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CONTRIBUTORS Gerald J. Bakus Paul R. Burkholder Leon S. Ciereszko Joseph H. Connell A. B. Cribb Louis H. Di Salvo Robert Endean Peter W. Glynn J. Frederick Grassle T. K. B. Karns Leonard Muscatine Yu. I. Sorokin Masashi Yamaguchi BIOLOGY AND GEOLOGY OF CORAL REEFS EDITED BY O. A. JONES Department of Geology University of Queensland St. Lucia, Brisbane Queensland, Australia R. ENDEAN Department of Zoology University of Queensland St. Lucia > Brisbane Queensland, Australia VOLUME II: Biology 1 Θ A C A D E M I C PRESS New York and London A Subsidiary of Harcourt Brace Jovanovich, Publishers 1973 COPYRIGHT © 1973, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 LIBRARY OF CONGRESS CATALOG CARD NUMBER: 72-84377 PRINTED IN THE UNITED STATES OF AMERICA To the Great Barrier Reef Committee, its office-bearers and its many members who have worked unremittingly for fifty years to further our knowledge of the Great Barner Reef; and to the memory of geologist Professor H. C. Richards, one of its founders, and biologist Professor E. /. Goddard, an early enthusiastic supporter LIST OF CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors' contributions begin. J. (325) GERALD BAKUS,* National Academy of Sciences, Washington, D.C. PAUL R. BuRKHOLDER,f Department of Marine Sciences, University of Puerto Rico, Mayaguez, Puerto Rico (117) LEON S. CIERESZKO, Department of Chemistry, The University of Okla- homa, Norman, Oklahoma (183) H. CONNELL, Department of Biological Sciences, University of California, Santa Barbara, California (205) JOSEPH A. B. CRTJBB, Department of Botany, University of Queensland, St. Lucia, Brisbane, Australia (47) Louis H. D I SALVO, Naval Biomédical Research Laboratory, University of California, Berkeley, California ( 1 ) Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland, Australia (389) ROBERT ENDEAN, W. GLYNN, Smithsonian Tropical Research Institute, Balboa, Canal Zone (271) PETER J. Woods Hole Océanographie Institution, Woods Hole, Massachusetts (247) FREDERICK GRASSLE, T. K. B. KARNS, Department of Chemistry, The University of Oklahoma, Norman, Oklahoma (183) LEONARD MUSCATINE, Department of Biology, University of California, Los Angeles, California (77) Yu. I. SOROKIN, Institute of Freshwater Biology, Academy of Sciences of the USSR, Borok, Garoslavl, USSR (47) MASASHI YAMAGUCHI, The Marine Laboratory, University of Guam, Agana, Guam (369) * Present address: Allan Hancock Foundation, University of Southern California, Los Angeles, California. f Deceased. xiii GENERAL PREFACE This four-volume work (two volumes covering geological and two biological topics) originated from an article on The Great Barrier Reefs of Australia written by the editors and published in the November 1967 issue of Science Journal. The prime aim of this treatise is to publish in one source as many as possible of the major advances made in the diverse facets of coral reef problems, advances scattered in a multitude of papers and published in a variety of journals. Initially a one-volume work was projected, but the wealth of material available led to this four-volume treatise. Two conta