Coelenterate 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.

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): -

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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, Dor­drecht, 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.

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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.

Foreword

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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

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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), Coelenterate Biology: Recent Research on Cnidaria and Ctenophora.

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© 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 evolution of the coelenterates themselves, is a somewhat murky topic as there are several meetings that could claim to have been the first. One of these was the hydra meeting organized by W. F. Loomis and H. M. Lenhoff at Coral Gables, Florida, in March 1961 (Lenhoff & Loomis, 1961) and another was the symposium arranged by L. M. Passano on behalf of the Com­parative Physiology Division of the American Society of Zoologists, held in Knoxville, Tennessee in December 1964 (Crowell, 1965).

Both of these were rather specialized gatherings, however, and it has come to be tacitly accepted that the first in the series of international coelenterate conferences was the one set up by W. J. Rees at the Zoological Society of London in March 1965, the papers from which were pub­lished in The Cnidaria and their Evolution (Rees, 1966). This conference was not, to my knowledge, planned as the first in a series, but several people in the field at the time felt that it would be a good thing to have another meeting within a few years. In October 1969, while attending the Primo Simposio Internacional de Zoofilogenia in Salamanca, Bernhard Werner and I met in a local tavern as a self-appointed, ad hoc committee to decide where the next conference should be held. The minutes of this meeting have not survived, but I recall that Werner had friends and contacts in Japan, and was planning to study at the Seto Marine Biology Laboratory during 1971. We agreed that Japan, the land of Taku Komai and Tohru Uchida, would be the ideal place to have

[ From the author's opening address as President of the Fifth International Conference on Coelenterate Biology, given at Southampton University, 9 July 1989.

the meeting, and Werner volunteered to explore the possibilities. Fortunately the Japanese coelen­terate people were fully in accord. The resulting conference (The Second International Sympo­sium on Cnidaria) was held at Shirahama and Kushimoto in October 1972 (Tokioka & Nishimura, 1973).

The decision to hold the third meeting in Canada was reached while the Japanese meeting was still in progress. During the last day of the conference, D. M. Ross issued a formal invitation on behalf of the Canadians present to host the next meeting in British Columbia. Our invitation was accepted. We decided, in order to keep the meeting within manageable limits, to have a specific focus rather than to invitc contributions in all areas, and the theme chosen was 'ecology and behaviour'. Even with this restriction, the conference, held in Victoria in May 1976, attracted about 150 participants from 18 countries (Mackie, 1976).

It is sad to have to record that three of the people most influential in establishing this series of conferences, W. J. Rees, B. Werner and D. M. Ross, have all now passed on. They have left a legacy from which we all benefit.

Fittingly for a meeting honouring Abraham Trembley, the fourth conference, held at Inter­laken in September 1979, was planned around the theme of developmental and cellular biology, stressing work on hydra. Hydra biology has always seemed to stand somewhat apart from the mainstream of coelenterate biology. Some no 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 concen­tration on hydra biology, was a substantial affair, gathering in participants from many countries and

x

yielding a 500 page book (Tardent & Tardent, 1980).

Now, nearly 25 years after the original London conference, we are back in England for the fifth conference in the series, with a gathering of people from 28 countries of 6 continents. Coelenterate biology is clearly not a declining field and there will continue to be a need for meetings of this sort. I am sure I voice the feelings of all those gathered here in Southampton, and in fact of everyone in the international community of coelenterate workers, in thanking our British hosts for the splendid job they have done in setting up the present meeting and arranging for the publication of the conference papers in an important and widely read scientific journal, Hydrobiologia.

References

Crowell, S. (ed.), 1965. Symposium on behavioral physiology of coelenterates. Am. Zool. 5(3): 335-589.

Lenhoff, H. M. & W. F. Loomis (eds), 1961. The biology of hydra and of some other coelenterates: 1961. University of Miami Press. Coral Gables. xvi + 467 pp.

Mackie, G. O. (ed.), 1976. Coelenterate ecology and behavior. Plenum Press, N.Y., xiv + 744 pp.

Rees, W. J. (ed.), 1966. The Cnidaria and their evolution. Symp. Zool. Soc. London 16. Academic Press, Lond., xviii + 449 pp.

Tardent, P. & R. Tardent (eds), 1980. Developmental and cellular biology of coelenterates. Elsevier/North Holland Biomedical Press, Amsterdam, xxiv + 499 pp.

Tokioka, T. & S. Nishimura (eds), 1973. Recent trends in research in coelenterate biology. PubIs Seto mar. bioI. Lab. 20: xlii + 793 pp.

Hydrobiologia 216/217: xi-xiv, 1991. R. B. Williams, P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds), Coelenterate Biology: Recent Research on Cnidaria and Ctenophora.

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© 1991 Kluwer Academic Publishers.

A selected bibliography of collective works concerning cnidarians and ctenophores (1961-1990)

R. B. Williams Noifolk House, Western Road, Tring, Hertfordshire, HP23 4BN, UK

Introduction

During the second half of the twentieth century we have seen the rapid development of a distinc­tive type of scientific publication, the 'collective work', comprising the proceedings of confer­ences, Festschriften and various other forms of 'multi-author' books with chapters on different aspects of broad areas of research; and usually edited by one or more workers in the field. They have special value in research, providing at once useful sources of specific information, subject reviews, general ideas, and bibliographic refer­ences; and so provide an ideal starting point for a search into background information for a new project. It is hoped that the present volume will similarly serve such purposes for workers on cnidarians and ctenophores, and its usefulness may perhaps be enhanced by including a selected bibliography of earlier collective works in the same field.

The selection covers an arbitrary period of 30 years, which takes in the proceedings of the first four international conferences on coelenterate biology [CB 1-4], the six international coral reef symposia [CR 1-6] and the five international sym­posia on fossil Cnidaria [FC 1-5]. The volumes included may be devoted completely to coelen­terates or they may be concerned with broader fields, such as aspects of ecology, physiology, be­haviour, experimental techniques, etc. In the main, their titles adequately reflect their contents and so it has not been considered necessary to classify the entries. Books included from the broader fields have been chosen on a subjective basis depending on my judgement of the interest

and importance to coelenterologists of only part of their contents, sometimes maybe just a single chapter. Hence, this list cannot be exhaustive nor, because of restricted space, is it possible to provide annotations. I must leave the interested reader to follow up references in order to assess their usefulness. This exercise might be made more profitable by the likelihood of picking up cross-fertilizing ideas and references from the more general works.

In the interests of bibliographical accuracy, multiple volumes of conference proceedings or other works are listed separately if the editor(s) and/or dates of the volumes are different. This also facilitates their retrieval from library cata­logues with name rather than title indices. Only the volumes of particular interest are listed of an annual series or a multivolume treatise. Encyclo­paedias are not included. On the rare occasions when the editor of a volume was not ascertain­able, it is listed under a corporate authorship or 'Anon.' as appropriate.

I thank Drs B. R. Rosen, P. F. S. Cornelius and E. A. Robson for helpful comments and some additional titles.

Bibliography

Academia Brasileira de Ciencias, 1981. Seminarios de biologia marinha. Rio de Janeiro, vi + 383 pp.

Adiyodi, K. G. & R. G. Adiyodi (eds), 1983·1989. Reproduc­tive biology of invertebrates. John Wiley & Sons, Chichester, voL 1, xxvi + 770 pp.; vol. 2, xxvi + 692 pp.; vol. 4A, xxiv + 463 pp.

Ali, M. A. (ed.), 1986. Nervous systems in invertebrates. Plenum Press, N.Y., viii + 675 pp.

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Anderson, P. A. V. (ed.), 1989. Evolution of the first nervous systems. Plenum Press, N.Y., xxiii + 423 pp.

Anon., 1974. Studies of venomous medusa Gonionemus ver­tens vertens. USSR Academy of Sciences, Vladivostok, 68 pp. [in Russian].

Anon., 1975. Marine flowers. Matsushita Electric Industrial Co. Ltd, Okinawa, 112 pp. [in Japanese and English].

Anon., 1977. Second symposium international sur les coraux et recifs coralliens fossiles. Memoires du Bureau de Re­cherches geologiques et minieres 89: i-xvi, 1-542. [FC 2]

Baker, J. T., R. M. Carter, P. W. Sammarco & K. P. Stark (eds), 1983. Proceedings of the inaugural Great Barrier Reef Conference. JCU Press, Townsville, xx + 545 pp.

Barnes, D. J. (ed.), 1983. Perspectives on coral reefs. Brian Clouston Publisher, Manuka, x + 277 pp.

Barnes, H. (ed.), 1967. Oceanography and marine biology. George Allen & Unwin Ltd, London, vol. 5, 653 pp.

Barnes, H. (ed.), 1969. Oceanography and marine biology. George Allen & Unwin Ltd, London, vol. 6, 563 pp.

Bayer, F. M. & A. J. Weinheimer (eds), 1974. Prostaglandins from Plexaura homomalla: ecology, utilization and conser­vation of a major medical marine resource. Stud. trop. Oceanogr. 12: xii + 165 pp.

Blaxter, J. H. S. & A. J. Southward (eds), 1987. Advances in marine biology. Academic Press, London, vol. 24, xii + 473 pp.

Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1982. Ad­vances in marine biology. Academic Press, London, vol. 19, xii + 381 pp.

Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1984. Ad­vances in marine biology. Academic Press, London, vol. 21, xii + 233 pp.

Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1985. Ad­vances in marine biology. Academic Press, London, vol. 22, xii + 259 pp.

Boardman, R. S., A. H. Cheetham & W. A. Oliver (eds), 1973. Animal colonies: development and function through time. Dowden, Hutchinson & Ross, Inc., Stroudsburg, xiv + 603 pp.

Bouillon, J., F. Boero, F. Cicogna & P. F. S. Cornelius (cds), 1987. Modern trends in the systematics, ecology, and evo­lution of hydroids and hydromedusae. Clarendon Press, Oxford, xxii + 328 pp.

Burnett, A. L. (ed.), 1973. Biology of Hydra. Academic Press, N.Y., xvi + 466 pp.

Cameron, A. M., B. M. Campbell, A. B. Cribb, R. Endean, J. S. Jell, O. A. Jones, P. Mather & F. H. Talbot (eds), 1974. Proceedings of the second international symposium on coral reefs. Great Barrier Reef Committee, Brisbane, vol. 1, x + 630 pp.; vol. 2, vi + 753 pp. [CR 2)

Cheng, T. C (ed.), 1971. Aspects of the biology of symbiosis. University Park Press, Baltimore, x + 327 pp.

Chia, F.-S. & M. E. Rice (eds), 1978. Settlement and meta­morphosis of marine invertebrate larvae. Elsevier, N.Y., xii + 290 pp.

Choat, J. H., D. Barnes, M. A. Borowitzka, J. C. Call, P. J. Davies, P. Flood, B. G. Hatcher, D. Hopley, P. A. Hutchings, D. Kinsey, G. R. Orme, M. Pichon, P. F. Sale, P. Sammarco, C. C. Wallace, C Wilkinson, E. Wolanski & O. Bellwood (eds), 1988. Proceedings of the sixth inter­national coral reef symposium. Townsville, vol. 1, viii + 285 pp.; vol. 2, xvi + 867 pp.; vol. 3, xiv + 584 pp.

[CR6] Corning, W. C, J. A. Dyal & A. O. D. Willows (eds), 1973.

Invertebrate learning. I. Protozoans through annelids. Plenum Press, N.Y., xviii + 296 pp.

Crowell, S. (ed.), 1965. Symposium on behavioral physiology of coelenterates. Am. Zoo!. 5(3): 335-589.

Delesalle, B., R. Galzin & B. Salvat (eds), 1985. Proceedings of the fifth international coral reef congress. Antenne Museum-EPHE, Moorea, vol. 1, x + 554 pp. [CR 5]

Dougherty, E. C., Z. N. Brown, E. D. Hanson & W. D. Hartman (eds), 1963. The lower Metazoa. Comparative biology and phylogeny. Univ. of California Press, Berkeley & Los Angeles, xii + 478 pp.

Doumenc, D. (ed.), 1987. Traite de zoologie, anatomie, syste­matique, biologie. Tome III, Fasc. 3, Cnidaires, Antho­zoaires. Masson, Paris, ii + 859 pp.

Dubinsky, Z. (ed.), 1990. Coral reefs. Elsevier, Amsterdam, xii + 550 pp.

Florkin, M. & B. T. Scheer (eds), 1968. Chemical Zoology. II. Porifera, Coelenterata and Platyhelminthes. Academic Press, N.Y., xx + 639 pp.

Gabrie, C & V. M. Harmelin (eds), 1985. Proceedings of the fifth international coral reef congress. Antenne Museum­EPHE, Moorea, vol. 6, xvi + 671 pp. [CR 5]

Gabrie, C & B. Salvat (eds), 1985. Proceedings of the fifth international coral reef congress. Antenne Museum­EPHE, Moorea, vol. 4, xiv + 583 pp. [CR 5]

Gabrie, C, J. L. Toffart & B. Salvat (eds), 1985. Proceedings of the fifth international coral reef congress. Antenne Museum-EPHE, Moorea, vol. 2, xliii + 428 pp. [CR 5]

Gabrie, C, J. L. Toffart & B. Salvat (eds), 1985. Proceedings of the fifth international coral reef congress. Antenne Museum-EPHE, Moorea, vo!' 3, xiv + 517 pp. [CR 5]

Giese, A. C. & J. S. Pearse (eds), 1974. Reproduction of marine invertebrates. I. Acoelomate and pseudocoelomate metazoans. Academic Press, N.Y., xii + 546 pp.

Gomez, E. D., C. E. Birkeland, R. W. Buddemeier, R. E. Johannes, J. A. Marsh & R. T. Tsuda (eds), 1981. The reef and man. Proceedings of the fourth international coral reef symposium. Marine Sciences Center, University of the Philippines, vol. 1, xii + 725 pp.; vol. 2, xii + 785 pp.

[CR4] Grant, P. T. & A. M. Mackie (cds), 1974. Chemoreception in

marine organisms. Academic Press, London, xii + 295 pp. Harmelin, V. M. & B. Salvat (eds), 1985. Proceedings of the

fifth international coral reef congress. Antenne Museum­EPHE, Moorea, vol. 5, xvi + 616 pp. [CR 5]

Harper, J. L., B. R. Rosen & 1. White (eds), 1986. The growth

and form of modular organisms. The Royal Society, London, iv + 250 pp.

Hayward, P. J. & J. S. Ryland (eds), 1990. The marine fauna of the British Isles and north-west Europe. Vol. 1, Intro­duction and protozoans to arthropods. Clarendon Press, Oxford, xvi + 628 + 44 pp.

Hessinger, D. A. & H. M. Lenhoff (eds), 1988. The biology ofnematocysts. Academic Press, San Diego, xii + 600 pp.

House, M. R. (ed.), 1979. The origin of major invertebrate groups. Academic Press, London, x + 515 pp.

Institut Oceanographique, 1985. Apport des coe1enteres ala biologie marine. Oceanis 11: 309-364.

Jackson, 1. B. C, L. W. Buss & R. E. Cook (eds), 1985. Popu­lation biology and evolution of clonal organisms. Yale University Press, New Haven, xiv + 530 pp.

Jell, P. A. & J. W. Pickett (eds), 1989. Fossil Cnidaria 5. Proceedings of the fifth international symposium on fossil Cnidaria including Archaeocyatha and spongiomorphs. Association of Australasian Palaeontologists, Brisbane, vi + 438 pp. [FC 5]

Jones, o. A. & R. Endean (eds), 1973-1977. Biology and geology of coral reefs. Vol. I, xviii + 410 pp.; vol. 2, xxii + 480 pp.; vol. 3, xxii + 435 pp.; vol. 4, xx + 337 pp. Academic Press, N.Y.

Keegan, B. F., 6 Ceidigh, P. & P. J. s. Boaden (eds), 1977. Biology of benthic organisms. Pergamon Press, Oxford, xxxiv + 630 pp.

Kinne, O. (ed.), 1970-1984. Marine ecology. A comprehen­sive, integrated treatise on life in oceans and coastal waters. I. Environmental factors (3 parts); II. Physiologi­cal mechanisms (2 parts); III. Cultivation (3 parts); IV. Dynamics (1 part); V. Ocean management (4 parts). John Wiley & Sons Ltd, London.

Koltun, V. M. & S. D. Stepanjants (eds), 1988. Porifera and Cnidaria. Modern and perspective investigations. USSR Academy of Sciences, Leningrad, 176 pp. [in Russian and English].

Larwood, G. & B. R. Rosen (eds), 1979. Biology and syste­matics of colonial organisms. Academic Press, London, xxxvi + 589 pp.

Lenhoff, H. M. (ed.), 1983. Hydra: research methods. Plenum Press, N.Y., xxxii + 463 pp.

Lenhoff, H. M. & W. F. Loomis (eds), 1961. The biology of hydra and of some other coelenterates: 1961. Univ. of Miami Press, Coral Gables, xvi + 467 pp.

Lenhoff, H. M. & P. Tardent (eds), 1985. From Trembley's polyps to new directions in research on Hydra: pro­ceedings of a symposium honoring Abraham Trembley (1710-1784). Societe de Physique et d'Histoire N aturelle, Geneva, vi + 229 pp. [Reprinted from Archs Sc. Geneve 38: 243-471.]

Lenhoff, H. M., L. Muscatine & L. V. Davis (eds), 1971. Experimental coelenterate biology. Univ. of Hawaii Press, Honolulu, x + 281 pp.

Mackie, G. o. (ed.), 1976. Coelenterate ecology and be-havior. Plenum Press, N.Y., 744 pp. [CB 3]

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Mariscal, R. N. (ed.), 1974. Experimental marine biology. Academic Press, N.Y., x + 373 pp.

Morris, S. C, 1. D. George, R. Gibson & H. M. Platt (eds), 1985. The origins and relationships oflower invertebrates. Clarendon Press, Oxford, xii + 397 pp.

Mukundan, C. & C S. G. Pillai (eds), 1972. Proceedings of the symposium on corals and coral reefs. Marine Biologi­cal Association of India, Cochin, xxii + 591 pp. [CR I]

Muscatine, L. & H. M. Lenhoff (eds), 1974. Coelenterate biology. Reviews and new perspectives. Academic Press, N.Y., x + SOl pp.

Naumov, D. V. & S. D. Stepanjants (eds), 1980. The theoretical and practical importance of the coelenterates. USSR Academy of Sciences, Leningrad, 148 pp. [in Russian].

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Contents

Fifth International Conference on Coelenterate Biology .......................................... v Some Conference participants (group photograph) .............................................. vi Foreword ................................................................................. vii Editorial preface ........................................................................... viii A history of the international conferences on coelenterate biology

by G. O. Mackie .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix A selected bibliography of collective works concerning cnidarians and ctenophores (1961-1990)

by R. B. Williams ........................................................................ xi

PROCEEDINGS

I. CELLULAR BIOLOGY The extracellular matrix (mesoglea) of hydrozoan jellyfish and its ability to support cell adhesion and spreading

by V. Schmid, A. Bally, K. Beck, M. Haller, W. K Schlage & C. Weber ....... '" . . . . . . . 3 Effects of tumor promoters and diacylglycerol on the transdifferentiation of striated muscle cells of the medusa Podocoryne carnea to RF -amide positive nerve cells

by E. Kurz & V. Schmid. . . . . . . . . .. .. . . . . . . . . . . . . . . . . .... . . . . . . . . . . . . ... . . .. . . . . . 7 The physiology of feeding in the ctenophore Pleurobrachia pileus

by A. G. Moss. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Behavioural and ultrastructural studies on the sedentary platyctenean ctenophore Vallicula multtfor­mis

by R. H. Emson & P. J. Whitfield ................................................. 27 Problems of being a cell in a soft body

by M. C. Holley. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35 An ultrastructural study of the polyp and strobila of Atorella japonica (Cnidaria, Coronatae) with respect to muscles and nerves

by A. Matsuno & S. Kawaguti .................................................... 39 Rhopalium development in Aurelia au rita ephyrae

by D. B. Spangenberg. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

II. DEVELOPMENTAL BIOLOGY Bud formation in the scyphozoan Cassiopea andromeda: epithelial dynamics and fate map

by D. K. Hofmann & M. Gottlieb ................................................. 53 Basic morphogenetic processes in Hydrozoa and their evolutionary implications: an exercise in rational taxonomy

by L. V. Beloussov .............................................................. 61 Regression and replacement of hydranths in thecate hydroids, and the structure of hydrothecae

by S. Crowell .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Differentiation of the interstitial cell line in hydrozoan planulae. I. RepopUlation of epithelial planulae

by V. J. Martin ................................................................. 75 A low-molecular weight factor from colonial hydroids affects body proportioning and cell differen­tiation

by G. Plickert .................................................................. 83

XVI

Polyp regeneration from isolated tentacles of Aurelia scyphistomae: a role for gating mechanisms and cell division

by G. E. Lesh-Laurie, A. Hujer & P. Suchy ......................................... 91

III. REPRODUCTION Corals and sea anemones Larval development and survivorship in the corals Favia favus and Platygyra lamellina

by y. Shlesinger & Y. Loya . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 101 Spawning of hermatypic corals in Bermuda: a pilot study

by S. C. Wyers, H. S. Barnes & S. R. Smith ........................................ 109 Annual cycle of gametogenesis and spawning in a tropical zoanthid, Protopalythoa sp.

by J. S. Ryland & R. C. Babcock .................................................. 117 Reproduction and developmental pathways of Red Sea Xeniidae (Octocorallia, Alcyonacea)

by Y. Benayahu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 125 Sperm morphology of scleractinians from the Caribbean

by S. C. C. Steiner .............................................................. 131 Embryonic and larval development of the sea anemone Haliplanella lineata from Japan

by Y. Fukui .................................................................... 137 Developmental pathways of anthozoans

by D. G. Fautin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 143

Life cycles The life cycle of Laodicea indica (Laodiceidae, Leptomedusae, Cnidaria)

by J. Bouillon, F. Boero & S. Fraschetti ............................................ 151 Notes on the morphology, ecology and life cyles of Fukaurahydra anthoformis and Hataia parva (Hydrozoa, Athecata)

by M. Yamada & S. Kubota ...................................................... 159 The ampullae and medusae of the calcareous hydrozoan Millepora complanata

by J. B. Lewis .................................................................. 165 Observations on the anatomy, behaviour, reproduction and life cycle of the cubozoan Carybdea sivickisi

by R. F. Hartwick. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 171 Distributional ecology and behaviour of the early life stages of the box-jellyfish Chironex fleckeri

by R. F. Hartwick. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 181 Oocyte development in four species of scyphomedusa in the northern Adriatic Sea

by M. Avian & L. Rottini Sandrini ................................................ 189 Reproduction of Pelagia noctiluca in the central and northern Adriatic Sea

by L. Rottini Sandrini & M. Avian ................................................ 197

IV. ECOLOGY Hydroids The growth patterns of some hydroids that are obligate epiphytes of seagrass leaves

by R. G. Hughes, S. Johnson & I. D. Smith. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 205 The growth and degeneration of the hydroid Sertularia perpusilla, an obligate epiphyte of leaves of the seagrass Posidonia oceanica

by R. G. Hughes, A. Garcia Rubies & J.-M. Gili . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 211

The settlement and growth of Sertularia cupressina (Hydrozoa, Sertulariidae) in Langstone Harbour, Hampshire, UK

XVll

by G. H. Schmidt & G. F. Warner ................................................. 215 Abundance and distribution of hydroids in a mangrove ecosystem at Twin Cays, Belize, Central America

by D. R. Calder ................................................................. 221 The adaptive pattern of growth and reproduction of the colonial hydroid Clavopsella michaeli

by S. Piraino ................................................................... 229

Zoanthids, corals and sea anemones Fission and the dynamics of genets and ramets in clonal cnidarian populations

by R. H. Karlson ................................................................ 235 Distribution and adaptive strategies of alcyonacean corals in Nanwan Bay, Taiwan

by C.-F. Dai ................................................................... 241 Effects of water movement on prey capture and distribution of reef corals [synopsis only]

by K. P. Sebens & A. S. Johnson .................................................. 247 How many damaged corals in Red Sea reef systems? A quantitative survey

by B. Riegl & B. Velimirov ....................................................... 249 The effects of Acanthaster planci predation on populations of two species of massive coral

by A. M. Cameron, R. Endean & L. M. DeVantier ................................... 257 Locomotion, asexual reproduction, and killing of corals by the corallimorpharian Corynactis califor-nica

by N. E. Chadwick & c. Adams .................................................. 263 Comparative ecology of subtidal actiniarians from the coasts of California and the Gulf of Maine, USA

by L. G. Harris 271

Associations Coral-crab association: a compact domain of a multilevel trophic system

by B. Rinkevich, Z. Wolodarsky & Y. Loya ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 279 The symbiotic relationship between the hydrocoral Millepora dichotoma and the barnacle Savignium milleporum

by P. A. Cook, B. A. Stewart & Y. Achituv ......................................... 285 Chemical recognition by hermit crabs of their symbiotic sea anemones and a predatory octopus

by W. R. Brooks ................................................................ 291 Mutualism between the antipatharian Antipathes jiordensis and the ophiuroid Astrobrachion constric­tum in New Zealand fiords

by K. R. Grange ................................................................ 297 Predation on Cnidaria by vertebrates other than fishes

by R. M. L. Ates ................................................................ 305

V. PELAGIC COELENTERATES Western Atlantic midwater hydrozoan and scyphozoan medusae: in situ studies using manned submersibles

by R. J. Larson, C. E. Mills & G. R. Harbison ...................................... 311 Swimming movements of ctenophores, and the mechanics of propulsion by ctene rows

by G. T. Matsumoto ............................................................. 319

xviii

Co-occurrence of hippopodiid siphonophores and their potential prey by P. R. Pugh .................................................................. 327

A review of cnidarians and ctenophores feeding on competitors in the plankton by J. E. Purcell ................................................................. 335

The role of prey stratification in the predation pressure by the cydippid ctenophore Mertensia ovum in the Barents Sea

by N. Swanberg & U. Bfunstedt ................................................... 343 Hydromedusae of the Mediterranean Sea

by J. Goy ..................................................................... , 351 Vertical distribution of epipelagic siphonophores at the confluence between Benguela waters and the Angola Current over 48 hours

by F. Pages & J.-M. Gili ......................................................... 355 Attraction of Aurelia and Aequorea to prey

by M. Needler Arai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 363

VI. ENDOSYMBIOSIS Amino acids as a nitrogen source for Chlorella symbiotic with green hydra

by P. J. McAuley ................................................................ 369 A review of some physiological and evolutionary aspects of body size and bud size of Hydra

by L. B. Slobodkin, P. Bossert, C. Matessi & M. Gatto. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 377 The stable level of coral primary production in a wide light range

by E. A. Titlyanov ............................................................... 383 Mechanisms of amplification of photosynthetically active radiation in the symbiotic deep-water coral Leptoseris fragi/is

by D. Schlichter & H. W. Fricke .................................................. 389

VII. SKELETOGENESIS The nature of the skeleton and skeletogenic tissues in the Cnidaria

by M. D'A. A. Le Tissier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 397 Chemistry and structure of skeletal growth rings in the black coral Antipathes fiordensis (Cnidaria, Antipatharia)

by W. M. Goldberg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 403

VIII. PALAEONTOLOGY Principles of early ontogeny in the rugose corals: a critical review

by J. Fedorowski ................................................................ 413 The Bainbridgia-Dualipora association (Cnidaria, Tabulata): palaeogeographical and palaeo­ecological implications

by F. Tourneur ................................................................. 419

IX. EVOLUTION AND SYSTEMATICS Crossing-experiments between Japanese populations of three hydrozoans symbiotic with bivalves

by S. Kubota ................................................................... 429 The variation of Halocordyle distich a (Cnidaria, Athecata) from the Brazilian coast: an environmen­tal indicator species?

by F. L. da Silveira & A. E. Migotto ............................................... 437 Samuraia tabularasa gen. nov., sp. nov. (Cnidaria, Hydrozoa, Hydrocorynidae), an intertidal hydroid from the Gulf of California, Mexico

by K. L. Mangin ................................................................ 443

XIX

A jellyfish-eating sea anemone (Cnidaria, Actiniaria) from Palau: Entacmaea medusivora sp. nov. by D. G. Fautin & W. K. Fitt ...................................... ,.............. 453

Taxonomic characters from the polyp tubes of coronate medusae (Scyphozoa, Coronatae) by G. Jarms .......................................................... " ........ 463

Morphological differences between, and reproductive isolation of, two populations of the jellyfish Cyanea in Long Island Sound, USA

by R. H. Brewer ................................................................ 471

X. BIOGEOGRAPHY The benthic deep-water siphonophore Rhodalia miranda and other coelenterates in the south-west Atlantic: ecological and oceanographical implication s

by K. Riemann-Zurneck .......................................................... 481 Hydromedusan distribution patterns

by S. van der Spoel ............................................................. 489 Siphonophores of upwelling areas of the Campeche Bank and the Mexican Caribbean Sea

by R. Gasca & E. Suarez ........................................................ 497

XI. GENETICS Molecular aspects Dead-end evolution of the Cnidaria as deduced from 5S ribosomal RNA sequences

by H. Hori & Y. Satow .......................................................... 505 Relationships within the family Actiniidae (Cnidaria, Actiniaria) based on molecular characters

by S. A. McCommas ............................................................ 509 The heat shock response in hydra: immunological relationship of hsp60, the major heat shock protein of Hydra vulgaris, to the ubiquitous hsp70 family

by T. C. G. Bosch & G. Praetzel .................................................. 513

Population structure and behaviour of sea anemones Effects of asexual reproduction on population structure of Sagartia elegans (Anthozoa, Actiniaria)

by P. W. Shaw .................................................................. 519 Gonad development of the sea anemone Anthopleura asiatica in clonal populations

by H. Fujii ..................................................................... 527 Experimental habituation of aggression in the sea anemone Actinia equina

by R. C. Brace & S.-J. Santer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 533 Acrorhagi, catch tentacles and sweeper tentacles: a synopsis of 'aggression' of actiniarian and scleractinian Cnidaria

by R. B. Williams ............................................................... 539

XII. PHYSIOLOGY AND BEHAVIOUR Transmitters Monoamine concentrations in Hydra magnipapillata

by N. Takeda & c. N. Svendsen .................................................. 549 Neuropeptides in coelenterates: a review

by C. J. P. Grimmelikhuijzen, D. Graff, O. Koizumi, J. A. Westfall & I. D. McFarlane .... 555 Peptides in the Hydrozoa: are they transmitters?

by A. N. Spencer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 565

Bioluminescence Observations on bioluminescence in some deep-water anthozoans

by P. J. Herring ................................................................. 573

xx

Propagation of bioluminescence in Euph.l'sa japonica hydromedusae (Tubulariidae) by G. O. Mackie ................................................................ 581

Sensory systems Interaction between the effects of light pulses of different chromatic content on Hydra attenuata and H. magnipapillata

by C. Taddei-Ferretti & A. Cotugno. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 589 The physiological control of feeding in corals: a review

by I. D. Lawn & I. D. McFarlane ................................................. 595 The senses of sea anemones: responses of the S S 1 nerve net to chemical and mechanical stimuli

by I. D. McFarlane & I. D. Lawn ................................................. 599

XIII. NEMA TOCYSTS Nematocysts of the Mediterranean hydroid Halocordyle disticha

by C. Ostman, S. Piraino & W. Kem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 607 A comparative analysis of nematocysts in Pelagia noctiluca and Rhizostoma pulmo from the North Adriatic Sea

by M. Avian, P. Del Negro & L. Rottini Sandrini .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 615 Differentiation and ultrastructure of nematocysts in Halammohydra intermedia (Hydrozoa, Cnidaria)

by C. Clausen .................................................................. 623 Clinical manifestations of jellyfish envenomation

by J. W. Burnett ................................................................ 629 Cubozoan jellyfish envenomation syndromes and their medical treatment in northern Australia

by P. J. Fenner ................................................................. 637 Envenomation by the box-jellyfish Chironex fleckeri: how nematocysts discharge

by R. Endean, J. F. Rifkin & L. Y. M. Daddow ..................................... , 641 Cnida discharge and the mechanism of venom delivery in Anemonia viridis (Cnidaria, Actiniaria)

by J. C. Thomason .............................................................. 649 Effects of controlled treatment with trypsin on the functional characteristics of isolated nematocysts of Calliactis parasitica and Aiptasia mutabilis (Cnidaria, Actiniaria)

by A. Salleo, G. La Spada, A. Brancati & P. Ciacco ................................. 655 The spatial distribution of cations in nematocytes of Hydra vulgaris

by I. Gerke, K. Zierold, J. Weber & P. Tardent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 661 Discharge of nematocysts isolated from aeolid nudibranchs

by P. G. Greenwood & L. K. Garrity .............................................. 671 The differentiation of cytoskeletal structures in nematocytes of Hydra

by U. Schertenleib & R. P. Stidwill ................................................ 679 Remodelling during the development of nematocysts in a siphonophore

by R. J. Skaer .................................................................. 685 Nematocysts of sea anemones (Actiniaria, Ceriantharia and Corallimorpharia: Cnidaria): nomen­clature

by K. W. England ............................................................... 691

Titles of presentations given during the conference but not included in the Proceedings ....... 699

Author index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 707 Taxonomic index .................................................................. 719 Subject index ..................................................................... 729