T urbellarian Biology978-94-011-2775...Hydrobiologia 227, 1991.S. Tyler (ed.), Turbellarian Biology. v Preface Turbellarian platyhelminths (or, as they are known now among cladistic

T urbellarian Biology

Developments in Hydrobiology 69

Series editor

H. J. Dumont

Turbellarian Biology

Proceedings of the Sixth International Symposium on the Biology of the Turbellaria,

held at Hirosaki, Japan, 7-12 August 1990

Edited by

Seth Tyler

With the Assistance of the Editorial Board

Wataru Teshirogi, Sachiko Ishida, Ernest Schockaert, Gunde Rieger, Reinhard Rieger, Ian R. Ball & Michael D. B. Burt

Reprinted from Hydrobiologia, vol. 227 (1991)

Kluwer Academic Publishers Dordrecht / Boston / London

Library of Congress Cataloging-in-Publication Data

ISBN 0-7923-1373-9

Published by Kluwer Academic Publishers, P.O. Box 17,3300 AA Dordrecht, The Netherlands.

Kluwer Academic Publishers incorporates

the publishing programmes of

D. Reidel, Martinus Nijhoff, Dr W. Junk and MTP Press.

Sold and distributed in the U.S.A. and Canada by Kluwer Academic Publishers, 101 Philip Drive, Norwell, MA 02061, U.S.A.

In all other countries, sold and distributed

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P.O. Box 322, 3300 AH Dordrecht, The Netherlands.

All Rights Reserved © 1991 Kluwer Academic Publishers

No part of the material protected by this copyright notice may be reproduced or utilized in any

form or by any means, electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without written permission from the copyright owners.

Hydrobiologia 227, 1991. S. Tyler (ed.), Turbellarian Biology. v

Preface

Turbellarian platyhelminths (or, as they are known now among cladistic systematists, free-living Plathelminthes) comprise a widely distributed assemblage of lower worms found in marine, freshwater, and even occasionally in terrestrial habitats. The phylum Platyhelminthes may be more widely known for its parasitic members since the major parasitic groups of the tapeworms, flukes, and their relatives are more speciose and have greater impact on everyday human life; but the turbelIarians are more diverse and, as inhabitants of virtually any aquatic habitat, are more widespread as well. Many of the lower turbellarians are rather simple in morphology and have served as models for ancestors of the Bilateria, i.e., the bulk of the animal phyla. Others are quite complex organisms, especially in the morphology of their reproductive systems which are highly specialized. The majority is free-living in aquatic habitats but a number of interesting parasitic and commensal species are found scattered among the higher turbellarian taxa.

But turbellarians are more than just taxonomic curiosities. They have served as illustrative models in research on a variety of basic life processes. For example, their high capacity for regeneration has made them the subject of a large literature in developmental biology, the occurrence of mixoploidy and other karyological oddities among turbellarians has been important in understanding evolution of the genome, and the fine structure and biochemistry of the nervous system in turbellarians is revealing important principles of the organization of so-called primitive neural systems.

Turbellarians have been the focus of a series of international symposia, the first held in 1970 when members of the American Society of Zoologists organized a symposium honoring the memory of the preeminent American student of the group, Libbie Hyman. The sixth in this series of symposia was held in Hirosaki, Japan, in August 1990; and papers presented there are compiled in this volume. Japan was a fitting site for this gathering; it has long been an important center of research on turbellarians, particularly on triclad turbellarians, with study of their regeneration, biogeography, genetics, ecology, ultrastructure, etc. by many internationally recognized scientists.

Professor Dr Wataru Teshirogi, Dean of the Faculty of Science at Hirosaki University, graciously organized the symposium, and his accomplishments in this task were truly awe-inspiring. Assisted by a local organizing committee with Dr Sachiko Ishida and members from other Japanese universities (see p. vi), he brought to fruition a highly orderly and outstandingly enjoyable symposium involving approximately 150 correspondents from 19 countries.

The festivities and interesting scientific developments at the symposium were dampened somewhat by announcement that one of the best-recognized specialists on turbellarians, Dr Eveline Du Bois-Reymond Marcus, died in 1990. In recognition of her many accomplishments and contributions to the study of turbellarians, the organizing committee of the symposium dedicates this volume to her memory. Dr Diva Diniz Correa has kindly provided a biography of Dr Marcus and a list of her publications; these appear on the following pages.

SETH TYLER

Hydrobiologia 227, 1991. VI S. Tyler (ed.), Turbellarian Biology.

Organizers of the Sixth International Symposium on the Biology of the Turbellaria Advisers Dr Shuji Tohno, President, Hirosaki University Dr Hisao Sugino, Emeritus Professor, Osaka Kyoiku University Dr Tetsuji Kido, Emeritus Professor, Kanazawa University Dr Kenji Osanai, Director, Marine Biological Station of Asamushi, Tohoku University

International Organizing Committee Chair: Dr Wataru Teshirogi,

Hirosaki University Dr Ian R. Ball

Memorial University of Newfoundland, Canada Dr M. D. B. Burt

University of New Brunswick, Canada

Local Organizing Committee Dr Sachiko Ishida

Hirosaki University Dr Michio Morita

Colorado State University Dr Saburo Ishii

Fukushima Medical College Dr Takashige Sakurai

Fukushima Medical College Dr Masuo Aikawa

Dokkyo University School of Medicine Dr N aoya Makino

Tokyo Medical College

Cooperative Staff Dr Shigeo Chiba

School of Allied Medical Sciences, Hirosaki University

Dr Emiko Furuta Dokkyo University School of Medicine

Dr Masashi Kodama Faculty of Education, Hirosaki University

Mr Yoshihiro Kon-no Sendai Ikuei Gakuen Senior High School

Dr Satoshi Kuchiiwa Hirosaki University School of Medicine

Dr Mitsuaki Kurotaki School of Allied Medical Sciences, Hirosaki University

Dr Ernest R. Schockaert Limburgs University, Belgium

Dr Seth Tyler University of Maine, USA

Dr Yasuko Shirasawa Tokyo Medical College

Dr Isao Hori Kanazawa Medical University

Dr Etsuo Asai Kanazawa Medical University

Dr Iwashiro Oki Osaka Environmental Project Authority

Dr Sachiko Tamura Osaka Prefectural Institute of Public Health

Dr Terufumi Yamasu Ryukyu University

Dr Tohsaku Nikara School of Allied Medical Sciences, Hirosaki University

Dr Takaharu Numakunai Marine Biological Station of Asamushi, Tohoku University

Dr Y oshitaka Obara Faculty of Science, Hirosaki University

Dr Keiko Ohkawa College of Liberal Arts, Hirosaki University

Dr Kazuo Sugiyama Faculty of Science, Hirosaki University

Dr Motoichi Yagihashi Ogaki Women's Junior College

Mr Makoto Yamada Kuroishi Senior High School

Hydrobiologia 227, 1991. S. Tyler (ed.), Turbellarian Biology. vii

Acknowledgments

The Sixth International Symposium on the Biology of the Turbellaria was made possible by the financial support of the following organizations:

Japan World Exposition Commemorative Fund Yoshida Foundation for Science and Technology The Prefectural Government of Aomori

(JEC Fund) Inoue Foundation for Science The Naito Foundation The Kajima Foundation Life Science Foundation of Japan

The Municipal Office of Hirosaki The Japanese Ministry of Education, Science and

Culture The Hirosaki University Foundation for Scienti

fic Research Encouragement

For editing of the Proceedings of the Symposium, financial support was also provided by the University of Maine.

The papers published in this Proceedings were subject to review by outside reviewers as well as by the editorial board. The editorial board gratefully acknowledges the help of the following reviewers:

Jaume Baguna Jay Boyd Best Ulrich Ehlers Werner Fischlschweiger Jacinto Garno Jan Hendelberg Isao Hori Susan J. Hunter Saburo Ishii Leonard Kass Masaharu Kawakatsu

Jorg Klima J erzy Kolasa Harold Koopowitz Michael Lesser Charles W. Major 1. Martelly Paul M. Martens Robert W. Mead Paul Moran Michio Morita Knud Jergen Pedersen

Stephen Prudhoe Maria Reuter Franklin L. Roberts Edward E. Ruppert J. Malcolm Shick Takao Shinozawa J. P. S. Smith, III Beate Sopott-Ehlers Wolfgang Sterrer Mary Beth Thomas Mary S. Tyler

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x

List of Participants and Contributors

H. Adam Department of Zoology Universitiit Salzburg Hellbrunnerstr. 34 A-5020 Salzburg Austria

Ajit Kumar Aditya Department of Zoology Visva-Bharati University Santiniketan - 731235 West Bengal India

Masuo Aikawa Department of Anatomy Dokkyo University School of Medi

cine Mibu, Tochigi 321-02 Japan

Bertil Akesson Department of Zoology University of Goteborg Box 25059 S-400 31 Boteborg Sweden

Etsuo Asai Department of Biology Kanazawa Medical University Uchinada-machi, Ishikawa-Ken 920-02 Japan

Peter Ax II. Zoologisches Institut und Museum

der Georg-August-Universitiit Berliner Str. 28 3400 Gottingen Germany

Renate Ax II. Zoologisches Institut und Museum


Jaume Baguiia Departament de Genetica Facultat de Biologia Universitad de Barcelona Diagonal 645 08028 Barcelona Spain

Celina Bedini Dipartimento di Science del Compor-

tamento Animale e dell'Uomo Universita di Pisa Via A. Volta 6 1-56100 Pisa Italy

Jay Boyd Best Department of Environmental Health Colorado State University Fort Collins, CO 80523 USA

David Bueno Departament de Genetica Facultat de Biologia Universitat de Barcelona Diagonal 645 08028 Barcelona Spain

Michael D. B. Burt Department of Biology Bag Service Number 45111 University of New Brunswick Fredericton, N. B., E3B 6EI Canada

Christine Cannon Queensland Museum P.O. Box 300 South Brisbane, Queensland 4101 Australia

Lester Cannon Queensland Museum P.O. Box 300 South Brisbane, Queensland 4101 Australia

Hydrobiologia 227, 1991. S. Tyler (ed.). Turbellarian Biology.

Shigeo Chiba School of Allied Medical Sciences Hirosaki University 66-1 Hon-cho Hirosaki 036 Japan

Dr Diva Diniz Correa Departamento de Zoologia Instituto de Biociencias Universidade de Sao Paulo C. P. 20520 01498 - Sao Paulo, SP Brazil

Marco Curini-Galletti Dipartimento di Scienze Dell'Ambiente e del Territorio Universita di Pisa Via Volta 6 56126 Pisa Italy

Eunice Judith Cardoso de Almeida Departamento de Biologia Instituto de Biociencias Universidade de Sao Paulo C. P. Jl461 05499 - Sao Paulo, SP Brazil

Alain De Vocht Dept. S. B. M. Limburgs Universitair Centrum B-3610 Diepenbeek Belgium

Sabine Dittmann Australian Institute of Marine

Science P. M. B. No.3 Townsville, M. c., Q 4810 Australia

Ulrich Ehlers II. Zoologisches Institut und Museum


Krister Eriksson Institutionen f6r Biologi Department of Biology Abo Akademi Porthansgatan 3 SF-20500 Abo 50 Finland

Alessandra Falleni Dipartimento di Biomedicina Speri-

mentale, Infettiva e Pubblica Sezione di Biologia e Genetica Universita di Pisa Via A. Volta 4 56100 Pisa Italy

Enrico Antonio Ferrero Department of Biology University of Trieste Via Alfonso Valerio 32 1-34127 Trieste Italy

Eudoxia Maria Froehlich Departamento de Zoologia Instituto de Biociencias Universidade de Sao Paulo C. P. 20520 01498 - Sao Paulo, SP Brazil

Hideki Fujino Department of Biological and Chemi-

cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan

Jun Fujita Department of Biological and Chemi-


Takeshi Fujiwara Miyagi Prefectural Freshwater

Fisheries Station Taiwa-cho, Miyagi 981-36 Japan

Takeshi Fukushima W. Alton Jones Cell Science Center 10 Old Barn Rd. Lake Placid NY 12946 USA

Emiko Furuta Department of Anatomy Dokkyo University School of Medi

cine Mibu, Tochigi 321-02 Japan

Elvezio Ghirardelli Dipartimento Biologia-Universita Via Valerio 32 Trieste 34/34 Italy

Toshiki Gotoh Hokkaido Sapporo Inakita Senior

High School 254 Teine Yamaguchi Sapporo 006 Japan

Vittorio Gremigni Dipartimento di Biomedicina Speri-


M. G. Gremigni-Querci Dipartimento di Biomedicina Speri-


Kenya Hakota Goshogawara Senior High School 3-3 Nakahirai-machi Goshogawara 037 Japan

Josef Hauser Universidade do Vale do Rio Dos

Sinos-UNISINOS Caixa Postal, 275 93.020 - Sao Leopoldo/RS Brazil

Jan Hendelberg Department of Zoology University of Goteborg P.O. Box 25059 S-400 31 Goteborg Sweden

Isao Hori Department of Biology Kanazawa Medical University Uchinada-machi Ishikawa-ken 920-02 Japan

Shinri Horiuchi Department of Biology College of General Education Osaka University Toyonaka Osaka 560 Japan

Taro Hoshina

Xl

Technical Division, Detergents and Personal Products

Nippon Lever B.V. Shibuya Higashiguchi Bldg. 22-3 Shibuya 2-chome Tokyo 150 Japan

Kouichi Hoshino Department of Biological and

Chemical Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan

Jie Hu Department of Computational Ma-

thematics Xian, Jiaotong University Xian, Shanxi People's Republic of China

Takeshi Inoue Department of Physiological Chemis-

try Faculty of Pharmaceutical Sciences Teikyo University Sagamiko, Kanagawa 199-01 Japan

xii

Sachiko Ishida Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Saburo Ishii Division of Cell Science Central Research Laboratory Fukushima Medical College I Hikariga-oka Fukushima 960-12 Japan

Boris I. Joffe Zoological Institute of the USSR Academy of Sciences 189034 Universitetskaya nab. I St Petersburg USSR

Ulf Jonde1ius Department of Zoology University of Goteborg P.O. Box 25059 S-400 31 Goteborg Sweden

Masaharu Kawakatsu Biological Laboratory Fuji Women's College Kita-16 Nishi-2, Kita-ku Sapporo (Hokkaido) 001 Japan

Masashi Kodama Faculty of Education Hirosaki University 1 Bunkyo-cho Hirosaki 036

Yoshihiro Kon-no Sendai Ikeui Gakuen Senior High

School 2-4 Miyagino Sendai 983 Japan

Yutaka Koshida Department of Biology College of General Education Osaka University Toyonaka Osaka 560 Japan

A. G. Kostenko Institute of Zoology of the Ukrainian

Academy of Sciences Lenin str. 15 Kiev 252650 USSR

Satoshi Kuchiiwa Department of Anatomy Hirosaki University School of Medi-

cine 5 Zaifu-cho Hirosaki 036 Japan

Toshiko Kuchiiwa Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Masato Kudoh Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Mitsuaki Kurotaki School of Allied Medical Sciences Hirosaki University 66-1 Hon-cho Hirosaki 036 Japan

Alberto Lanfranchi Dipartimento di Scienze Del Com-

portamento Animale e Dell'Uomo Via Volta 6 1-56100 Pisa Italy

A. Legniti Institut fUr Zoologie Universitat Innsbruck TechnikerstraBe 25 A-6020 Innsbruck Austria

Lei Li Department of Computational Ma-

thematics Xian, Jiaotong University Xian, Shanxi People's Republic of China

Paula Lindroos Institute of Biology Abo Akademi Porthansgatan 3-5 SF-20500 Abo Finland

Sridam Mahapatra Department of Zoology Visva-Bharati University Santiniketan-731235 West Bengal India

Naoya Makino Biology Laboratory Tokyo Medical College 6-1-1 Shinjuku Shinjuku-ku Tokyo 160 Japan

Yuri V. Mamkaev Zoological Institute of the Academy

of Sciences Universitetskaya nab. I 199034 St Petersburg USSR

Roberto Marzari Department of Biology University of Trieste Via Alfonso Valerio 32 1-34127 Trieste Italy

Ryoichi Matsuda W. Alton Jones Cell Science Center,

Inc. 10 Old Barn Road Lake Placid, NY 12946 USA

Akira Matsuo Department of Oral Anatomy School of Dentistry Higashi-Nippon-Gakuen University Ishikari-Tobetsu Hokkaido 061-02 Japan

Nancy Mead Department of Biology University of Nevada Reno, NV 89557 USA

Robert W. Mead Department of Biology University of Nevada Reno, NV 89557 USA

Hideo Minegishi The Senior High School of Japan

Women's University Toyokawa-cho, Takada Bunkyo-ku Tokyo 112 Japan

Humberto 1. A. Moretto Casilla de Correo 113 Sucursal 13 Buenos Aires Rep. Argentina

David Morita Department of Anatomy &

Neurobiology Colorado State University Fort Collins, CO 80523 USA

Michio Morita Department of Anatomy &


Shoko Morita Department of Anatomy &


Hitoshi Murayama 4-301-2, Nakazawa Nagaoka, Niigata-ken 940 Japan

Galena-Vanzetti Murina Institute of Biology of South Seas Academy of Sciences Uk.SSR 2, Nahimov St. Sevastopol, 335000 USSR

Yozo Naoki Mukainakano Gakuen High School 2-3 Saikawa, Mukainakano Morioka, Iwate 020 Japan

Fumio Ni-imura Biological Laboratory Nagano Women's Junior College 11-3, Miwa 9-chome Nagano 380 Japan

Tohsaku Nikara School of Allied Medical Sciences Hirosaki University 66-1 Hon-cho Hirosaki 036 Japan

Yuji Nishimura Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Takaharu N umakunai Marine Biological Station of Asamu-

shi Tohoku University 9-Sakamoto, Asamushi Aomori 039-34 Japan

Yoshitaka Obara Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Jean Ogren Department of Biology Wilkes University Wilkes-Barre, PA 18766 USA

Robert E. Ogren Department of Biology Wilkes University Wilkes-Barre, PA 18766 USA

xiii

Keiko Ohkawa College of Liberal Arts (Biology) Hirosaki University 1 Bunkyo-cho Hirosaki 036 Japan

Kazunori Ohnishi Department of Dermatology Faculty of Medicine Gunma University Maebashi Gunma 371 Japan

Iwashiro Oki Osaka Environmental Project

Authority MFC7F 1-2-15 U chihon-machi Chuoh-ku Osaka 540 Japan

Hidefumi Orii Division of Morphogenesis, NIBB Nishinomiya 38, Myodaiji-cho Okazaki, Aichi 444 Japan

Kenji Osanai Marine Biological Station of Asamu-

shi Tohoku University 9-Sakamoto, Asamushi Aomori 039-34 Japan

Irmeli Palmberg Department of Biology Faculty of Education Abo Akademi PB 31 I 65101 Vasa Finland

Uday Raj Institute of Marine Resources The University of the South Pacific Suva 1168 Fiji

XIV

S. Reindl Institut fUr Zoologie Universitat Innsbruck TechnikerstraBe 25 A-6020 Innsbruck Austria

Dietmar Reiter Institut fUr Zoologie Universitat Innsbruck TechnikerstraBe 25 A-6020 Innsbruck Austria

Maria Reuter Institutionen fOr Biologi Department of Biology Abo Akademi Porthansgatan 3 SF-20500 Abo 50 Finland

Reinhard Rieger Institut fUr Zoologie Universitat Innsbruck TechnikerstraBe 25 A-6020 Innsbruck Austria

Klaus Rohde Department of Zoology University of New England Armidale, N.S.W. 2351 Australia

Rafael Romero Departament de Genetica Facultat de Biologia Universitat de Barcelona Diagonal 645 08028 Barcelona Spain

Akira Sakurai Department of First Anatomy School of Medicine Kobe University Kobe 650 Japan

Norita Sakurai Department of Biological and Chemi-


Takashige Sakurai Division of Cell Science Central Research Laboratory Fukushima Medical College I Hikariga-oka Fukushima 960-12 Japan

W. Salvenmoser Institut fUr Zoologie Universitat Innsbruck TechnikerstraBe 25 A-6020 Innsbruck Austria

Massimo Sarperi Dipartimento di Biomedicina Speri-


Ken Sasaki Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Yuri Sasaki Biology Laboratory Tokyo Medical College 6-1-1 Shinjuku Shinjuku-ku Tokyo 160 Japan

Koh Satoh Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Ernest Schockaert Dept. S.B.M. Limburgs Universitair Centrum B-361O Diepenbeek Belgium

Naomi Seo Biology Laboratory Tokyo Medical College 6-1-1 Shinjuku Shinjuku-ku, Tokyo 160 Japan

Takao Shinozawa Department of Biological and Chemi-


Takashi Shirakawa Fuji Rebio Inc. 51 Komiya-cho, Hachiohji Tokyo 192 Japan

Mikio Shirasawa Biology Laboratory Tokyo Medical College 6-1-1 Shinjuku Tokyo 160 Japan

Yasuko Shirasawa Biology Laboratory Tokyo Medical College 6-1-1 Shinjuku Shinjuku-ku Tokyo 160 Japan

P. Simonsberger Department of Zoology Universitat Salzburg Hellbrunnerstr. 34 A-5020 Salzburg Austria

A. I. Solonchenko Institute of Biology of South Seas of

the Academy of Sciences Ukraina S.S.R. 2 N akhimov Pr. Sevastopol' 335000 USSR

Beate Sopott-Ehlers II. Zoologisches Institut und Museum

der Georg-August-Universitat Berliner Str. 28 3400 Gottingen Germany

Marc Aureli Soriano Departament de Genetica Facultat de Biologia Universitat de Barcelona Diagonal 645 08028 Barcelona Spain

Kazuo Sugiyama Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Lauro Sumoy 56 Old Mill Lane West Harford, CT 06107 USA

Ken-ichi Tajika Department of Biology Nihon University School of Medicine Oyaguchi, Itabashi Tokyo 173 Japan

Masako Tajika Department of Biology Nihon University School of Medicine Oyaguchi, Itabashi Tokyo 173 Japan

Masayuki Takai Biology Laboratory Saga Medical School Nabesima Saga 840-01 Japan

Sachiko Tamura Osaka Prefectural Institute of Public

Health Nakamichi-l-chome 3-69 Higashinari-ku Osaka 537 Japan

Hisanobu Teshima Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Shukuko Teshirogi Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Wataru Teshirogi Department of Biology Faculty of Science Hirosaki University Hirosaki 036 Japan

Oleg Timoshkin 664033 Limnological Institute Irkutsk Russia USSR

Shuji Tohno Administration Bureau Hirosaki University 1 Bunkyo-cho Hirosaki 036 Japan

Seth Tyler Department of Zoology Murray Hall University of Maine Orono, ME 04469 USA

Katsumi Wakabayashi Hormone Assay Center Institute of Endocrinology Gunma University Maebashi Gunma 371 Japan

Kenji Watanabe Himeji Institute of Technology Faculty of Science Department of Life Science Himeji 671-22 Japan

M. Wikgren Institute of Biology Abo Akademi Porthansgatan 3-5 SF-20500 Abo 50 Finland

Leigh Winsor Department of Zoology James Cook University of North

Queensland Townsville, Queensland, Q 4811 Australia

Motoichi Yagihashi Biological Laboratory Oogaki Women's Junior College 606 Nakano-cho Oogaki 503 Japan

Cecilia Mikie Yamada Departamento de Biologia Instituto de Biociencias Universidade de Sao Paulo c.P. 11461 05499 - Sao Paulo, SP Brazil

Keiko Yamada

xv

Department of Medical Zoology Dokkyo University School of Medi-

cine 880, Kitakobayashi, Mibu Tochigi 321-02 Japan

Makoto Yamada Kuroishi Senior High School 65 Nishigaoka Kuroishi 036-03 Japan

Tatsuya Yamada Ogano High School Ogano-machi, Chichibu-gun Saitama 368-01 Japan

Y oshihisa Yamashita Central Research Laboratories New Drug Development Research

Center, Inc 452-1 Toiso, Eniwa Hokkaido 061-14 Japan

Nobuko Yamasu Department of Biology College of Liberal Arts Ryukyu University Senbaru 1, Nishihara Okinawa 903-01 Japan

XVI

Terufumi Yamasu Department of Biology College of Liberal Arts Ryukyu University Senbaru 1, Nishihara Okinawa 903-01 Japan

Chifumi Yamaya Noboribetsu Ohtani Senior High

School 2-1,2-1 Sakuragi-cho, Noboribetsu Hokkaido 059-03 Japan

Bong Y. Yoo Department of Biology Bag Service Number 45111 University of New Brunswick Fredericton, N.B., E3B 6EI Canada

W ataru Yoshida Department of Biological and Chemi-


Yasuhiro Y oshizawa Department of Biological and Chemi-


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Contents

Preface............. .... .. ... ....... .... ... ....................................... v

Committees of the Symposium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vi Acknowledgments ................................................................. Vll

Group photograph ................................................................. Vlll

List of Participants and Contributors ................................................. x

Dedication Eveline du Bois-Reymond Marcus (1901-1990) ........................................ XXI

Biography of Dr Marcus by Diva Diniz Correa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. xxiii List of publications by Dr Marcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. XXIV

Regeneration and asexual reproduction Differentiation during asexual reproduction and regeneration in a microturbellarian

by Irmeli Palmberg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Studies of the budding process in Convolutriloba retrogemma (Acoela, Platyhelminthes)

by Jan Hendelberg & Bertil Akesson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Differentiation of epidermal cells in the regenerating planarian Dugesia japonica

by Isao Hori ................................................................. 19 Effect of timing of cutting on patterning and proportion regulation during regeneration of the

planarian Dugesia tigrina by Robert W. Mead ........................................................... 25

Inhibition of planarian regeneration by melatonin by Yasuhiro Yoshizawa, Katsumi Wakabayashi & Takao Shinozawa ................. 31

Differences in blastema-associated proteins according to position along body axis in regenerating planarians

by Elvezio Ghirardelli, Roberto Marzari & Enrico A. Ferrero. . . . . . . . . . . . . . . . . . . . . . . . 41 Dedifferentiation and redifferentiation of the penis of the freshwater planarian Bdellocephala brunnea

by M. Kudoh, H. Teshima & W. Teshirogi ....................................... 47 Pharyngeal regeneration in the land planarian Bipalium kewense

by Yasuko Shirasawa & Naoya Makino. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 See also article by Hoshina & Teshirogi, p. 61

Cancer Two-stage carcinogenesis in the planarian

by Michio Morita. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 Formation of malformed pharynx and neoplasia in the planarian Bdellocephala brunnea following

treatment with a carcinogen by Taro Hoshina & Wataru Teshirogi ..................................... , . . . . . . 61

Regeneration and the relationship between regeneration and development of cancerous tumors by Josef Hauser .............................................................. 71

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Markers for planarian tissues Monoclonal antibodies as markers of specific cell types and regional antigens in the freshwater

planarian Dugesia (G.) tigrina by Rafael Romero, Joan Fibla, David Bueno, Lauro Sumoy, Marc A. Soriano & Jaume Bagufia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 73

Production of cell- and tissue-specific monoclonal antibodies in the freshwater planarian Phagocata vivida

by Takashi Shirakawa, Akira Sakurai, Takeshi Inoue, Ken Sasaki, Yuji Nishimura, Sachiko Ishida & Wataru Teshirogi ..................................................... 81

Presence of rhodopsin-like proteins in the planarian head by Jun Fujita, Noriko Sakurai & Takao Shinozawa ................................ 93

Development and reproductive systems Analytical studies of the ultrastructure and movement of the spermatozoa of freshwater triclads

by Sachiko Ishida, Yoshihisa Yamashita & Wataru Teshirogi . . . . . . . . . . . . . . . . . . . . . . . . 95 Ultrastructural features of cocoon-shell globules in the vitelline cells of neoophoran platyhelminths

by Vittorio Gremigni & Alessandra Falleni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 105 An electron-microscopic study of syncytium formation during early embryonic development of the

freshwater planarian Bdellocephala brunnea by Takeshige Sakurai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 113

Organization and differentiation of the body-wall musculature in Macrostomum (Turbellaria, Macrostomidae)

by Reinhard Rieger, W. Salvenmoser, A. Legniti, S. Reindl, H. Adam, P. Simonsberger & S. Tyler ..................................................................... 119

Ultrastructural aspects of nervous-system and statocyst morphogenesis during embryonic development of Convoluta psammophila (Turbellaria, Acoela)

by Enrico A. Ferrero & Celina Bedini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 131 Developmental sequence for the copulatory organs of Kontikia mexicana with remarks on taxonomic significance (Turbellaria: Tricladida: Geoplanidae)

by Robert E. Ogren ........................................................... 139 Notes on the biology of the freshwater planarian Dugesia bengalensis (Platyhelminthes Turbellaria

Tricladida) by Ajit Kumar Aditya & Sridam Mahapatra ...................................... 145

Karyology Further survey of chromosomal polymorphisms in the freshwater planarian Polycelis auriculata

by Wataru Teshirogi, Fumio Ni-imura & Sachiko Ishida ............................ 147 Karyological and taxonomic studies of Dugesia japonica from the Southwest Islands of Japan - II

by Sachiko Tamura, Iwashiro Oki & Masaharu Kawakatsu ......................... 157 Karyology of four land-planarian species of the genus Bipalium from Japan

by Iwashiro Oki, Sachiko Tamura, Robert E. Ogren & Masaharu Kawakatsu .......... 163 Cytogenetic studies of two land-planarian species from Brazil: Geoplana marginata and Issoca

rezendei (Tricladida, Terricola) by Eunice J. C. de Almeida, Cecilia M. Yamada & Eudoxia M. Froehlich ............. 169

Analysis of ploidy in a planarian by flow cytometry by Kouichi Hoshino, Kazunori Ohnishi, Wataru Yoshida & Takao Shinozawa ......... 175

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Cellular processes Food ingestion by planarian intestinal phagocytic cells - a study by scanning electron microscopy

by Saburo Ishii & Takashige Sakurai ............................................ 179 Experiments with culture media for planarian cells

by Takeshi Fukushima & Ryoichi Matsuda ....................................... 187 Phagocytic response of planarian reticular cells to heat-killed bacteria

by Michio Morita ............................................................. 193

Nervous system On the number and spatial distribution of catecholamine-containing (GA-positive) neurons in some

higher and lower turbellarians - a comparison by B. I. loffe ................................................................. 201

Catecholamines demonstrated by glyoxylic-acid-induced fluorescence and HPLC in some microturbellarians

by Maria Reuter & Krister Eriksson ............................................. 209 Are there differences between proseriates and lower flatworms in ultrastructure of the nervous

system? by Maria Reuter .............................................................. 221

Immunoreactivity to a specific echinoderm neuropeptide in the nervous system of the flatworm Macrostomum hystricinum marinum (Turbellaria, Macrostomida)

by D. Reiter & M. Wikgren .................................................... 229

Sensory structures Comparative morphology of photoreceptors in free-living plathelminths - a survey

by Beate Sopott-Ehlers ........................................................ 231 Comparative morphological studies on the visual systems in a binocular and a multi-ocular species

of freshwater planarian by Toshiko Kuchiiwa, Satoshi Kuchiiwa & Wataru Teshirogi ........................ 241

Ultrastructure of the photoreceptors of Allostoma sp. (Plathelminthes, Prolecithophora) by Alberto Lanfranchi ......................................................... 251

The multiple eyes of Polycelis. 1. Relation between the number of eyes and body length by Masuo Aikawa & Atsumi Shimozawa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 257

Comparative morphology of statocysts in the Plathelminthes and the Xenoturbellida by Ulrich Ehlers .............................................................. 263

See also article by Ferrero & Bedini, p. 131

Ultrastructure of other systems Fine structure and function of ocelli and sagittocysts of acoel flatworms

by Terufumi Yamasu .......................................................... 273 Extracellular matrix in some microturbellarians

by Paula Lindroos & Maria Reuter .............................................. 283 Anatomy and ultrastructure of the proboscis in Mesorhynchus terminostylis (Platyhelminthes,

Rhabdocoela) by Alain De Vocht ............................................................ 291

Unciliated body surface in three species of the Umagillidae (Dalyellioida, Platyhelminthes) by Ulf londelius .............................................................. 299

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Phylogeny On the phylogenetic significance of sagittocysts and copulatory organs in acoel turbellarians

by Yu. V. Mamkaev & A. G. Kostenko .......................................... 307 The evolution of protonephridia of the Platyhelminthes

by Klaus Rohde .............................................................. 315 Turbellaria Lecithoepitheliata: morphology, systematics, phylogeny

by O. A. Timoshkin ........................................................... 323

Taxonomy Polyclad turbellarians collected on the Osaka University Expedition to Viti Levu, Fiji, in 1985, with

remarks on distribution and phylogeny of the genus Discoplana by K.-I. Tajika, U. Raj, S. Horiuchi, Y. Koshida .................................. 333

Temnocephalan symbionts of the freshwater crayfish Cherax quadricarinatus from northern Australia

by Lester R. G. Cannon ....................................................... 341 Methods for taxonomic and distributional studies of terrestrial flatworms (Tricladida: Terricola)

by L. Winsor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 349 On the taxonomic status of a Dugesia species from Buenos Aires (Platyhelminthes: Turbellaria:

Tricladida) by Humberto J. A. Moretto .................................................... 353

See also article by Ogren, p. 139

Biogeography and ecology Geographical distribution of Polycelis (Seidlia) auriculata in Japan

by Masaharu Kawakatsu, Tatsuya Yamada, Hitoshi Murayama & Y6z6 Naoki ........ 355 Northern circumpolar distribution of brackish-water plathelminths

by Peter Ax .................................................................. 365 Plathelminths in tropical intertidal sediments of northeastern Australia

by Sabine Dittmann ........................................................... 369 Toxicology of planarians

by Jay Boyd Best & Michio Morita .............................................. 375

Commensals Commensals of Mytilus galloprovincialis in the Black Sea: Urastoma cyprinae (Turbellaria) and

Polydora ciliata (Polychaeta) by G.-V. Murina & A. I. Solonchenko ........................................... 385

See also article by Cannon, p. 341

History of science History of the study of Turbellaria in Japan

by Masaharu Kawakatsu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 389

This volume is dedicated to the memory of Eveline du Bois-Reymond Marcus

(6 October 1901-31 January 1990)

Hydrobiologia 227: xxiii-xxvi, 1991. S. Tyler (ed.), Turbellarian Biology.

Dr Eveline du Bois-Reymond Marcus

Diva Diniz Correa

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Departamento de Zoologia, Instituto de Biociencias, Universidade de sao Paulo, C. P. 20520, 01498 -sao Paulo, SP, Brazil

Eveline du Bois-Reymond Marcus was born in Berlin, Germany, October 6th, 1901, as the youngest daughter of Rene and Frieda du BoisReymond; she died in Sao Paulo, Brazil, January 31st, 1990.

She grew up in an environment of people interested in science. Her father was a teacher of physiology at Berlin University, and her grandfather, Emile du Bois-Reymond, also a teacher of physiology in the same University, was famous as a scientist. When she was only nine years old, she started to watch small animals in her father's microscope; and she continued so, watching small animals, almost her whole life, until her health began to decline.

From 1918 to 1919 she attended courses in laboratory techniques, and from 1920 to 1923 she worked as a technician in the universities of Bonn, G()ttingen, and Berlin. From 1923 to 1924 she started attending zoology courses at the University of Berlin, thinking about a career as a researcher in zoology. She did not finish her studies, however, because there she met a teacher of zoology, Ernst Marcus; they were married March 11, 1924, and started a life together that lasted 44 years, thoroughly dedicated to zoology. From their marriage to the day of his death, June 30, 1968, it became impossible to talk about their scientific lives separately.

In 1924 she started a close collaboration with her husband, first in Berlin (1924-1936) and later in Sao Paulo (1936-1968); this included study of several invertabrate groups, such as Protozoa, Ctenophora, Turbellaria, Nemertinea, Oligochaeta, Archiannelida, Tardigrada, Onychophora, Bryozoa, Pantopoda, and Gastropoda. They

dedicated a great many hours to zoology. It was sometimes said that he was the 'writer' and she was the 'technician', but in fact all subjects were always discussed by both. All their published papers were the result of this close work together, but, to justify his full-time position as a professor, only his name appeared as author at first, as she never accepted a remunerated job. While all their first papers list only E. Marcus as author, her collaboration was always mentioned in the first lines of the publications; in addition, many new species were named for her, as 'evelinae'. Eventually, her name started to appear, sometimes with his, as 'Marcus, E. & E.', sometimes as only 'Marcus, E. du Bois-Reymond'.

In 1936 they left Berlin to live in Sao Paulo, Brazil, as Ernst Marcus had been invited to teach in the Faculdade de Filosofia, Ciencias e Letras of the Universidade de Sao Paulo. In 1940 they became Brazilian citizens and commemorated the fact by purchasing a gold medal with the symbol of the Brazilian Republic.

Looking back to their list of publications, we recognize that some groups received a special preference and were exhaustively explored by them. Among these was the large group of the turbellarians.

During the Second World War (1939-1945), they could not travel to the sea coast due to their German origin. They then dedicated their time to the study of freshwater animals. In 1945 Ernst Marcus presented to the University of Sao Paulo a large thesis about microturbellarians and received for that the position of Full Professor of Zoology.

Soon after the end of the war, they started to

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travel to the coast of Sao Paulo State where they found an immense and unexplored number of turbellarian species and of other marine groups. In number of written pages, number of drawings, number of new species, or number of keys in their publications about turbellarians, their productivity was so large as to be nearly impossible to grasp. Also in other animal groups, particularly the Bryozoa, Tardigrada, and Gastropoda, the number of papers they published is extraordinarily large. The full list of their publications, dating from 1924 to 1970, includes about 220 titles. From 1970 to 1985, Eveline Marcus, then alone, published about 30 papers dealing mostly with opisthbranch molluscs from warm waters of the Western Atlantic Ocean.

In 1973 she received the title of Honorary Member of the Brazilian Malacological Society and in 1979 was elected Honorary Member of the London Malacological Society. In 1976 the University of Sao Paulo, Brazil, awarded her the title of Doctor Honoris Causa owing to her significant contribution to the development of zoological research in Brazil; the University of Marsaille, France, awarded her the title of Doctor Honoris Causa in 1988. In 1976 she became co-editor of the publication Studies on the Neotropical Fauna.

Considering these accomplishments, zoologists the world over accept the Marcus name (Ernst Marcus and Eveline du Bois-Reymond Marcus) as among the first line of the most important zoologists of our century. We, her friends and colleagues, all agree that she was a remarkable woman.

Publications by Dr Eveline du Bois-Reymond Marcus

Marcus, Ev., 1943. Protozoa associados a vermes Iimnicos. An. Acad. bras. Cienc. 15: 359-371, 1 t.

Marcus, Ev., 1943. Sabre Tardigrados Brasileiros. Comun. Zoo!. Mus. Hist. Nat. Montev. 1(13): 1-20.

Marcus, Ev., 1944. Notes on the freshwater Oligochaeta from Brazi!. Comun. Zool. Mus. Hist. Nat. Montev. 1(20): 1-18.

Marcus, Ev., 1946. On a new Brazilian form, Fredericella sultana, etc. Comun. Zoo!. Mus. Hist. Nat. Montev. 2(31): 1-10.

Marcus, Ev., 1946. On a new archianne1id, Saccocirrus gab-

riellae. Comun. Zool. Mus. Hist. Nat. Montev. 2(37): 1-6. Marcus, Ev., 1947. naidids and tubificids from Brazi!. Comun.

Zoo!. Mus. Hist. Nat. Montev. 2(44): 1-18. Marcus, Ev., 1947. Nerilla mediterranea from Brazil. Comun.

Zool. Mus. Hist. Nat. Montev. 2(45): 1-6. Marcus, Ev. 1948. An Amazonian heteronemertine. Bol. Fac.

Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 13: 93-109. Marcus, Ev., 1948. Further archiannelids from Brazil. Comun.

Zoo!. Mus. Hist. Nat. Montev. 2(48): 1-17. Marcus, Ev., 1949. Further notes on naidids .... Comun. Zool.

Mus. Hist. Nat. Montev. 3(51): I-II. Marcus, Ev., 1949. Phoronis ovalis from Brazil. Bo!. Fac.

Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 14: 157-172.

Marcus, Ev., 1950. A marine tubificid from Brazil. Comun. Zool. Mus. Hist. Nat. Montev. 3(59): 1-6.

Marcus, Ev., 1950. A new loxosomatid from Brazil. Bo!. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 15: 193-202.

Marcus, Ev., 1951. On South American geoplanids. Bo!. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 16: 217-256.

Marcus, Ev., 1951. Two freshwater Turbellaria from Nata!. Ann. Natal Mus. 12(1): 73-80.

Marcus, E. & Ev. Marcus, 1951. Contributions to the Natural History of Brazilian Turbellaria. Comun. Zoo!. Mus. Hist. Nat. Montev. 3(63): 1-25.

Marcus, Ev., 1952. A hermaphrodite pantopod. An. Acad. bras. Cienc. 24: 23-30.

Marcus, Ev., 1952. On South American Malacopoda. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zoo!. 17: 189-209.

Marcus, Ev., 1952. A new tubificid from the Bay of Montevideo. Comun. Zool. Mus. Hist. Nat. Montev. 3(56): 1-6.

Marcus, Ev., 1953. Some South American triclads. An. Acad. bras. Cienc. 25: 65078.

Marcus, Ev., 1953. The opisthobranch Pseudovermis. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 18: 109-127.

Marcus, Ev., 1953. Bryozoa from Lake Titicaca. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool.; 18: 149-163.

Marcus, E. & Ev. Marcus, 1954. The nudibranch Rhodope from South America. Comun. Zool. Mus. Hist. Nat. Montev. 4(68): 1-8.

Marcus, E. & Ev. Marcus, 1954. Uber Philinoglossacea und Acochlidiacea. Kieler Meeresforsch. 10: 215-223.

Marcus, E. & Ev. Marcus, 1954. On Onchidella indolens (Gould, 1852). Bol. Inst. Oceanogr. 5: 87-94.

Marcus, Ev., 1955. Chave dos Polycladida do litoral de Sao Paulo. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 19: 281-288.

Marcus, E. & Ev. Marcus, 1955. A new Peripatus from Minas Gerais, Brazil. An. Acad. bras. Cienc. 27: 189-193.

Marcus, E. & Ev. Marcus, 1955. Uber Sandopisthobranchier. Kieler Meeresforsch. 11: 230-243.

Marcus, Ev., 1955. On Turbellaria and Polygordius from the Brazilian coast. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 20: 19-53.

Marcus, E. & Ev. Marcus, 1956. On two sacoglossan slugs from Brazil. Am. Mus. Novit. (1796): 1-21.

Marcus, E. & Ev. Marcus, 1956. On the tectibranch gastropod Cylindrobulla. An. Acad. bras. Cienc. 28: 119-128.

Marcus, E. & Ev. Marcus, 1955. Sea-hares and side gilled slugs from Brazil. Bol. Inst. Oceanogr. 6: 3-49.

Marcus, Ev., 1957. On Turbellaria. An. Acad. bras. Cienc. 29: 153-191.

Marcus, Ev., 1957. Neue Entoprocten aus der Gegend von Santos. Zool. Anz. 159: 68-75.

Marcus, E. & Ev. Marcus, 1957. Turbellaria. Expl. HydrobioI. Lacs. Kivu, Edouard et Albert. Res. Sci. V. 3(2): 27-52.

Marcus, E. & Ev. Marcus, 1957. On Phyllaplysia engeli. Basteria 21(4-5): 53-66.

Marcus, Ev., 1958. On some Prosobranchia from the coast of Sao Paulo. Bol. Inst. Oceanogr. 7: 3-29.

Marcus, Ev., 1958. Vallicula multiformis Rankin, 1956, from Brazil. Bol. Inst. Oceanogr. 7: 87-91.

Marcus, E. & Ev. Marcus, 1958. Opistobranchia aus dem Schill von Helgoland. Kieler Meeresforsch. 14: 91-96.

Marcus, Ev., 1958. On South American Turbellaria. An. Acad. bras. Cienc. 30: 391-417.

Marcus, Ev., 1959. Ein neuer Pantopode aus Foraminiferensand. Kieler Meeresforsch. 15: 105-107.

Marcus, E. & Ev. Marcus, 1959. Notes on Aplysia. Bol. Inst. Oceanogr. 8: 3-21.

Marcus, E. & Ev. Marcus, 1959. Studies on Olividae. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 22: 99-188.

Marcus, E. & Ev. Marcus, 1959. On the reproduction of Olivella. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 22: 189-200.

Marcus, E. & Ev. Marcus, 1959. Turbellaria from Madeira and Azores. Lunds University Expedition, Report 3, 1957. Bol. Mus. Munic. Funchal 12(30): 15-42.

Marcus, Ev., 1960. Tardigrada from Cura.;;ao, Bonaire and Los Testigos. Stud. Fauna Cura,<ao Other Caribb. lsI. 10: 52-57.

Marcus, E. & Ev. Marcus, 1960. Opisthobranch aus dem Roten Meer und von den Malediven. Akad. Wiss. Lit. Abh. math.-naturwiss. KI. (Mainz) 1959, no. 12, p. 871-934.

Marcus, Ev., 1960. Notes on the freshwater polychaete Lycastopsis from Cura.;;ao. Stud. Fauna Caracao Other Caribb. lsI. 10: 58-63.

Marcus, E. & Ev. Marcus, 1960. Opisthobranchs from American Atlantic warm waters. Bull Mar. Sci. Gulf Caribb. 10: 129-203.

Marcus, E. & Ev. Marcus, 1960. Some opisthobranchs from the Northwestern Gulf of Mexico. Publ. Inst. Mar. Sci. Univ. Tex. 6: 251-264.

Marcus, E. & Ev. Marcus, 1961. On Hastula cinerea. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 23: 25-34.

Marcus, E. & Ev. Marcus, 1961. Siphonaria hispida. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 23: 107-130.

Marcus, E. & Ev. Marcus, 1961. On Tricolia affinis cruenta. Bol. Fac. Filos. Cienc. Let. Sao Paulo Ser. Zool. 23: 171-198.

Marcus, E. & Ev. Marcus, 1961. On Coryphellina rubrolineata. etc. Proc. Malac. Soc. Lond. 34: 224-227.

Marcus, E. & Ev. Marcus, 1962. On Stylocheilus citrinus, etc. Proc. Malac. Soc. Lond. 35: 16-19.

Marcus, E. & Ev. Marcus, 1962. A new species of Gnathodoridacea. An. Acad. bras. Cienc. 34: 269-275.

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Marcus, E. & Ev. Marcus, 1962. On lunulitiform Bryozoa. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool. 24: 281-324.

Marcus, E. & Ev. Marcus, 1962. Opisthobranchs from Florida and the Virgin Islands. Bull. mar. Sci. Sci. GulfCaribb. 12: 450-488.

Marcus, E. & Ev. Marcus, 1963. On Leucozonia nassa. Bol. Fac. Filos. Cienc. Let. Sao Paulo Ser. Zool. 24: 11-30.

Marcus, E. & Ev. Marcus, 1963. On Uncancylus ticagus. Bol. Fac. Filos. Cienc. Let. Sao Paulo Ser. Zool. 24: 217-254.

Marcus, E. & Ev. Marcus, 1963. A new Pycnogonum from Brazil. Bol. Inst. Oceanogr. 12: 3-10.

Marcus, E. & Ev. Marcus, 1963. Mesogastropoda von der Kuste Sao Paulo. Akad. Wiss. Lit. Abh. math.-naturwiss. KI. (Mainz), pp. 1-105.

Marcus, E. & Ev. Marcus, 1963. Opisthobranchs from the Lesser Antilles. Stud. Fauna Cura.;;ao Other Caribb. lsI. 19: 1-76.

Marcus, E. & Ev. Marcus, 1964. On Brazilian supralitoral and brackish water snails. Bol. Inst. Oceanogr. 13: 41-52.

Marcus, E. & Ev. Marcus, 1964. Berzeichnis der euthyneuren Meereschnecken Brasiliens. Beitr. Neotrop. Fauna 3: 195-206.

Marcus, E. & Ev. Marcus, 1964. On Cerithium atratum, etc. Bull. mar. Sci. Gulf Caribb. 14: 494-510.

Marcus, E. & Ev. Marcus, 1965. On Rissolina chesnelii (Michaud, 1830). Proc. Malac. Soc. Lond. 36: 163-172.

Marcus, E. & Ev. Marcus, 1965. On the dove-shell Anachis pulchella (Blainev.) An. Acad. bras. Cienc. 36: 359-366.

Marcus, E. & Ev. Marcus, 1965. On Haliotinella patinaria. Bull. mar. Sci. 15: 211-215.

Marcus, E. & Ev. Marcus, 1966. On Brazilian supratidal and estuarine Snails. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool., 25: 19-104.

Marcus, E. & Ev. Marcus, 1966. On two Ellobiidae from southern Brazil. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo Ser. Zool., 25: 425-466.

Marcus, Ev., 1965. Drei neue neotropische Turbellarien. Sitzungsber. Finn. Akad. Wiss. 5: 129-135.

Marcus, E. & Ev. Marcus, 1966. Opisthobranchs from tropical West Africa. Stud. Trop. Oceanogr. (Miami) 4(9): 152-208.

Marcus, E. & Ev. Marcus, 1966. Systematische Ubersicht der Polykladen. Zool. Beitr. 12: 319-344.

Marcus, E. & Ev. Marcus, 1967. American opisthobranch mollusks. Stud. Trop. Oceanogr. (Miami) (6): 256 p.

Marcus, E. & Ev. Marcus, 1967. Opisthobranchs from the southwestern Caribbean Sea. Bull. mar. Sci. 17: 597-628.

Marcus, E. & Ev. Marcus, 1967. Some Opisthobranchs from Sapello Island, Georgia, U.S.A. Malacologia 6: 199-222.

Marcus, E. & Ev. Marcus, 1968. Uber einige Subulinidae (Pulmonata) von Sao Paulo. Beitr. Neotrop. Fauna 5: 186-208.

Marcus, E. & Ev. Marcus, 1968. On the prosobranchs Ancilla dimidiata and Marginella fraterculus. Proc. Malac. Soc. Lond. 38: 55-69.

Marcus, E. & Ev. Marcus, 1968. Polycladida from Cura,<ao and faunistic ally related regions. Stud. Fauna Cura.;;ao Other Caribb. lsI. 26: 1-133.

Marcus, E. & Ev. Marcus, 1968. Neue brasilianische Loxosomen. Zool. Beitr. 14: 203-212.

xxvi

Marcus, E. & Ev. Marcus, 1968. Flabellina engeli, a new nudibranch from Cura~ao. Beaufortia 15(200): 139-142.

Marcus, Ev., 1968. Ernst Marcus. Atti. Soc. ital. Sci. nat. Mus. Stor. nat. Milano, 108: 7-12.

Marcus, E. & Ev. Marcus, 1969. Some Opisthobranchs from Ivory Coast. Bull. Inst. fro Afr. Noire Ser. A, 30(4): 1334-1352.

Marcus, E. & Ev. Marcus, 1969. Euthyneure Meeresschnecken Brasilien. (2). Beitr. Neotrop. Fauna 6(1): 1-15.

Marcus, E. & Ev. Marcus, 1969. Opisthobranchian and lamellarian Gastropoda collected by the ·Vema'. Am. Mus. Novit. (2368): 1-33.

Marcus, E. & Ev. Marcus, 1970. Opisthobranch mollusks from the southern tropical Pacific. Pac. Sci. 24: 155-179.

Marcus, E. & Ev. Marcus, 1970. Opisthobranchs from Cura~ao and faunistic ally related regions. Stud. Fauna Cura~ao Other Caribb. lsI. 33: 1-129.

Marcus, E. & Ev. Marcus, 1970. Some gastropods from Madagascar and West Mexico. Malacologia 10(1): 181-223.

Marcus, Ev., 1970. Opisthobranchs from northern Brazil. Bull. mar. Sci. 20(4): 922-951.

Marcus, Ev., 1971. On some euthyneuren gastropods from the Indian and Pacific Oceans. Proc. Malac. Soc. Lond. 39: 355-369.

Marcus, Ev., 1972. On some Acteonidae (Gastropoda Opisthobranchia). Pap. Avulsos Dep. Zool. (Sao Paulo) 25(19): 167-188.

Marcus, Ev., 1972. On some opisthobranchs from Cananeia. Bol. Zool. BioI. Mar. (Nova Ser.) 27: 207-228.

Marcus, Ev., 1972. On some opisthobranchs from Florida. Bull. mar. Sci. 22(2): 284-308.

Marcus, Ev., 1972. Notes on some opisthobranch gastropods from Chesapeake Bay. Chesapeake Sci. 13(4): 300-317.

Marcus, Ev., 1972. On the Anaspidea (Gastropoda Opisthobranchia) of the warm waters of the Western Atlantic. Bull. mar. Sci. 22(4): 841-874.

Marcus, Ev., 1973. On the genus Bosellia (Mollusca Opisthobranchia). Bull. mar. Sci. 23(4): 811-823.

Marcus, Ev., 1972. Lista dos Opisthobranchia (Mollusca Gastropoda) coletados pelo Laboratorio de Ciencias do Mar, Recife, Brasil. Trab. Oceanogr. Univ. Fed. Pernambuco 13: 71-82.

Marcus, Ev., 1974. On some Cephalaspidea from the Western Atlantic. Bull. mar. Sci. 24(2): 300-371. figs. \-111.

Marcus, Ev. & Hughes, H. 1974. Opisthobranchs molluscs from Barbados. Bull. mar. Sci. 24(3): 498-532. figs. 1-56.

Marcus, Ev., 1975. On an albinistic nudibranch. Cienc. Cult. (Sao Paulo) 27: 893.

Marcus, Ev., 1975. On Kentrodoris and Jorunna (Gastropoda Opisthobranchia). Bol. Zool. (1): 11-68, 49 figs.

Marcus, Ev., 1976. A taxonomic survey of the genus Toledoni a Dall, 1902 (Opisthobranchia, Diaphanidae). Zool. Scr. 5: 25-33.

Marcus, Ev. & Gallagher, S. 1976. A new species of Dendrodoris from Florida. Veliger 18: 353-356. figs. 1-9.

Marcus, Ev., 1976. Opisthobranchia von Santa Marta, Colombia. Stud. Neotrop. Fauna 11: 119-150. figs. 1-33.

Marcus, Ev., 1976. Marine euthyneuren gastropods from Brazil (3). Stud. Neotrop. Fauna 11: 5-23. figs. 1-14.

Marcus, Ev., 1977. On the genus Tornatina and related forms. J. Molluscan Stud. Suppl. 2, 35 p. 88 figs.

Marcus, Ev., 1977. On a pleurobranchid from the Mediterranean. Veliger 19: 418-421., 1 pI. 7 figs.

Marcus, Ev., 1977. Adolf Remane, Dr. Phil, Dr. rer. nat. h.c. (lO.VLLL. 1898 - 22. XLL. 1976). Cienc. Cult. (Sao Paulo) 29(4): 481.

Edmunds, M. & Ev. Marcus. 1977. On Favorinus auritulus Marcus and Favorinus branchialis (Muller). J. Moll. Stud. 43: 200-201.

Marcus, Ev., 1977. An annotated checklist of the Western Atlantic warm water opisthobranchs. J. Moll. Stud. Suppl. 4: 1-22.

Marcus, Ev., 1978. The Western Atlantic species of Onchidella Pulmonata. Sarsia 63: 221-224, 15 figs.

Marcus, Ev., 1978. On a new species of Bosellia. Bol. Zool. 3: 1-6, 5 figs.

Marcus, Ev., 1979. The Atlantic species of Onchidella (Gastropoda Pulmonata), part. 2. Bol. Zool. 4: 1-38, 56 figs.

Marcus, Ev., 1979. Mollusca Opisthobranchia. Resultats Scientifiques des Campagnes de la Calypso, Paris, Masson. fasc. 11: 131-137, 12 figs.

Marcus, Ev., 1980. Review of Western Atlantic Elysiidae (Opisthobranchia, Ascoglossa) with a description of a new species of Elysia. Bull. mar. Sci. 30: 54-79. 60 figs.

Marcus, Ev., 1982. Systematics of the genera of the order Ascoglossa (Gastropoda). J. Molluscan Stud. Suppl. 72 figs.

Marcus, Ev., 1982. A polyclad turbellarian from oysters in the Gulf of California. Bol. Zool. 7: 171-180, 12 figs.

Marcus, Ev., 1983. The Western Atlantic Tritoniidae. Bol. Zool. 6: 177-214.80 figs.

Marcus, Ev. & T. Gosliner, 1984. A Review of the family Plerobranchaeidae (Mollusca, Opisthobranchia). Ann. S. Afr. Mus. 93: (1): 1-52, 25 figs.

Marcus, Ev., 1984. Bryozoa. Manual de Identificacao dos Invertabrados Limnicos do Brasil. 66: 1-26, 36 figs.

Marcus, Ev., 1985. The Western Atlantic warm water Notaspidea (Gastropoda, Opisthobranchia), part 2. Bol. Zool. 8: 43-76, 66 figs.

Marcus, Ev., 1985. Notes on some Brazilian Bryozoa Ectoprocta. Bol. Zool. 8: 137-143,5 figs.

Marcus, Ev., 1985. Catalogue of the Western Atlantic warm water Opisthobranchia. 10: The Western Atlantic warm water Notaspidea. part 3. Umbraculacea. Bol. Zool. 9: 1-15,25 figs.

Marcus, Ev., 1985. On the genus Austrodoris (Mollusca Opisthobranchia) and a new species. Bol. Zool. 9: 213-223, 10 figs.

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