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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
by Kluwer Academic Publishers Group,
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
der Georg-August-Universitiit Berliner Str. 28 3400 Gottingen Germany
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
der Georg-August-Universitiit Berliner Str. 28 3400 Gottingen Germany
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-
cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan
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-
mentale, Infettiva e Pubblica Sezione di Biologia e Genetica Universita di Pisa Via A. Volta 4 56100 Pisa Italy
M. G. Gremigni-Querci Dipartimento di Biomedicina Speri-
mentale, Infettiva e Pubblica Sezione di Biologia e Genetica Universita di Pisa Via A. Volta 4 56100 Pisa Italy
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 &
Neurobiology Colorado State University Fort Collins, CO 80523 USA
Shoko Morita Department of Anatomy &
Neurobiology Colorado State University Fort Collins, CO 80523 USA
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-
cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan
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-
mentale, Infettiva e Pubblica Sezione di Biologia e Genetica Universita di Pisa Via A. Volta 4 56100 Pisa Italy
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-
cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan
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-
cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan
Yasuhiro Y oshizawa Department of Biological and Chemi-
cal Engineering Faculty of Engineering Gunma University Kiryu, Gunma 376 Japan
XVll
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
XV111
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
XIX
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
xx
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
xxiii
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
XXIV
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.
xxv
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.