Table Of ContentCoelenterate Biology:
Recent Research on Cnidaria and Ctenophora
Developments in Hydrobiology 66
Series editor
H. J. Dumont
Coelenterate Biology:
Recent Research on Cnidaria
and Ctenophora
Proceedings of the Fifth International Conference on
Coelenterate Biology, 1989
Edited by
R. B. Williams, P. F. S. Cornelius,
R. G. Hughes and E. A. Robson
(on behalf of theCoelenterate Society of the United Kingdoml
Reprinted from HVdrobiologia, vols 216/217 (1991)
Springer-Science+Business Media, B.V.
Library of Congress Cataloging-in-Publication Data
International Conference on Coelenterate Biology (5th 1989
University of Southamptonl
Coelenterate blo1oQY recent research on Cnidaria and Ctenophora
proceedings of the F'fth International Conference on Coelenterate
ßioloQY, 1989 I edited by R.ß. Williams ... let al.J.
p. cm. -- <Developments in hydrobiology ; 661
Conference held at the Unlversity of Southampton, July 10-14,
1989.
Reprinted from Hydrobiologia.
ISBN 978-94-010-5428-7 ISBN 978-94-011-3240-4 (eBook)
DOI 10.1007/978-94-011-3240-4
1. Coelenterata--Congresses. 2. Ctenophora--Congresses.
1. Wi 11 iams, R. ß. (Raymond B.), 1945- I!. Hydrobiologia.
III. Title. 11!. Series.
QL375.1535 1989
593' ,5--dc20
ISBN 978-94-010-5428-7
Printed on acidjree paper
All Rights Reserved
© 1991 Springer Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 1991
Softcover reprint ofthe hardcover 1st edition 1991
No part ofthe 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.
v
Fifth International Conference on Coelenterate Biology, 1989
University of Southampton, UK
10-14 July 1989
Conference President: Prof. G. O. Mackie (Victoria, B.C., Canada)
Vice-Presidents: Prof. J. Bouillon (Bruxelles, Belgium)
Prof. G. Chapman (London, UK)
Prof. A. P. M. Lockwood (Southampton, UK)
Organizing Committee (representing the Coelenterate Society of the United Kingdom): -
Chairman: Dr R. G. Hughes (School of Biological Sciences, Queen Mary & Westfield College, Mile End
Road, London El 4NS)
Vice-Chairman: Dr P. F. S. Cornelius (Dept of Zoology, British Museum (Natural History), Cromwell
Road, London SW7 5BD)
Hon. Secretary: Dr E. A. Robson (Dept of Pure & Applied Zoology, The University, P.O. Box 228,
Reading RG6 2AJ)
Hon. Treasurer: Dr M. A. Carter (School of Biological Sciences, Portsmouth Polytechnic, King Henry I
Street, Portsmouth PO 1 2DY)
Hon. Member: Dr J. A. E. B. Hubbard (Geological Studies Group, King's College London, The Strand,
London WC2R 2LS)
Hon. Chief Editor: Dr R. B. Williams (Norfolk House, Western Road, Tring, Herts. HP23 4BN)
Hon. Associate Editors: Dr P. F. S. Cornelius
Dr R. G. Hughes
Dr E. A. Robson
The Conference logo, depicting an actinula larva, was devised by Prof. Garth Chapman on the basis
of an illustration by G. J. Allman (1812-98). This planktonic form was selected by Prof. Chapman as
the symbolic unification of the two dominant cnidarian life-cycle stages of polyp and medusa.
Bibliographic references to these Proceedings should be in one of the following forms: -
1) For the whole volume:
Williams, R. B., P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds), 1991. Coelenterate Biology:
Recent Research on Cnidaria and Ctenophora. Proceedings of the Fifth International Conference on
Coelenterate Biology, 1989. Developments in Hydrobiology 66. Kluwer Academic Publishers, Dor-
drecht, xx + 742 pp.
2) For individual papers:
Holley, M. c., 1991. Problems of being a cell in a soft body. Hydrobiologia 216/217: 35-38.
VI
Hydrobiologia 216/217: vii-viii, 1991.
R. B. Williams, P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds), Vll
Coelenterate Biology: Recent Research on Cnidaria and Ctenophora.
Foreword
The Fifth International Conference on Coelenterate Biology was held at the University of Southampton
from Monday to Friday, 10-14 July 1989. Over 300 people attended altogether, of whom 250 were
participants. The presentations comprised 164 lectures, 2 films and 64 posters divided into 41 sessions,
each under the direction of a session organizer. The topics were Cellular Biology, including Nematocysts
and Skeletogenesis; Developmental Biology and Reproduction; Ecological Physiology and Symbiosis;
Ecology including Conservation and Pollution; Evolution and Systematics including Palaeontology,
Regional Ecology and Biogeography; Genetics; Physiology and Behaviour; and Historical Aspects. This
diversity contrasted with some of the previous International Conferences (p. ix), reflecting the Organizing
Committee's decision not to restrict the scope of the subject matter and recognizing the value of
interactions between specialists in different fields. On Tuesday evening informal workshops on
Coelenterate Muscles (led by Dr M.-L. Hernandez-Nicaise), Nematocysts (led by Prof. J. W. Burnett)
and Phylogeny (led by Dr B. R. Rosen and Prof. B. G. Gardiner) were held in Conn aught Hall.
The Conference was generally regarded as being particularly successful. This was undoubtedly due to
the foresight and hard work of my colleagues on the organizing Committee, the efforts of the session
organizers, the invaluable contributions of the other helpers (including Mr & Mrs D. McAllen and Miss
M. Matthews), and the professionalism of the University Conference Office. In the early stages of
planning, the Organizing Committee was chaired successively by Prof. Garth Chapman and Dr Brian
Rosen with Dr Paul Cornelius as its Hon. Secretary. For various reasons these people had to resign or
take a less active role. To all I extend my thanks. One of the reasons that Southampton was finally chosen
as the venue was that most of the participants could be accommodated in one residence. This encouraged
informal discussions and new friendships, stimulated by the great diversity of interests of the participants.
The reception of participants was slightly manic but good humoured and set the tone for the atmosphere
of relaxed friendliness which subsequently pervaded the Conference. The social programme was highly
successful. The Conference was opened by the president, Prof. George Mackie, at an informal reception
on the evening of 9 July. Next evening a wine reception and a trade exhibition, including books, journals
and scientific equipment, were held in Conn aught Hall. On Wednesday afternoon the Conference was
given a Civic Reception by the Mayor of Southampton in The Hall of Aviation Museum, which was
followed by a shipboard evening meal while cruising in perfect conditions on The Solent and River
Hamble. The Conference Banquet was on Thursday evening. The greater part of the organization
associated with the Southampton venue fell on Dr Mike Carter, acting as local secretary, who received
a standing ovation from the participants and his committee colleagues for his tireless enterprise.
The Organizing Committee is grateful to The Royal Society of London for a grant and an interest-free
loan which enabled it to begin the organization of the Conference and to assist some overseas visitors
who otherwise could not have attended. Much-appreciated support was also received from the British
Academy, the British Council, the Commonwealth Foundation, the Monsanto Company (USA), the
United Nations Environment Programme and the University of Victoria (Canada).
During the Conference the procedure for arranging the venue for the next International Coelenterate
Conference was discussed, there being no perpetual committee. It was agreed that the Organizing
Committee of this Fifth Conference should receive offers and choose the venue for the next. Hence, it
is intended that Dr S. van der Spoel (Institute of Taxonomic Zoology, PO Box 4766, 1009 AT,
Amsterdam) and his colleagues in The Netherlands will organize the Sixth International Conference, for
June 1995.
R. G. HUGHES
Chairman, Organizing Committee
Vlll
Hydrobiologia 216/217: vii-viii, 1991.
R. B. Williams, P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds),
Coelenterate Biology: Recent Research on Cnidaria and Ctenophora.
Editorial preface
Devotees of the Cnidaria and Ctenophora will all agree that it is impossible to separate aesthetic and
scientific interests in these animals. They are among the most beautiful of any, rivalling all others in their
graceful forms and often vivid colours. They are deceptively simple, often colonial invertebrates, almost
exclusively marine, and include the benthic hydroids, sea anemones, reef-building corals, the familiar
(though atypical) freshwater hydra; and the planktonic medusae, siphonophores and ctenophores (comb
jellies), as well as some larval forms of various other Cnidaria.
Coelenterates provide excellent experimental models for understanding more complicated organisms,
and in addition have considerable biological importance in their own rights. Accordingly, much research
has been concentrated on coral reef ecosystems, including the growth, calcification, reproduction and
algal symbioses of corals; on the taxonomic study of species diversity; on popUlation studies on dynamics,
social interactions and genetics in corals and anemones; on the cell differentiation, morphogenesis, life
cycles and longevity of hydroids and anemones; on the physiological analysis of behaviour; on the
biochemistry and significance of bioluminescence; on the unique cnidarian stinging system, including its
toxins and medical importance; on the effects of pelagic predators on fisheries; on the exploitation and
conservation of corals; and on biogeographical and ecological problems including the monitoring and
control of pollution.
Aspects of all these - general reviews, specific advances in research, and descriptions of new taxa -
were addressed at the Fifth International Conference on Coelenterate Biology. Ten years had elapsed
since the last international conference on coelenterates and much had been achieved in these varied and
fruitful fields of research. This volume of proceedings contains some of the fascinating and exciting
insights of researchers from many parts of the world, and like its predecessors will, it is hoped, provide
a source book of lasting value.
A total of97 texts out of the 230 presentations given are published here. The remainder, some of which
will appear elsewhere, are listed under title and author( s) with the addresses of the first authors for the
convenience of readers who may wish to follow up topics of interest. No presentations given on historical
aspects are published here because it was considered more appropriate to submit them to specialist
journals: some will appear in the Archives of Natural History. The papers and the list of remaining titles
are arranged under the broad subject headings of the Conference (p. vii). The author, taxonomic and
subject indices prepared by P. F. S. Cornelius and R. B. Williams provide a detailed guide to the contents
of the published papers and assist in finding authors and titles of the unpublished works.
I thank all the authors who offered papers of such high scientific merit for these proceedings: without
their enthusiasm, patience and understanding, this publication would not have appeared. I am grateful
also to the many referees who freely gave their time and advice. Special thanks go to my co-editors: it
should be mentioned that all the editorial work had to be done during whatever spare evenings, weekends
and holidays were available. I would also like to thank Prof. H. M. Lenhoff who organized the collation
of the historical papers. Mrs S. E. Williams made considerable contributions to administration and proof
reading. Thanks are also due to Wi! R. Peters of Kluwer Academic Publishers for his help and advice
during the production of this volume.
R. B. WILLIAMS
Chief Editor, Editorial Committee
Hydrohiologia 216/217: ix-x, 1991.
R. B. Williams. P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds), ix
Coelenterate Biology: Recent Research on Cnidaria and Ctenophora.
© 1991 Kluwer Academic Publishers.
A history of the international conferences on coelenterate biologyl
G. O. Mackie
Department of Biology, University of Victoria, Victoria, B.C., V8W 2Y2, Canada
The origin of this series of conferences, like the the meeting, and Werner volunteered to explore
evolution of the coelenterates themselves, is a the possibilities. Fortunately the Japanese coelen-
somewhat murky topic as there are several terate people were fully in accord. The resulting
meetings that could claim to have been the first. conference (The Second International Sympo-
One of these was the hydra meeting organized by sium on Cnidaria) was held at Shirahama and
W. F. Loomis and H. M. Lenhoff at Coral Kushimoto in October 1972 (Tokioka &
Gables, Florida, in March 1961 (Lenhoff & Nishimura, 1973).
Loomis, 1961) and another was the symposium The decision to hold the third meeting in
arranged by L. M. Passano on behalf of the Com- Canada was reached while the Japanese meeting
parative Physiology Division of the American was still in progress. During the last day of the
Society of Zoologists, held in Knoxville, conference, D. M. Ross issued a formal invitation
Tennessee in December 1964 (Crowell, 1965). on behalf of the Canadians present to host the
Both of these were rather specialized next meeting in British Columbia. Our invitation
gatherings, however, and it has come to be tacitly was accepted. We decided, in order to keep the
accepted that the first in the series of international meeting within manageable limits, to have a
coelenterate conferences was the one set up by specific focus rather than to invitc contributions
W. J. Rees at the Zoological Society of London in all areas, and the theme chosen was 'ecology
in March 1965, the papers from which were pub- and behaviour'. Even with this restriction, the
lished in The Cnidaria and their Evolution (Rees, conference, held in Victoria in May 1976,
1966). This conference was not, to my knowledge, attracted about 150 participants from 18
planned as the first in a series, but several people countries (Mackie, 1976).
in the field at the time felt that it would be a good It is sad to have to record that three of the
thing to have another meeting within a few years. people most influential in establishing this series
In October 1969, while attending the Primo of conferences, W. J. Rees, B. Werner and D. M.
Simposio Internacional de Zoofilogenia in Ross, have all now passed on. They have left a
Salamanca, Bernhard Werner and I met in a local legacy from which we all benefit.
tavern as a self-appointed, ad hoc committee to Fittingly for a meeting honouring Abraham
decide where the next conference should be held. Trembley, the fourth conference, held at Inter-
The minutes of this meeting have not survived, but laken in September 1979, was planned around the
I recall that Werner had friends and contacts in theme of developmental and cellular biology,
Japan, and was planning to study at the Seto stressing work on hydra. Hydra biology has
Marine Biology Laboratory during 1971. We always seemed to stand somewhat apart from the
agreed that Japan, the land of Taku Komai and mainstream of coelenterate biology. Some no
Tohru Uchida, would be the ideal place to have doubt think that it is the mainstream. Either way,
Abraham Trembley has a lot to answer for! The
Swiss meeting, despite (or because of) its concen-
[ From the author's opening address as President of the
Fifth International Conference on Coelenterate Biology, tration on hydra biology, was a substantial affair,
given at Southampton University, 9 July 1989. gathering in participants from many countries and
x
yielding a 500 page book (Tardent & Tardent, References
1980).
Now, nearly 25 years after the original London Crowell, S. (ed.), 1965. Symposium on behavioral physiology
of coelenterates. Am. Zool. 5(3): 335-589.
conference, we are back in England for the fifth
Lenhoff, H. M. & W. F. Loomis (eds), 1961. The biology of
conference in the series, with a gathering of people
hydra and of some other coelenterates: 1961. University
from 28 countries of 6 continents. Coelenterate
of Miami Press. Coral Gables. xvi + 467 pp.
biology is clearly not a declining field and there Mackie, G. O. (ed.), 1976. Coelenterate ecology and
will continue to be a need for meetings of this sort. behavior. Plenum Press, N.Y., xiv + 744 pp.
Rees, W. J. (ed.), 1966. The Cnidaria and their evolution.
I am sure I voice the feelings of all those gathered
Symp. Zool. Soc. London 16. Academic Press, Lond.,
here in Southampton, and in fact of everyone in
xviii + 449 pp.
the international community of coelenterate
Tardent, P. & R. Tardent (eds), 1980. Developmental and
workers, in thanking our British hosts for the cellular biology of coelenterates. Elsevier/North Holland
splendid job they have done in setting up the Biomedical Press, Amsterdam, xxiv + 499 pp.
Tokioka, T. & S. Nishimura (eds), 1973. Recent trends in
present meeting and arranging for the publication
research in coelenterate biology. PubIs Seto mar. bioI. Lab.
of the conference papers in an important and
20: xlii + 793 pp.
widely read scientific journal, Hydrobiologia.
