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. --

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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 [ From the author's opening address as President of the Swiss meeting, despite (or because of) its concen• 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 conference, we are back in England for the fifth of coelenterates. Am. Zool. 5(3): 335-589. 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. I am sure I voice the feelings of all those gathered Rees, W. J. (ed.), 1966. The Cnidaria and their evolution. 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. present meeting and arranging for the publication Tokioka, T. & S. Nishimura (eds), 1973. Recent trends in 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. Hydrobiologia 216/217: xi-xiv, 1991. R. B. Williams, P. F. S. Cornelius, R. G. Hughes & E. A. Robson (eds), Xl Coelenterate Biology: Recent Research on Cnidaria and Ctenophora. © 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 and importance to coelenterologists of only part of their contents, sometimes maybe just a single During the second half of the twentieth century chapter. Hence, this list cannot be exhaustive nor, we have seen the rapid development of a distinc• because of restricted space, is it possible to tive type of scientific publication, the 'collective provide annotations. I must leave the interested work', comprising the proceedings of confer• reader to follow up references in order to assess ences, Festschriften and various other forms of their usefulness. This exercise might be made 'multi-author' books with chapters on different more profitable by the likelihood of picking up aspects of broad areas of research; and usually cross-fertilizing ideas and references from the edited by one or more workers in the field. They more general works. have special value in research, providing at once In the interests of bibliographical accuracy, useful sources of specific information, subject multiple volumes of conference proceedings or reviews, general ideas, and bibliographic refer• other works are listed separately if the editor(s) ences; and so provide an ideal starting point for and/or dates of the volumes are different. This a search into background information for a new also facilitates their retrieval from library cata• project. It is hoped that the present volume will logues with name rather than title indices. Only similarly serve such purposes for workers on the volumes of particular interest are listed of an cnidarians and ctenophores, and its usefulness annual series or a multivolume treatise. Encyclo• may perhaps be enhanced by including a selected paedias are not included. On the rare occasions bibliography of earlier collective works in the when the editor of a volume was not ascertain• same field. able, it is listed under a corporate authorship or The selection covers an arbitrary period of 30 'Anon.' as appropriate. years, which takes in the proceedings of the first I thank Drs B. R. Rosen, P. F. S. Cornelius four international conferences on coelenterate and E. A. Robson for helpful comments and biology [CB 1-4], the six international coral reef some additional titles. symposia [CR 1-6] and the five international sym• posia on fossil Cnidaria [FC 1-5]. The volumes included may be devoted completely to coelen• Bibliography terates or they may be concerned with broader fields, such as aspects of ecology, physiology, be• Academia Brasileira de Ciencias, 1981. Seminarios de haviour, experimental techniques, etc. In the biologia marinha. Rio de Janeiro, vi + 383 pp. main, their titles adequately reflect their contents Adiyodi, K. G. & R. G. Adiyodi (eds), 1983·1989. Reproduc• and so it has not been considered necessary to tive biology of invertebrates. John Wiley & Sons, Chichester, voL 1, xxvi + 770 pp.; vol. 2, xxvi + 692 pp.; classify the entries. Books included from the vol. 4A, xxiv + 463 pp. broader fields have been chosen on a subjective Ali, M. A. (ed.), 1986. Nervous systems in invertebrates. basis depending on my judgement of the interest Plenum Press, N.Y., viii + 675 pp. Xll

Anderson, P. A. V. (ed.), 1989. Evolution of the first nervous Choat, J. H., D. Barnes, M. A. Borowitzka, J. C. Call, P. J. systems. Plenum Press, N.Y., xxiii + 423 pp. Davies, P. Flood, B. G. Hatcher, D. Hopley, P. A. Anon., 1974. Studies of venomous medusa Gonionemus ver• Hutchings, D. Kinsey, G. R. Orme, M. Pichon, P. F. Sale, tens vertens. USSR Academy of Sciences, Vladivostok, P. Sammarco, C. C. Wallace, C Wilkinson, E. Wolanski & 68 pp. [in Russian]. O. Bellwood (eds), 1988. Proceedings of the sixth inter• Anon., 1975. Marine flowers. Matsushita Electric Industrial national coral reef symposium. Townsville, vol. 1, Co. Ltd, Okinawa, 112 pp. [in Japanese and English]. viii + 285 pp.; vol. 2, xvi + 867 pp.; vol. 3, xiv + 584 pp. Anon., 1977. Second symposium international sur les coraux [CR6] et recifs coralliens fossiles. Memoires du Bureau de Re• Corning, W. C, J. A. Dyal & A. O. D. Willows (eds), 1973. cherches geologiques et minieres 89: i-xvi, 1-542. [FC 2] Invertebrate learning. I. Protozoans through annelids. Baker, J. T., R. M. Carter, P. W. Sammarco & K. P. Stark Plenum Press, N.Y., xviii + 296 pp. (eds), 1983. Proceedings of the inaugural Great Barrier Crowell, S. (ed.), 1965. Symposium on behavioral physiology Reef Conference. JCU Press, Townsville, xx + 545 pp. of coelenterates. Am. Zoo!. 5(3): 335-589. Barnes, D. J. (ed.), 1983. Perspectives on coral reefs. Brian Delesalle, B., R. Galzin & B. Salvat (eds), 1985. Proceedings Clouston Publisher, Manuka, x + 277 pp. of the fifth international coral reef congress. Antenne Barnes, H. (ed.), 1967. Oceanography and marine biology. Museum-EPHE, Moorea, vol. 1, x + 554 pp. [CR 5] George Allen & Unwin Ltd, London, vol. 5, 653 pp. Dougherty, E. C., Z. N. Brown, E. D. Hanson & W. D. Barnes, H. (ed.), 1969. Oceanography and marine biology. Hartman (eds), 1963. The lower Metazoa. Comparative George Allen & Unwin Ltd, London, vol. 6, 563 pp. biology and phylogeny. Univ. of California Press, Berkeley Bayer, F. M. & A. J. Weinheimer (eds), 1974. Prostaglandins & Los Angeles, xii + 478 pp. from Plexaura homomalla: ecology, utilization and conser• Doumenc, D. (ed.), 1987. Traite de zoologie, anatomie, syste• vation of a major medical marine resource. Stud. trop. matique, biologie. Tome III, Fasc. 3, Cnidaires, Antho• Oceanogr. 12: xii + 165 pp. zoaires. Masson, Paris, ii + 859 pp. Blaxter, J. H. S. & A. J. Southward (eds), 1987. Advances in Dubinsky, Z. (ed.), 1990. Coral reefs. Elsevier, Amsterdam, marine biology. Academic Press, London, vol. 24, xii + 550 pp. xii + 473 pp. Florkin, M. & B. T. Scheer (eds), 1968. Chemical Zoology. Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1982. Ad• II. Porifera, Coelenterata and Platyhelminthes. Academic vances in marine biology. Academic Press, London, Press, N.Y., xx + 639 pp. vol. 19, xii + 381 pp. Gabrie, C & V. M. Harmelin (eds), 1985. Proceedings of the Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1984. Ad• fifth international coral reef congress. Antenne Museum• vances in marine biology. Academic Press, London, EPHE, Moorea, vol. 6, xvi + 671 pp. [CR 5] vol. 21, xii + 233 pp. Gabrie, C & B. Salvat (eds), 1985. Proceedings of the fifth Blaxter, J. H. S., F. S. Russell & M. Yonge (eds), 1985. Ad• international coral reef congress. Antenne Museum• vances in marine biology. Academic Press, London, EPHE, Moorea, vol. 4, xiv + 583 pp. [CR 5] vol. 22, xii + 259 pp. Gabrie, C, J. L. Toffart & B. Salvat (eds), 1985. Proceedings Boardman, R. S., A. H. Cheetham & W. A. Oliver (eds), of the fifth international coral reef congress. Antenne 1973. Animal colonies: development and function through Museum-EPHE, Moorea, vol. 2, xliii + 428 pp. [CR 5] time. Dowden, Hutchinson & Ross, Inc., Stroudsburg, Gabrie, C, J. L. Toffart & B. Salvat (eds), 1985. Proceedings xiv + 603 pp. of the fifth international coral reef congress. Antenne Bouillon, J., F. Boero, F. Cicogna & P. F. S. Cornelius (cds), Museum-EPHE, Moorea, vo!' 3, xiv + 517 pp. [CR 5] 1987. Modern trends in the systematics, ecology, and evo• Giese, A. C. & J. S. Pearse (eds), 1974. Reproduction of lution of hydroids and hydromedusae. Clarendon Press, marine invertebrates. I. Acoelomate and pseudocoelomate Oxford, xxii + 328 pp. metazoans. Academic Press, N.Y., xii + 546 pp. Burnett, A. L. (ed.), 1973. Biology of Hydra. Academic Press, Gomez, E. D., C. E. Birkeland, R. W. Buddemeier, R. E. N.Y., xvi + 466 pp. Johannes, J. A. Marsh & R. T. Tsuda (eds), 1981. The reef Cameron, A. M., B. M. Campbell, A. B. Cribb, R. Endean, and man. Proceedings of the fourth international coral reef J. S. Jell, O. A. Jones, P. Mather & F. H. Talbot (eds), symposium. Marine Sciences Center, University of the 1974. Proceedings of the second international symposium Philippines, vol. 1, xii + 725 pp.; vol. 2, xii + 785 pp. on coral reefs. Great Barrier Reef Committee, Brisbane, [CR4] vol. 1, x + 630 pp.; vol. 2, vi + 753 pp. [CR 2) Grant, P. T. & A. M. Mackie (cds), 1974. Chemoreception in Cheng, T. C (ed.), 1971. Aspects of the biology of symbiosis. marine organisms. Academic Press, London, xii + 295 pp. University Park Press, Baltimore, x + 327 pp. Harmelin, V. M. & B. Salvat (eds), 1985. Proceedings of the Chia, F.-S. & M. E. Rice (eds), 1978. Settlement and meta• fifth international coral reef congress. Antenne Museum• morphosis of marine invertebrate larvae. Elsevier, N.Y., EPHE, Moorea, vol. 5, xvi + 616 pp. [CR 5] xii + 290 pp. Harper, J. L., B. R. Rosen & 1. White (eds), 1986. The growth Xlll

and form of modular organisms. The Royal Society, Mariscal, R. N. (ed.), 1974. Experimental marine biology. London, iv + 250 pp. Academic Press, N.Y., x + 373 pp. Hayward, P. J. & J. S. Ryland (eds), 1990. The marine fauna Morris, S. C, 1. D. George, R. Gibson & H. M. Platt (eds), of the British Isles and north-west Europe. Vol. 1, Intro• 1985. The origins and relationships oflower invertebrates. duction and protozoans to arthropods. Clarendon Press, Clarendon Press, Oxford, xii + 397 pp. Oxford, xvi + 628 + 44 pp. Mukundan, C. & C S. G. Pillai (eds), 1972. Proceedings of Hessinger, D. A. & H. M. Lenhoff (eds), 1988. The biology the symposium on corals and coral reefs. Marine Biologi• ofnematocysts. Academic Press, San Diego, xii + 600 pp. cal Association of India, Cochin, xxii + 591 pp. [CR I] House, M. R. (ed.), 1979. The origin of major invertebrate Muscatine, L. & H. M. Lenhoff (eds), 1974. Coelenterate groups. Academic Press, London, x + 515 pp. biology. Reviews and new perspectives. Academic Press, Institut Oceanographique, 1985. Apport des coe1enteres ala N.Y., x + SOl pp. biologie marine. Oceanis 11: 309-364. Naumov, D. V. & S. D. Stepanjants (eds), 1980. The Jackson, 1. B. C, L. W. Buss & R. E. Cook (eds), 1985. Popu• theoretical and practical importance of the coelenterates. lation biology and evolution of clonal organisms. Yale USSR Academy of Sciences, Leningrad, 148 pp. [in University Press, New Haven, xiv + 530 pp. Russian]. Jell, P. A. & J. W. Pickett (eds), 1989. Fossil Cnidaria 5. Oliver, W. A., W. J. Sando, S. D. Cairns, A. G. Coates, 1. G. Proceedings of the fifth international symposium on fossil Macintyre, F. M. Bayer & 1. E. Sorauf(eds), 1984. Recent Cnidaria including Archaeocyatha and spongiomorphs. advances in the paleobiology and geology of the Cnidaria. Association of Australasian Palaeontologists, Brisbane, Proceedings of the fourth international symposium on vi + 438 pp. [FC 5] fossil Cnidaria (and Archaeocyathids and Stromato• Jones, o. A. & R. Endean (eds), 1973-1977. Biology and poroids). Palaeontographica Americana 54: 1-557. [FC 4] geology of coral reefs. Vol. I, xviii + 410 pp.; vol. 2, Osin6lska, H. (ed.), 1980. Third international symposium on xxii + 480 pp.; vol. 3, xxii + 435 pp.; vol. 4, xx + 337 pp. fossil cnidarians. Acta Palaeontologica Polonica 25: Academic Press, N.Y. 292-679. [FC 3] Keegan, B. F., 6 Ceidigh, P. & P. J. s. Boaden (eds), 1977. Parker, S. P. (ed.), 1982. Synopsis and classification ofliving Biology of benthic organisms. Pergamon Press, Oxford, organisms. McGraw-Hill, N.Y., vol. 1, xx + 1166 pp. xxxiv + 630 pp. Rees, W. J. (ed.), 1966. The Cnidaria and their evolution. Kinne, O. (ed.), 1970-1984. Marine ecology. A comprehen• Academic Press, London, xviii + 449 pp. [CB I] sive, integrated treatise on life in oceans and coastal Reverberi, G. (ed.), 1971. Experimental embryology of waters. I. Environmental factors (3 parts); II. Physiologi• marine and fresh-water invertebrates. North-Holland, cal mechanisms (2 parts); III. Cultivation (3 parts); IV. Amsterdam, xxiv + 587 pp. Dynamics (1 part); V. Ocean management (4 parts). John Russell, F. E., H. Gonzalez, S. B. Dobson & J. A. Coats Wiley & Sons Ltd, London. (eds), 1984. Bibliography of venomous and poisonous Koltun, V. M. & S. D. Stepanjants (eds), 1988. Porifera and marine animals and their toxins. Office of Naval Research, Cnidaria. Modern and perspective investigations. USSR Los Angeles, viii + 416 pp. Academy of Sciences, Leningrad, 176 pp. [in Russian and Russell, F. S. (ed.), 1963. Advances in marine biology. English]. Academic Press, London, vol. I, xiv + 410 pp. Larwood, G. & B. R. Rosen (eds), 1979. Biology and syste• Russell, F. S. & M. Yonge (eds), 1978. Advances in marine matics of colonial organisms. Academic Press, London, biology. Academic Press, London, vol. 15, xiv + 563 pp. xxxvi + 589 pp. Riitz1er, K. & I. G. Macintyre (eds), 1982. The Atlantic Lenhoff, H. M. (ed.), 1983. Hydra: research methods. Plenum barrier reef ecosystem at Carrie Bow Cay, Belize, I: struc• Press, N.Y., xxxii + 463 pp. ture and communities. Smithsonian Institution Press, Lenhoff, H. M. & W. F. Loomis (eds), 1961. The biology of Washington, xiv + 539 pp. hydra and of some other coelenterates: 1961. Univ. of Ryland, J. S. & P. A. Tyler (eds), 1989. Reproduction, Miami Press, Coral Gables, xvi + 467 pp. genetics and distributions of marine organisms. Olsen & Lenhoff, H. M. & P. Tardent (eds), 1985. From Trembley's Olsen, Fredensborg, viii + 469 pp. polyps to new directions in research on Hydra: pro• Shelton, G. A. B. (ed.), 1982. Electrical conduction and be• ceedings of a symposium honoring Abraham Trembley haviour in 'simple' invertebrates. Oxford University Press, (1710-1784). Societe de Physique et d'Histoire N aturelle, Oxford, x + 567 pp. Geneva, vi + 229 pp. [Reprinted from Archs Sc. Geneve Sheppard, C & S. M. Wells (eds), 1988. Coral reefs of the 38: 243-471.] world. Vol. 2, I + 389 pp. United Nations Environment Lenhoff, H. M., L. Muscatine & L. V. Davis (eds), 1971. Programme, Gland. Experimental coelenterate biology. Univ. of Hawaii Press, Sims, R. W. (ed.), 1980. Animal identification: a reference Honolulu, x + 281 pp. guide. Vol. 1, marine and brackish water animals. British Mackie, G. o. (ed.), 1976. Coelenterate ecology and be- Museum (Natural History), London, x + III pp. havior. Plenum Press, N.Y., 744 pp. [CB 3] XIV

Smith, D. C. & Y. Tiffon (eds), 1980. Nutrition in the lower Thorpe, W. H. & D. Davenport (eds), 1964. Learning and Metazoa. Pergamon Press, Oxford, viii + 180 pp. associated phenomena in invertebrates. Anim. Behav. Sokolov, B. S., V. N. Dubatolov & A. B. Ivanovski (eds), Suppl. 1: viii + 190. 1974-1975. Ancient Cnidaria. Publishing House Nauka, Tokioka, T. & S. Nishimura (eds), 1973. Recent trends in Novosibirsk, vol. 1, 364 pp.; vol. 2, 280 pp. [mainly in research in coelenterate biology. Pubis Seto mar. bioI. Lab. Russian and English]. [FC I) 20: i-xxxii, 1-793, xxxiii-xlii. [eB 2] Stoddart, D. R. & R. E. Johannes (eds), 1978. Coral reefs: United Nations Environment Programme, 1984. Workshop research methods. UNESCO, Paris, xvi + 581 pp. on jellyfish blooms in the Mediterranean. xvi + 221 pp. Stoddart, D. R. & M. Yonge (eds), 1971. Regional variation Usherwood, P. N. R. & D. R. Newth (eds), 1975. 'Simple' in Indian Ocean coral reefs. Academic Press, London, nervous systems. Edward Arnold, London, 506 pp. xxxvi + 584 pp. Vernberg, W. B. (ed.), 1974. Symbiosis in the sea. University Stoddart, D. R. & M. Yonge (eds), 1978. The Northern Great of South Carolina Press, Columbia, xvi + 276 pp. Barrier Reef. The Royal Society, London, Wells, S. M. (ed.), 1988. Coral reefs of the world. Vol. 1, viii + 194 + 172 pp. xlviii + 373 pp. United Nations Environment Programme, Tardent, P. & R. Tardent (eds), 1980. Development and Gland. cellular biology of coelenterates. Elsevier/North-Holland, Wells, S. M. & M. D. Jenkins (eds), 1988. Coral reefs of the Amsterdam, xxiv + 499 pp. [eB 4] world. Vol. 3, I + 329 pp. United Nations Environment Taylor, D. L. (ed.), 1977. Proceedings: third international Programme, Gland. coral reef symposium. Rosenstiel School of Marine and Wells, S. M., R. M. Pyle & N. M. Collins (eds), 1983. The Atmospheric Science, Miami, vol. 1, xxiv + 656 pp.; vol. 2, mCN invertebrate red data book. IUCN, Gland, xxviii + 628 pp. [CR 3] 1 + 632 pp. xv

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 multtfor• mis 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 differen• tiation 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 XVll

The settlement and growth of Sertularia cupressina (Hydrozoa, Sertulariidae) in Langstone Harbour, Hampshire, UK 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 constric• tum 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 palaeo• ecological 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 environmen• tal 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. NEMATOCYSTS 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): nomen• clature 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