taphius, 44 compressa, 231
Janthinidae, 10 costata, 230-232, 440, 452
japónica, Radix, 256, 404, 405, 407 lasmodon, Ventridens, 422
jayense, Sphaerium lacustre, 146, 147,
153, 154 /aia, 438, 452
jugulariSj Lymnaea stagnalis, 256 Zaia, Lastena, 438, 452
5
478 MALACOLOGIA
Lateorbis, 35 lineatus, Planaxis, 167

lateralis, Musculus, 167, 173-175 lineolata. Plagióla, 444, 452
Latia, 18, 63, 67, 68 linnaeus, Planorbis, 47
neritoides, 313 Liotia sp,, 167
Latiidae, 11, 18, 19, 63 Liotiinae, 13
latissima, bigamia recta, 446, 452 Litiopa
Laubellidae, 9 melanostoma, 170, 171
lauricochae, Planorbis, 41 litteratam, Cerithium, 169, 172
Taphius, 41 Littorinacea, 9
lavalleeana, Persicula, 169-172 Littorinidae, 9
Lavigeriidae, 15 Lobiger
lawae, Ventridens, 422 souverbii, 167
L axis pira, 1 Lomanotidae, 12
Leila, 189, 206 Lophospiridae, 8
Leilinae, 203, 206, 207 Lortiellinae, 205
Lemintina, 15 Loxonematacea, 10
lentas, He li soma, 37 Loxonematidae, 10
Planorbis, 37 lucidus, Drepanotrema, 39
lenzi, Anisus, 39 Planorbis, 39
Drepanotrema, 39 Luciellidae, 8
Lepetellidae, 9 lugubris, Planorbis, 42
Lepetidae, 9 Taphius, 42
Leptodea, lundii, Planorbis, 47
fragilis, 444, 452 lutescens, Planorbis, 42
laevissim,a, 444, 452 Taphius, 42
leptodon, 444, 452 Lymnaea
leptodon, Leptodea, 444, 452 jagularis, stagnalis, 256
Leptognatha, 21 lacustris, stagnalis, 256
Leucodiscus, 14 rhodani, stagnalis, 256
Leucorhynchia, 14 stagnalis, 331-335
leucosphaera. Trivia, 166 stagnalis jugalaris, 256
levistriatas Planorbis,
, 43 stagnalis lacustris, 256
Taphius, 43 stagnalis rhodani, 256
lewisi, Dysnomia, 450, 452 Lymnaeacea, 11, 18
liebmanni, Planorbis, 45 Lymnaeidae, 11, 19, 67
Taphius, 45 Lymnaeinae, 19
Ligum.ia Macluritacea, 8
latissima, recta, 446, 452 Macluritidae, 8
recta latissim.a, 446, 452 macnabiarms Drepanotrema,
, 40
Lilax, 15 Planorbis, 40
lilljeborgi, Pisidium, 142, 147, 152, 153, Macrom-phalina, 14
155 Madrellidae, 12
lilljeborgi cristatum, Pisidium, 142, magelioides, Anachis, 170
147, 153-155 Magilidae, 10
Lima sp,, 167, 174 mainense, Pisidium walkeri, 146, 153
Limacidae, 12 Maizaniidae, 9
Limapontiidae, 12 Margaritiferidae, 203, 204
limayana, Planorbis, 39 margínala, Alasmidonta, 228-232, 440,
limayanum, Drepanotrema, 39 452
limayanus, Planorbis peregrinas, 43 Mar¿^¿ne//a
limayanus Taphius,
, 43 /acíea, 166, 167
limosa. Radix, 256, 405, 407 sp., 167
INDEX, VOL. 1 479
Marginellidae, 10 Milacidae, 12
Margine Hops is milium, Pisidium, 142, 147, 152-155
serrei, 167, 169-172 minor, Alasmidonta, 450
Mathildidae, 10, 14, 15 minor, Helisoma, 37
Maturipupa, 13 Planorbis caribaeus, 37
maya, Planorbis, 46 minutulus, Planorbis, 47
Taphius, 46 minutas, Promenetas, 47
medianum, Pisidium ferrugineam, 142, mirabilis, Pedipes, 166, 167
153 Mitracea, 10
Medionidus Mitridae, 10, 16
conradicus, 444, 452 Mitrinae, 16
Meekospira, 14 modicella, Fossaria, 256, 407
Meekospiridae, 10, 14 m,odicella rustica, Fossaria, 256
Megalomphalus , 14 Modulidae, 10
Megalonaias Modulus
gigantea, 434, 452 modulus, 166, i 72
megas, Australorbis bahiensis, , 43 sp., 167
Taphius, 43 modulus. Modulus, 166, 172
megasom.a, Bulimnea, 256 m.oesta, Caranculina, 444, 452
Megaspiridae, 11, 18, 19 Monocondylaea, 189, 206
Megatyloma, 14 Monocondylaeinae, 189, 203, 206
Melanellacea, 10, 16 monodon, Ventridens, 420
Melanellidae, 10, 16 monodonta, Cum.berlandia, 432, 450, 452
Melaniidae, 10 montanensis, Stagnicola, 240, 256
Melanopsidae, 10, 15 montanas, Planorbis, 41
melanostom.a, Litiopa, 170, 171 Taphius, 41, 43
melleus, Drepanotrema, 39 Tropicorbis, 43
Planorbis, 39 moreletianas, Planorbis, 47
Melongenidae, 10, 16 m.oricandi, Planorbis, 48
mendozanus, Planorbis, pfeifferi, 46 Murchisoniacea, 9
meniscus, Drepanotrem,a, 39 Murchisoniidae, 9
Planorbis, 39 Muricacea, 10
menziesi, Hyridella, 200 Muricidae, 10, 16
mercatoria, Colum,bella, 165, 169-172
m,eridaensis , Planorbis, 46 lateralis, 167, 173-175
Mesogastropoda, 9, 15, 17, 18 mutandaensis Bulinus tropicus,
, 117
Mesurethra, 11, 19, 66 Mutela, 205
metanevra, Quadrula, 434, 452 bourguignati, 181
Metoptomatidae, 9 Mutelacea, 186, 203, 205
mexicanas, Helisom,a, 37 Mutelidae, 187, 203, 205
Planorbis, 37, 47 Mycetopoda, 189, 205
Microdomatacea, 9 Mycetopodella, 189, 205
Microdomatidae, 9 Mycetopodidae, 186, 187, 189, 203, 205
Microhedylidae, 11 Mycetopodinae, 189, 203, 205
Micromelaniidae, 9 Myrrhinidae, 13, 22
m,icrom.phalus Planorbis,
, 45 mysaarus, Helisoma, 37
Taphius, 45 Planorbis, 37
Micromya Naricidae, 10
nebulosa, 446, 450, 452 Nasonia, 239
picta, 446, 452 Nassariidae, 10, 16
trabalis, 446, 452 Nassidae, 10
vanuxemensis, 446, 452 nasutas, Bulinus, 118
480 MALACOLOGIA
natalensis, Bulinus, 393, 395, 397 Notodiaphanidae, 11

" ?natalensis" , Bulinus, 393, 395, 397 Notodorididae, 12
natalensis, Radix, 119 Nudibranchia, 12, 20, 66
Naticacea, 10 nuttalli, Tresus, 321-327
Naticidae, 10 Obliquaria
nebulosa, Micromya, 446, 450, 452 reflexa, 440, 452
Neogastropoda, 7, 10, 16 Obovaria
Neopisidium, 140 retusa, 442, 452
Neoplanorbidae, 19 subrotunda, 442, 452
Neoplanorbinae, 19, 64 obrussa, Fossaria, 269, 270
Neptuneidae, 10, 16 Obstructio, 35
Nerinea, 16 obstructus, Planorbis, 45
Nerineacea, 11, 16, 17 Taphius, 45
Nerineidae, 11, 17 obtusale, Pisidium, 142, 147, 152-156
Nerinellidae, 11, 17 obtusale rotundatum, Pisidium, 142, 147,
Neritacea, 9, 13 153-156
Neritidae, 9 obtusale ventricosum, Pisidium, 142, 147,
neritoides, Latia, 313 153,154, 156,
Neritopsidae, 9 obtusale vesiculare, Pisidium, 156
Nesovitrea obvolutus, Planorbis, 45
electrina, 270 Taphius, 45
nicaraguanus, Helisoma, 37 occidentale, Sphaerium, 146, 147, 152,
Planorbis, 37 153
nigellus, Drepanotrema, 38 ocellata, Nitidella, 166
Planorbis, 38 Odontostomidae, 12
nigricanus, Planorbis, 42 Odostom.ia
Taphius, 42 canaliculata, 166, 167
nigrilabris, Planorbis, 44 laevigata, 166, 172
Taphius, 44 Oleacina, 20
nitida, Nitidella, 172 Oleacinacea, 12, 20
Nitidella, Oleacinidae, 12, 20
laevigata, 166 Oleidae, 12
nitida, 172 olivaceus, Planorbis, 42
ocellata, 166 Taphius, 42
sp., 167 Olividae, 10
nitidum, Caecum, 165, 167, 169, 172 ollula. Radix, 404, 405, 407
nitidum, Pisidium, 142, 147, 152, 153, ollula pervia, Fossaria, 256
155 Omalaxidae, 10, 14
nitidum ontortum, Pisidium, 142, 147, 153 Omalogyra, 14
nitidum. pauperculum, Pisidium, 142, Omalogyra sp., 170-172
147, 153 Omalogyridae, 9, 14
nitidum, Sphaerium, 146, 147, 155 Omphalotrochidae, 8
niveus, Triptyckus, 166 Onchidiacea, 13
Nododelphiniilidae, 9 Onchidiidae, 13, 22
Nonsuctoria, 12, 21 Onchidorididae, 12, 21
nordestensis, Tropicorbis, 41 onychia. Radix {Omia), 404, 405, 407
noronhensis, Drepanotrema^ 40 Onychochilidae, 8
Planorbis, 40 Oocorythidae, 16
notabilis, Anadara, 167, 173 Opisthobranchia, 8, 66
Notaeolidiidae, 12, 21 orbiculata, Lampsilis, 448, 452
Notaspidea, 8, 12, 16, 20, 21 orbiculus, Planorbis, 45
Notobranchaeidae, 12 Taphius, 45
INDEX, VOL. 1 481
orbignyana, Planorbis tenagophilus, 43 partumeium, Sphaerium, 146, 147, 152,

Taphius, 43 153
Orbitestellidae, 9 parva, Fossaria, 256, 407
Orculidae, 11 Parviturbo sp., 167
Oreohelicidae, 12, 20 Parviturboides sp., 170, 171
Oriostomatacea, 9 parvus, Planorbis, 48
Oriostomatidae, 9 patagónica, Anodontites, 180
omatas, Cyclostremiscus, 165, 169-172 patagónicas, Planorbis peregrinas, 43
Orthalicidae, 12 Taphius, 43
Orthurethra, 11, 19, 66 Patellacea, 9
Otinidae, 11 Patellidae, 9
Otoconchidae, 12 poMcispiratus, Helisoma, 37
ovata, Lampsilis, 215, 448, 452 Planorbis, 37
ovatOf Radix, 256, 407 paaperculum, Pisidium nitidum,, 142,
ovata ventricosa, Lampsilis, 448, 450, 147, 153,
452 Paxyodon, 191, 201, 204
oviforme, Pleurobema, 438, 452 alatas, 201
Ovulidae, 10, 16 Payettia, 18
ovuliformis, Bullata, 167, 170, 172 Payettiidae, 11, 18
Oxyloma sp., 270 Payettiinae, 18
Oxynoacea, 12 paysanduensis, Planorbis, 43
Oxynoe, Taphius, 43
antillarum, 165, 166, 169, 171 pecterosa, Actinonaias, 442, 450, 452
Oxynoidae, 12
Pachygnatha, 12, 21 mirabilis, 166, 167
Paedophoropodidae, 10 pedrinas, Planorbis, 43
Palaeotrochacea, 9 Taphius, 43
Palaeotrochidae, 9
Palaeozygopleuridae, 10 /&/, 44Ô, 450
Paleoanodonta, 189 pellucidum., Pisidium, compressum, 142
Paleomutela, 189 peninsularis , Drepanotrema, 40
pallidus, Planorbis, 46 Planorbis, 40
Taphius, 45 pentlandi, Planorbis, 41
palustris, Stagnicola, 256 Taphius, 41
desidiosa, Stagnicola, 256 Peraclidacea, 11
elodes, Stagnicola, 256 Peraclididae, 11
palustris group, Stagnicola, 119 pereger. Radix, 256, 407
panamensis, Planorbis, 47 peregrinas, Planorbis, 43
panuco, Drepanotrema, cultratum, 40 Taphius, 43
Paosia, 14 perforabas, Planorbis, 47
paparyensis, Segmentina, 44 Per¿sí¿c^z¿a
Taphius, 44 agria, 166, i 67
papillosus, Ischnochiton, 168 PersicwZa
parallela, Ferrissia, 61 lavalleeana, 169-172
parallelipipedon, Diplodon, 199 palcherrima, 166, 167
Parasita, 7, 11, 16 perStriatalus, Gyraulus, 404, 409, 411
Paratxirbinidae, 9 pertenuis, Helisom,a, 37
ParmacellLdae, 12 Planorbis, tenuis, 37
parodizi, Diplodon, 199 peravianus, Helisoma, 37
paropseides, Drepanotrema, 39 Planorbis, 37, 47
Planorbis, 39 Peraviella, 17
PartxiUdae, 11, 19 pervia, Fossaria ollula, 256
482 MALACOLOGIA
petenensis, Planorbis, 47 cruciatum, 140, 147, 151-154
petricola, Acrorbis, 41 dubium, 146, 147, 152, 153
pfeifferi, Planorbis, 47 fallax, 142, 147, 152-154
pfeifferi gaudi, Planorbina, 118 ferrugineum, 142, 147, 152-153, 155
Phanerobranchia, 12, 21 ferrugineum medianum, 142, 153
Phanerotrematidae, 8 henslowanum., 142, 147
Phasianellidae, 9, 13 idahoense, 146, 147, 152, 154, 155
Phenacolepadidae, 9 insigne, 140, 147, 152-155
Philinacea, 11 laevigatum, compressum, 142
Philinidae, 11 lilljeborgi, 142, 147, 152, 153, 155
Philinoglossacea, 18 lilljeborgi cristatum, 142, 147, 153-155
Philinoglossidae, 11, 18 mainense, walkeri, 146, 153
Philinoglossoidea, 7, 11, 17, 18 medianum., ferrugineum,, 142,153
philippianus , Planorbis, 47 milium, 142, 147, 152-155
Philomycidae, 12, 20 nitidum, 142, 147, 152, 153, 155
PhylUdiidae, 12, 21 nitidum contortum, 142, 147, 153
Phylliroidae, 12 nitidum, pauperculum, 142, 147, 153
Phymatopleuridae, 9 obtusale, 142, 147, 152-156
Physa obtusale rotundatum, 142, 147, 153-156
acuta, 404, 409 obtusale ventricosum, 142, 147, 153,
gyrina, 269, 270 154, 156
integra, 119 obtusale vesiculare, 156
sp., 119 pauperculum, nitidum, 142, 147, 153
Physidae, 11, 19, 67 pellucidum, compressum, 142
Physopsis, 116 punctiferum, 141, 147, 151-155
picth, Micromya, 446, 452 punctiferum armatum, 151, 153
picta, Stomatella, 166, 171 punctiferum, simplex, 152
Pilidae, 9 roperi, casertanum, 153
pilsbryi, Stagnicola, 265 rostratum, compressum, 142
pilsbryi, Ventridens, 418 rotundatum, obtusale, 142, 147, 153-
Pisidiinae, 140 156
Pisidium, 140, 146 sargenti, adamsi, 141, 153
adamsi, 141, 147, 152, 153, 155 simplex, punctiferum, 152
adamsi sargenti, 141, 153 subtruncatum, 142, 147, 155
aequilaterale , 142 variabile, 146, 147, 152, 153, 155
amnicum, 146, 147 ventricosum, obtusale, 142, 147, 153,
armatum, punctiferum, 151, 153 154, 156
arrosum, compressum, 142 vesiculare, obtusale, 156
casertanum, 142, 147, 152, 153, 155, walkeri, 146, 147, 152-155
269, 270 walkeri mainense, 146, 153
casertanum roperi, 153 pistiae, Drepanotrema cimex, 40
compressum, 142, 147, 152, 153
compressum arrosum, 142 lineolata, 444, 452
compressum confertum, 142 Plagiothyridae, 9
com.pressum laevigatum, 142 Planaxidae, 10
compressum pellucidum, 142 P/anax¿s
compressum, rostratum, 142 lineatus, 167
confertum, compressum, 142 sp., 166
contortum, nitidum,, 142, 147, 153 Planitrochidae, 9
conventus, 140, 147, 155 Planorbarius
cristatum, lilljeborgi, 142, 147, 153- comeus, 48, 119
155 Planorbidae, 11, 18, 19, 33, 63, 67
INDEX, VOL. 1 483
Planorbina, 35 concavospira, 42
gaudi, pfeifferi, 118 concentratus, andecolus, 41
glabrata, 119 confusus, 42
pfeifferi gaudi, 118 contrerasi, 36
Planorbis costaricensis, 36
aeruginosus, 46 costatus, 47
36 coton, 36
affinis,
albescens, 42 cultratus, 39
albicans, 44 cumingianus, 42
anatinus, 38 cumingii, 47
ancylostomus, 36 decipiens, 45
andecolus, 41 declivis, 39, 44
dentatus, 44
angulatus, 40
anitensis, 39 dentiens, 44
dentifer, 42, 46
antiguensis, 42
dentiferus, 44
antillarum, 42
depressissimus, 40
applanatus, tenuis, 36
duenasianus, 40
aracasensis, 38
dunkerianus, 44
arakanensis, 44
edentatus, dentiferus, 44
argentinensis, 43
edentulus, 44
auriculatus, 36
equatorius, 37
bahiensis, 43
esperanzensis, 38
barbadensis, 39
eucosmius, 34, 42
bavayi, 40
exaggeratus, tenuis, 37
becki, 42
ferrugineus, 42
belizensis, 36
fieldii, 46
berendtii, 44 filocinctus, 46
biangulatus, 43 fovealis, 37
blauneri, 42 fragilis, 37
boetzkesi, 46 fuscus, 46
bolivianas, 42 glabratus, 34, 42
bonariensis, kermatoides, 39 gracilentus, 46
boucardi, tenuis, 36 guadaloupensis , 42
boucardianus, 46 guatemalensis , 37
brazilianus, 47 gundlachi, 46
caloderma, 41 haldemani, 38, 45
cannarum, 44 havanensis, 45
canonicus, 43 heloicus, 40
capillaris, 47 helophilus, 45
caribaeus, 36 heteropleurus, 41
castaneonitens, 40 hindsianus, 46
catillus, 39 hondurasensis, 46
centimetralis , 43 humilis, 37
chapalensis, tenuis, 36 immunis, 42
chemhitziana, tenagophilus , 43 incertulus, 44
chiapasensis , ancylostomus, 36 incertus, 44
chilensis, 43 intermedius, 37
cimex, 40 involutus, 38
circularis, 46 lsabel, 38
cifcumlineatus, 40 isthmicus, 46
clevei, 43 Jacobeanus, 46
colon, 36 jamaicensis, 39, 48
484 MALACOLOGIA
janeirensis, 44 perforatas, 47
juvenilis, tenuis, 37 pertenuis, tenuis, 37
kermatoides, 40 peruvianas, 37, 47
kuhnerianus, 46 petenensis, 47
kuhniana, 46 pfeifferi, 47
lanerianus, 39 philippianas, 47.
lauricochae, 41 planalatas, AA
lentas, 37 planus, 47
levistriatus, 43 poeyanus, 40
liebmanni, 45 pronas, 41
limayana, 39 pacaraensis, AA
limayanus, peregrinas, 43 pulchellus, 40
linnaeus, 47 paras, 39
lucidus, 39 raimondi, 47
lugubris, 42 redfieldi, 39
lundii, 47 refalgens, 42
lutescens, 42 retusus, 47
macnabianus, 40 reventlowi, 48
maya, 46 riisei, 45
melleus, 39 roía, 48
mendozanus, pfeifferi, 46 salleanas, 40, 41
meniscus, 39 salvini, 37
meridaensis, 46 santacrazensis, 41
mexicanas, 37, 47 schrammi, 45
micromphalus, 45 simplex, 48
minor, caribaeus, 37 sinuosus, 37
minatulus, 47 solidulus, 41
montanas, 41 solidus, 37
moreletianus, 47 stagnicola, 45
moricandi, 48 stramineus, AA
mysöntnts, 37 strebeliana, 37
nicaraguanus, 37 Striatalus, 42
nigellus, 38 strictus, 45
nigricans, 42 subpronus, 41
nigrilabris, AA swcczn^MS, 39
noronhensis, 40 sumichrasti, 39
obstractus, 45 surinamensis, 39
obvolutas, 45 tancredii, 40
olivaceus , 42 taeniatus, 39
orbiculus, 45 tenagophilus, 43
orbignyana, tenagophilus, 43 tenuis, 37
pallidus, 46 tenuissimus, 40
panamensis, 47 tepicensis, 45
paropseides, 39 terverianus, 45
parvus, 48 titicacensis, 41
patagonicus, peregrinas, 43 trigyras, 41
pauc is piratas, 37 /5, 38
paysanduensis, 43 tamidus, 38
pedrinas, 43 uhdci, tenuis, 38
peninsularis, 40 umbilicatas, 47
pentlandi, 41 ungalatas, 40
peregrinas, 43 uruguayensis, 45
INDEX, VOL. 1 485
vaughani, eucosmius, 42 Poteriidae, 9

viridis^ 43 prisons, Pleiodon, 190
weinlandi, 45 Prisodon, 201, 204
wyldi, 38 Prisodontini, 201, 203, 204
xerampeliims, 43 Procerithiidae, 10
ysabelensiSf 38 Procymbuliidae, 11
Planorbula Prograngerella, 13
dentiens edentulay 44 Promenetus
planulatus, Planorbis, 44 minutus, 47
TaphiuSf 44 promis, Planorbis, 41
planus, Planorbis, 47 Taphius, 41
Platyacridae, 9 Proptera
Platyceratacea, 9 alata, 444, 452
Platyceratidae, 9 Prosobranchia, 8
Platytaphius, 35, 41 Protancylinae, 19, 63
PZeiodon Protancylus, 19, 63
priscuSf 190 Provermicularia, 15

plenum, Pleurobema, cordatum, 436, psammoica, Castalina, 200
452 Pseudobroc hidium, 15
PZei/zobasMS Pseudoglauconia, 14
cooperianus, 436, 452 Pseudomelaniacea, 10, 14, 15
cyphyus, 436, 452 Pseudomelaniidae, 10, 14
Plethospiridae, 9 Pseudomesalia, 14, 15
PZeurobema, Pseudophoracea, 9
coccineum, cordatum, 438, 452 Pseudophoridae, 9
cordatum, 436, 452 Pseudorhytidopilus, *18
cordatum coccineum, 438, 452 Pseudorotella, 14
cordatum plenum, 436, 452 Pseudosacculidae, 10
cordatum pyramidatum, 436, 452 Pseudostomatella
oviforme, 438, 452 coccinea, 166, 172
plenum, cordatum, 436, 452 Pseudosuccinea
pyramidatum, cordatum, 436, 452 columella, 256
Pleurobranchaeea, 12, 20 sp., 119
Pleurobranchidae, 12 Pseudotritonia, 21
10, 15, 16 Pseudovermidae, 13, 21
Pleuroceridae,
Pleurodiscidae, 11 Pseudozygopleuridae, 10
Pleuroprocta, 12, 21 Pterotracheidae, 10
Pleurotomariacea, 8 Ptychobranchus
Pleurotomariidae, 9 fasciolaris, 440, 452
pucaraensis, Planorbis, 44
Pneumodermatidae^ 12
40 Taphius, 44
poeyanus, Drepanotrema,
puelchana, Anodontites, 186
Planorbis, 40
pulchella, Anachis, 165, 166, 169-171
Polyceridae, 12, 21
pulchellum. Caecum, 165, 169-172
Polygyracea, 12, 20
pulchellus, Cyclostremiscus, 170, 171
Polygyratia, 20
pulchellus, Drepanotrema, 40
Polygyridae, 12, 20
Polytremariidae, 9 Planorbis, 40
pulcherrima, Persicula, 166, 167
Pomatiasidae, 9
Pulmonata, 8, 67
Porcelliidae, 9
punctiferum, Pisidium, 141, 147, 151-
Porostomata, 12, 21
155
Portlockiellidae, 9
punctiferum armatum, Pisidium, 151, 153
Potamididae, 10
6
486 MALACOLOGIA
punctifemm simplex, Pisidium, 152 reventlowi, Planorbis, 48 ^
Pupillacea, 11, 19 Rhipidodonta, 191, 201, 202, 205

Pupillidae, 11 Rhodacm,ea, 64
Pupinidae, 9 cahawbensis, 60, 61
Purpurinidae, 9 Rhodacmeidae, 19
paras, Drepanotrema, 39 rhodani, Lymnaea stagnalis, 256
Planorbis, 39 Rhodope, 20
Pusia, 1 Rhodopidae, 12
hanleyi, 166 Rhodopoidea, 12
sp., 167 rhomboideum, Sphaerium, 146, 147, 152,
pastulosa, Quadrala, 434, 452 153
pyramidatam, Plenrobema cordatam, rhuacoicus, Diplodon, 201
436, 452 Rhytidacea, 11
Pyramidella, 16 Rhytididae, 11, 20
Pyramide liasp., 166 riiseana, Atys, 166
Pyramidellacea, 11, 17 riisei, Planorbis, 45
Pyramidellidae, 11, 14-17 Taphius, 45
Pyramidulidae, 11 Ringiciilidae, 11
Pyrenidae, 10 Rissoacea, 14
9, 13,
Pyrgocythara sp,, 166 Rissoella, 14
Pyrgophysa, 116 caribaea, 165, 169-172
Pyxipom,a, 15 Rissoellidae, 9, 14
Quadrala Rissoidae, 9, 14
cylindrica, 434, 452 A¿sso¿no,
intermedia, 450 öryerea, 170-172
metanevra, 434, 452 Rochanaia, 190
pastulosa, 434, 452 rohlfsi, Bulinus trancabas, 117, 393,395
Radix roperi, Pisidium, casertanum, 153
aaricularia, 256, 407 Rostellorbis, 14
natalensis, 119 rostratam, Pisidium. compressum, 142
///, 404, 405, 407 roía, Planorbis, 48
ovata, 256, 407 rotandatam, Pisidium obtasale, 142,
pereger, 256, 407 147, 153-156
japónica, 256, 404, 405, 407 rotandus, Diplodon, 200
onychia, (Omia), 404, 405, 407 ragosus, Strophitus, 231, 440, 452
limosa, 256, 405, 407 Runcinidae, 11
raimondi, Planorbis, 47 rustica, Fossaria modicella, 256
rustica, Fossaria, 405, 407
Raphischismatidae, 9
Raphistomatidae, 8
ryckholti, Sphaerium lacustre, 146, 147,
153, 154
Rathouisiidae, 13, 22
Sacoglossa, 8, 12, 17, 20, 21, 66
recta latissima, bigamia, 446, 452
Sagdidae, 12, 20
Rectidentidae, 205 salleanus, Planorbis, 40, 41
redfieldi, Drepanotrema, 39 salvini, Helisoma, 37
Planorbis, 39 Planorbis, 37
reflexa^ Obliquaria, 440, 452 santacrazensis, Planorbis, 41
reflexa, Stagnicola, 256 sargenti, Pisidium adamsi, 141, 153
refalgens, Planorbis, 42 Scalacea, 10, 15
Taphius, 42 scalaris, Bulinus, 118
reticulatas, Bulimis, 118 Scalidae, 10, 15
retasa, Obovaria, 442, 452 Scaphandridae, 11
Retusidae, 11 Sc/2¿smo/>e
retusus, Planorbis, 47 cingulata, 165, 166, 169-172
5
INDEX, VOL. 1 487
Schizogoniidae, 9 Taphius, 41
Schleschiella, 201, 205 solidus, Helisoma, 37
schmiererianus, Taphius, 44 Planorbis, 37
Tropic orbis, 44 sonorensis, Helicodiscus lineatus, 47
schrammi, Planorbis, 45 Sphaeriinae, 146
Taphius, 45 Sphaerium, 146
schubarii, Drepanotrema, 39 comeum, 146, 147
Hippeutis, 39 /abaie, 146, 147, 153, 154
Scissilabra, 14 jayense, lacustre, 146, 147, 153, 154
Scissurellidae, 9 lacustre, 146, 147, 152, 153
Scyllaeidae, 12 lacustre jayense, 146, 147, 153, 154
securis, Sphaerium, 146, 147, 152, 153, lacustre ryckholti, 146, 147, 153, 154
155 nitidum, 146, 147, 155
Segmentella, 15 occidentale, 146, 147, 152, 153
Segmentina, partumeium, 146, 147, 152^ 153
donbilli, 44 rhomboideum, 146, 147, 152, 153
geoscopus, obstructa, 44 ryckholti, lacustre, 146, 147, 153, 154
hemisphaerula, 404, 411 securis, 146, 147, 152, 153, 155
paparyensis, 44 striatinum, 147, 152, 153
Seguenzia, 15 sulcatum, 147, 152, 153, 155
Seguenziidae, 10, 15 transversum, 147, 151-154
57 souverbii, Lobiger, 167
adamsi, 166, 172 Spiratellacea, 11
sericinus, Bulinus, 117, 393, 395 Spiratellidae, 11
sericinus, Bulinus truncatus, 391 Spiraxidae, 11, 20
Serpulorbis, 15 Spiraxinae, 20
serrata, Stagnicola em.arginata, 256 spirillus, Gyraulus, 404, 409, 411
serrei, Marginellopsis, 167, 169-172 Spiroglyphus , 1
shimeki, Taphius, 44 Spirostylidae, 10
Tropic orbis, 44 Spurillidae, 22
Sigmurethra, 11, 19, 66 stagnalis, Lymnaea, 331-335
siliquoidea, Lampsilis, 230, 231 stagnalis jugularis, Lymnaea, 256
simmonsi, Drepanotrema, 41 stagnalis lacustris, Lymnaea, 256
simplex, Pisidium punctiferum, 152 stagnalis rhodani, Lymnaea, 256
Planorbis, 48 Stagnicola, 239
Sinuitidae, 8 Stagnicola ißinkleyia), 239
Sinuopeidae, 8
sinuosus, Helisom.a, 37 bulimoides techella, 239, 254
Planorbis, 37 caperata, 256, 261
Siphonariacea, 11 catascopium, 256
Siphonariidae, 11, 18, 66, 67 desidiosa, palustris, 256
Siphonium, 15 elodes, 269-271
Skeneidae, 9, 13, 14 elodes, palustris, 256
Skeneopsidae, 9, 14 emarginata serrata, 256
smithii, Condylocardia, 167, 173, 174 e xi lis, 256
Solariacea, 14 lance ata, 256
Solariidae, 10, 14 montanensis, 240, 256
Solariorbis, 14 palustris, 256
Solaropsis, 20 palustris desidiosa, 256
Soleolifera, 8, 13, 22, 66 palustris elodes, 256
solidulus, Diplodon, 191, 200 palustris group, 119
solidulus, Planorbis, 41 pilsbryi, 265
488 MALACOLOGIA
reßexa, 256 Planorbis, 39
serrata, emarginataj 256 Suctoria, 12, 21
techella, bulimoides, 239, 254 sulcata, Dysnomia, 450, 452
umbrosa, 256 sulcatum, Sphaerium, 147, 152, 153, 155
stagnicola, Planorbis, 45 sumichrasti, Drepanotrema, 39
Taphius, 45 Planorbis, 39
Starke yna, 14 suppressus, Ventridens, 424
stefanensis, Bartlettia, 190, 206 surinamensis, Drepanotrema, 39
Stenacmidae, 16, 18 Planorbis, 39
Stenoglossa, 10, 15, 16 Symmetrocapulidae, 9
Stenothyridae, 9 Synceratidae, 9
Stephopoma, 15 Syrnolopsidae, 10, 13
Stiliferidae, 10 Systrophiidae, 12, 20
Stiligeridae, 12 taeniatus, Drepanotrema, 39
Stomatella Planorbis, 39
picta, 166, 171 Tamanovalva, 20
Stomatellidae, 9 Tamanovalvacea, 20
stramineus, Planorbis, 44 Tamanovalvidae, 20
Taphius, 44 Tamsiella, 189, 206
strebeliana. He lis orna, 37 tancredii, Drepanotrema, 40
Planorbis, 37 Planorbis, 40
Streptacididae, 11, 17 Taphiinae, 41
Streptaxacea, 11 Taphius, 35, 41
Streptaxidae, 11, 20 albescens, 42
Streptoneura, 7, 8 albicans, 44
Streptostylinae, 20 andecolus, 41
striata. Bulla, 170, 172 anodonta, 45
striatinum, Sphaerium, 147, 152, 153 antiguensis, 42
striatulus, Planorbis, 42 antillarum, 42
Taphius, 42 arakanensis, 44
s trie tus, Planorbis, 45 argentinensis , 43
Taphius, 45 atacamensis, 45
Strombacea, 10, 16 bahiensis, 43
Strombidae, 10 becki, 42
berendtii, 44
rugosus, 231, 440, 452 biangulatus, 43
Strophocheilacea, 11, 19 blauneri, 42
Strophocheilidae, 11, 19 bolivianas, 42
Struthiolariidae, 10 caloderma, 41
Stylommatophora, 7, 11, 19, 66, 67 cannarum, 44
subpronus, Planorbis, 41 eanoTÚcus, 43
Taphius, 41 centimetralis, 43
subrotunda, Fusconaia, 434, 452 chemnitziana, 43
subrotunda, Obovaria, 442, 452 chilensis, 43
subtfuncatum, Pisidium, 142, 147, 155 christophorensis , 42
Subiilinidae, 11 clevei, 43
Subulitacea, 10, 14 concavospira, 42
Subulitidae, 10 concentratus, 41
succinea, Assiminea, 165, 169-172 confusus, 42
Succineacea, 11 costatus, 41
Succineidae, 11 cumingianus, 42
succineus, Drepanotrema, 39 decipiens, 45
INDEX, VOL. 1 489
declivis, 44 pronas f 41
dentatus, 44 pucaraensiSf 44
dentiens, 44 reßilgens, 42
dentifer, 42 riisei, 45
dentiferuSf 44 schmiereriamiSy 44
donbilliy 44 schrammiy 45
dunkerif 45 shimeki, 44
dunkeriamis, 44 solidulus, 41
edentatus, 44 stagnicola, 45
edentulüf 44 stramineus, 44
edentuluSf 44 striatulus, 42
eucosmius, 42 s trie tus f 45
ferrugineus, 42 subpronus, 41
geoscopus, 44 íaíe¿, 45
glabratus, 42, 283, 339-351 tenagophilus , 43
guadaloapensis, 42 tepicensis, 45
haldemani, 45 terverianus, 45
havanensis, 45 thermaluSf 47
helophilus, 45 titicacensis, 41
heteropleurus , 41 trigyruSf 41
immunis, 42 uruguayensis, 45
incertuluSy 44 vaughani, 42
incertuSf 44 viridis, 43
inßexuSf 45 weinlandi, 45
insularunif 44 xerampelinus, 43
isthmicus, 46 tarda, Ferrissia, 61
janeirensis, 44 íaíe¿, Taphius, 45
lauricochae, 41 Tropicorbis, 45
levistriatus , 43 techella, Stagnicola, bulimoides, 239,
Uebmanni, 45 254
limayanus, 43
lugubriSf 42 fasciata, 166, 171, 172
lutescens, 42 Teinostoma, 14
maya, 46 Telarma, 21
megas j 43 Temnotropidae, 9
micromphalus, 45 Tenagodus, 15
montanus, 41, 43 tenagophilus, Planorbis, 43
nigricans^ 42 Taphius, 43
nigrilabris, 44 tenebricosus, Anodontites, 190
obstructus, 45 tenuis, Helisoma, 37
obvolutuSf 45 Planorbis, 37
olivaceus, 42 tenuissimus, Drepanotrema, 40
orbiculus, 45 Planorbis, 40
orbignyana, 43 tepicensis, Planorbis, 45
palliduSf 45 Taphius, 45
paparyensis, 44 Terebracea, 16
patagonicus, 43 Terebridae, 10
paysanduensis, 43 Tergipedidae, 21
pedrinus, 43 terverianus, Planorbis, 45
pentlandi, 41 Taphius, 45
peregrinas, 43 tessellata, Tricolia, 170, 172
planulatus, 44 Testacellacea, 12
490 MALACOLOGIA
Testacellidae, 12 Tritoniidae, 12, 21
Thaisidae, 10 Trivia
thalassicola, Tricolia, 166, 172 leucosphaera, 166
Thecosomata, 8, 11 Helisoma,
trivolvis, 291
theloides, Ventridens, 420 Trochacea, 9
thermalus, Taphius, 47 Trochaclisidae, 9
Thiaridae, 10, 13, 14, 15 Trochidae, 9, 15
Thliptodontidae, 12 Trochomorphidae, 12
Thyrophorellidae, 12 Trochonematacea, 9
Thysanophoridae, 12, 20 Trochonematidae, 9
titicacensis, Planorbis, 41 Trochotomidae, 9
Taphius, 41 Troostella, 17
Titiscaniidae, 9 Tropic or bis, 35
tokyoensis, Gyraulus, 404, 409, 411 dunkeri, orbiculus, 45
Tonnacea, 10 insularum, havanensis, 44
Tonnidae, 10, 16 montanus, 43
Tornatellinidae, 19 nordestensis, 41
Tornidae, 9, 14 schmiererianus 44 ,
T orre hry Sias, 20 shimeki, 44

Toxoglossa, 10, 16 tatet, 45
trabalis, Micromya, 446, 452 tropicus, Bulinus, 73
Trachysmldae, 9 tropicus tropicus, Bulinus, 117, 393,395
Trajanella, 14 tropicus tropicus, Bulinus, form
Trajanellidae, 14 depressus, 117
transversum, Sphaerium, 147, 151-154 truncata, Truncilla, 444, 452
trapesialis forbesianus, Anodontites, Truncatella sp., 166
181-183 Truncatellidae, 9
Tresus truncatula, Fossaria, 404, 407
mittalli, 321-327 truncatus truncatus, Bulinus, 391, 393,
tricarinata. Arene, 170-172 395
Trichotropididae, 10 truncatus truncatus, Bulinus, form
Tricolia innesi, 117
adamsi, 166, 167, 171 dybowskii, 117
bella, 166, 172
tessellata, 170, 172 donaciformis , 444, 452
thalassicola, 166, 172 truncata, 444, 452
Trigonochlamydidae, 12 tuberculata, Cyclonaias, 436, 452
trigonus, Bulirms truncabas, 117 tuberculata granifera, Cyclonaias, 436,
trigyrus, Planorbis, 41 452
Taphius, 41 Tubinidae, 9
Trimusculidae, 11 tumens, Helisoma, 38
Triophidae, 12 Planorbis, 38
Triphora sp., 166 tumidus, Helisom,a, 38
Triphoridae, 10, 15 Planorbis, 38
Triplodon, 204 Turbinidae, 9, 13
Triptychus
niveus, 166 abrupta, 170-172
triquetra, Dysnomia, 450, 452 sp., 170, 171
Triquetrana, 204 Turridae, 10
Tritigonia Turritellidae, 10, 15
verrucosa, 436, 452 ugandae, Bulinus, 117, 118
Tritonia, 21 uhdei, Helisoma, 38
INDEX, VOL. 1 491
Planorbis tenuis, 38 theloides, 420

umbilicatus, Planorbis, 47 virginicus, 425
Umbraciilacea, 12, 20 Vermetidae, 10, 14, 15
Umbraculidae, 12 Vermicularia, 15
umbrosa, Stagnicola, 256 Veronicellacea, 13, 22
Uncancylus, 64 Veronicellidae, 13, 22
undata, Fusconaia, 434, 452 verrucosa, Tritigonia, 436, 452
ungulatus, Drepanotrema, 40 Vertiginidae, 11
Planorbis, 40 vesiculare, Pisidium obtusale, 156
Unionacea, 186, 203, 204 Vexillinae, 16
Unionidae, 203, 204 virginicus, Ventridens, 425
Unioninae, 203 viridis, Planorbis, 43
Unnamed superfamily, 11 Taphius, 43
Urocoptidae, 12 Vitrinella sp., 166, 167
Urocyclidae, 12 Vitrinellidae, 9, 13, 14
uruguayensis, Planorbis, 45 Vitrinidae, 12
Taphius, 45 Viviparacea, 9, 13
vallei, Amphithalamus , 165, 166, 169- Viviparidae, 9, 13
172 Volutacea, 10, 16
Valloniidae, 11 Volutidae, 10
Valvatacea, 9 Volvatella, 17
Valvatidae, 9 walkeri, Dysnomia florentina, 448, 452
Vanikoridae, 10 walkeri, Pisidium, 146, 147, 152-155
vanuxemensis Micromya,
, 446, 452 walkeri mainense, Pisidium, 146, 153
variabile, Cerithium, 165, 170 Weeksiidae, 8, 13
variabile, Pisidium, 146, 147, 152, 153, weinlandi, Planorbis, 45
155 Taphius, 45
variabilis, Diplodon, 199 wyldi, Helisoma, 38
varium, Bittium, 165, 169-172 Planorbis, 38
Vasidae, 10 wymanni, Anodontites, 180
vaughani, Planorbis eucosmius, 42 Xancidae, 10
Taphius, 42 Xenophoridae, 10, 16
Vayssiereidae, 12, 21 xeram,pelinus, Planorbis, 43
Velainellidae, 9 Taphius, 43
Velesunio ysabelensis, Drepanotrema, 38
ambiguus, 192 Planorbis, 38
Velesunioninae, 203, 205 Zalophancylus, 19
ventricosa, Lampsilis, 215, 230 zanzebaricus, Bulinus tropicus, 393, 395
Lampsilis ovata, 448, 450, 452 zanzibaricus , Bulinus tropicus, 117
ventricosum, Pisidium obtusale, 142, 2e&ra, Area, 167, 173, 174
147, 153, 154, 156 Zephyrinidae, 21
Ventridens Zonitacea, 12, 20
cerinoideus, 424 Zonitidae, 12
coelaxis, 422 Zygitidae, 9
collisella, 418 Zygopleuridae, 10
decussatus, 420
gularis, 424
lasmodon, 422
lawae, 422
m.onodon, 420
pilsbryi, 418
suppressus, 424
Vol. 1, No. 3 MALACOLOGIA June 1964
CONTENTS
Page
R. POHLO
Ontogenetic changes of form and mode of life in
Tresus nuttalli (Bivalvia: Mactridae) 321
J. LEVER, J. JAGER and A. WESTERVELD

A new anaesthetization technique for fresh water snails,
tested on Lymnaea stagnalis 331
CM. YAGER and H. W. HARRY

The uptake of radioactive Zinc, Cadmium and Copper
by the freshwater snail, Taphius glabratus 339
H. W. HARRY
The anatomy of Chilina ßuctuosa Gray reexamined, with
prolegomena on the phylogeny of the higher limnic
Basommatophora (Gastropoda: Pulmonata) 355
J. B. BURCH
Cytological studies of Planorbidae (Gastropoda: Basommatophora).
I. The African subgenus Bulinus s.s 387
J. B. BURCH, J. E. WILLIAMS, Y. HISHINUMA and R. NATARAJAN

Chromosomes of some Japanese freshwater snails
(Basommatophora: Branchiopulmonata) 403
L. HUBRICHT
The bidentate species of Ventridens (Stylommatophora:
Zonitidae) 417
J. K. NEEL and W. R. ALLEN

The mussel fauna of the upper Cumberland Basin before
itsimpoundment 427
Date Due

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HARVARD UNIVERSITY

Museum of Comparative Zoology

International Journal of Malacology

Revista Internacional de Malacologia

Journal International de Malacologie

Internationale Malakologische Zeitschrift

At least 50 copies of MALACOLOGIA were mailed to subscribers (including a free

Cytological studies of Planorbidae (Gastropoda: Basommatophora).

J. B. BURCH and M. PATTERSON

J. B. BURCH, J. E. WILLIAMS, Y. HISHINUMA and R. NATARAJAN

The anatomy of Chilina fluctuosa Gray reexamined, with prolegomena

H. W. HARRY and D. V. ALDRICH

J. LEVER, J. C. JAGER and A. WESTERVE LD

J. K. NEEL and W. R. ALLEN

J. J. PARODIZ and A. A. BONETTO

I. STIGLINGH, J. A. VAN EEDEN and P. A. J. RYKE

D. W. TAYLOR and N. F. SOHL

D. W. TAYLOR, H. J. WALTER and J. B. BURCH

H. VAN DER SCHALIE

G. L. WARMKE and L. R. ALMODOVAR

M. YAGER and H. W. HARRY

TAXONOMIC AND NOMENCLATURAL CHANGES

Mutelacea, Parodiz and Bonetto, 1963, 186

Architectonicacea, Taylor and Sohl, 1962, 14

VOL. 1 NO. 1 OCTOBER 1962

ifiüS. W' "^"^' '

International Journal of Malacology

Revista Internacional de Malacologia

Journal International de Malacologie

Internationale Malakologische Zeitschrift

EDITORIAL BOARD CONSEJO EDITORIAL

. H. BARNARD N. A. HOLME W. L. PARAENSE

fut avec l'aide d'une subven-

MALACOLOGIA, AN INTERNATIONAL JOURNAL OF MALACOLOGY

The founders of MALACOLOGIA hope tions obtained, establishment of a self-

MALACOLOGIA, UN JOURNAL INTERNATIONAL DE MALACOLOGIE

Les fondateurs de MALACOLOGIA es- pour un journal pareil. Un questionnaire

MALACOLOGIA, UNA REVISTA INTERNACIONAL DE MALACOLOGIA

artículos y monografías que resultan muy Melbourne R. Carriker Norman F. Sohl

AN OUTLINE OF GASTROPOD CLASSIFICATION^'^

^Publication authorized by the Director, U. S. Geological Survey

Asterisks denote groups that are known

Amphibulimidae (note 43) 8 15. Order Notaspidea (note 50) 18

Infraorder Aulacopoda 610 Umbraculidae 6

14 TAYLOR AND SOHL

The name Acmeidae used by Wenz

OUTLINE OF GASTROPOD CLASSIFICATION 15

tectonicidae in operculum, radula, and dae on the basis of reproductive charac-

thirses Montfort, 1808; Anguillospira for the Triphoridae is probably justified,

OUTLINE OF GASTROPOD CLASSIFICATION 17

tilus, 70(3): 141-142.

Aeolidiinae. Glaucinae of Thiele are clausilioids. Nautilus, 75: 33-36.

Nautilus, 75: 142-145.

of Rathouisiidae and Veronicellidae from genus Epitonium in the western Atlantic,

1956, The family Melongenidae in

the western Atlantic. Johnsonia, 3: 161-188.

of Geology, Stanford University; Harald A. comparée. Paris (privately printed). 12:

OUTLINE OF GASTROPOD CLASSIFICATION 23

(Jeffreys). J. Marine Biol. Assoc. U. K. LE LOUP, Eugène, 1953, Gastéropodes. Inst.

London, 33: 114-126. Brazil (2). J. Linnean Soc. London, Zool.,

18-19. thobranchiate gastropods. Amer. Mus. Nat.

of the Californian Province, based upon an California. Ve liger, 3, Suppl. pt. 1, 84 p.

enskaps— och Vitterhets-Samhailes Handlin- 232, Zool. (22): 99-188.