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Coralline Algae, A First Synthesis Author H.W. Johansen, Ph.D. Associate Professor of Botany Department of Biology Clark University Worcester, Massachusetts Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business First published 1981 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1981 by CRC Press, Inc. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright. com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not- for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging in Publication Data Johansen, H William. Coralline algae, a first synthesis Bibliography: p. Includes index. 1. Corallinaceae. I . Title. QK569.C8J64 589.4’1 80-14028 ISBN 0-8493-5261-4 A Library of Congress record exists under LC control number: 80014028 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-315-89182-8 (hbk) ISBN 13: 978-1-351-07092-8 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com PREFACE As the title implies, this book is a first step at pulling together the voluminous but scattered information on coralline algae. Much can be said about these omnipresent plants of the sea, and the purpose here is to provide a coherent framework of data and discussion. This si the time to make this step, because research on coralline algae si now resulting in a surge of papers. What was once a rather neglected group of seaweeds si now becoming relatively well-known. Marine biologists are rapidly recog- nizing the important role that coralline algae play in the sea. And, the old idea that coralline algae are an extraordinarily difficult group to work with si also being dis- pelled. More than most other seaweed groups, the Corallinaceae are well-defined. Hence, it si easy to scan papers and quickly pick out information on this group. There are two types of papers: those in which the research is focused on corallinea lgae and those in which data on these algae are part of a larger whole. A great deal of important literature was published from 1700 to the early 1900s. I have incorporated some of this, that which seems pertinent, especially in the chapter on taxonomy, but many others have been excluded. Rather, the aim of this book is to give a state-of-the-art presentation and to emphasize recent publications. Therefore, 1 have included asm uch information as possible upt oa nd including 1979. Another facet that reflects the well-defined status of the coralline family si the spec- ialized vocabulary that has crept into use. 1 hope the glossary will help in this regard. This book is aimed at marine biologists in general, and I have tried to keep the writing such that nonphycologists will not have undue difficulty with it. It is true, however, thats ome chapters, for example those on interal structures and reproduction, will bem ostu nderstandable by phycologists. In selecting chapter topics I kept in mind coralline algae- the plants. tA the same time I recognize that of the wider interest to marine biologists si the role that these plants play in the seas. Hence, the 12 chapters form 3 groups: (1) plant structure and organization, (2) ecology, and (3) taxonomy and phylogeny. Hopefully, there is an approximately even distribution in these three areas. Coverage in the area of physiol- ogy si lacking, however. The closest this book comes to physiology si in Chapter 6, on calcification - a must with corallinael gae. Many people have helped with this book, both in reading for content and grammar, and in logistics, such as typing, drawing, and procuring publications. 1 thank the fol- lowing people for reading parts of the manuscript or for exchanging ideas about cor- alline algae with me: W. H. Adey, V. Ahmadjian, H. Akioka, W. Andrake, L. F. Austin, E. A. Boger, M. A. Borowitzka, J. J. Brink, J. Cabioch, Y. M. Chamberlain, M. Chihara, B. J. Colthart, G. F. Elliot, D. J. Garbary, B. T. Gittins, M. H. Hom- mersand, S. E. Johnson, L. Irvine, P. A. Lebednik, M. Lemoine, M. M. Littler, T. Masaki, P. J. Matty, R. E. Meslin, J. D. Milliman, J. N. Norris, R. F. Nunnemacher, R. B. Searles, P. C. Silva, R. A. Townsend and W. J. Woelkerling. And there are surely others that I have inadvertently omitted. In spite of their help, it is possible that there are errors in the text, and for these I am fully responsible. Many thanks go to Elizabeth M. Rogers and Inis C. Cook, as well as Terry Rey- nolds, Rene Baril, Teri McCall, and Roxanne Rawson of the Secretarial Pool, Clark University, for hours of tedious typing. Thanks also go to R. B. Parker, R. E. Leven- baum, Irene W. Walch, and Marion Henderson for help with illustrations and refer- ences. The Cooperation of CRC Press is much appreciated: Lisa Levine Eggenberger, Benita Budd Segraves, and B. J. Starkoff. The initial days of work on this book in a lovely Montana cabin were possible because of Sylvia and Neil R. Schroeder, and S. S. Cook, Jr. (Bud). I would like to express my sincere gratitude to G. F. Papenfuss for starting me off on my study of coralline algae and for his continued interest. Much credit also goes to my family, Eric J., Brian F., Edith L., and Fredrik Johansen, as well as to Frances L. Pedusey, Carolee A. Virgilio, and Barbara J. Johansen for their understanding and support duringt he years when I was devoted to studying plants of the sea. H. W. nesnahoJ THAEU THOR H. William Johansen, Ph. D., is Associate Professor of Botany, Clark University, Worcester, MA. He obtained a B. A. degree from San Jose State University, an M.A. Degree from San Franciso State University ni 1961, and a Ph.D. degree from the Uni- versity of California in Berkeley in 1966. From 6691 to 8691 he had an N.I.H. post- doctoral fellowship, part of which was spent in South Africa and Europe, and part ni Berkeley. From 8691 to the present, he has held a position at Clark University in the Department of Biology. Dr. Johansen has been actively involved in teaching plant and marine-oriented courses. His research deals with coralline algae, particularly systematics and ecology. Recently he has traveled to the Gulf of California, Japan, and Canada to study these algae. He si a member of the American Association for the Advancement fo Science, the International Phycological Society, the American Phycological Society, the British Phycological Society, SigmXai , and the Western Society of Naturalists. Since his career ni research began, Dr. Johansen has published more than 03 papers and abstracts, mostly on coralline algae. Coralline Algae, A First Synthesis si his first book. Thibso ok si dedicated to my parents, Edith and Fredrik and my sons, Eric and thel ate Brian. TABLE OF CONTENTS Chapter 1 ScopeandDiversity ........................................................ 1 Introduction .............................................................. 1 Basicstructure ............................................................ 1 Roleintheoceans ......................................................... 8 Classification ............................................................ I0 Sunimary ................................................................ 10 Chapter 2 evitategeV Cytology ....................................................... 13 Introduction ............................................................. 13 Cellwalls ............................................................... 13 Primary Pit.Connections ................................................... 16 Secondary Pit.Connections ................................................. 16 CellFusions .............................................................. 19 Nuclei ................................................................... 20 Organellesand Inclusions .................................................. 20 SporeGermination ........................................................ 23 Sporeling Growth and Development ......................................... 25 Meristems ............................................................... 27 Epithallia ................................................................ 29 TrichocytesandMegacells .................................................. 33 Summary ................................................................ 36 Chapter 3 erutcurtS of enillaroC detalucitranoN Algae ................................... 39 Introduction ............................................................. 39 Lithophylloideae .......................................................... 41 Lithophyllum Series ....................................................... 42 Dermatolithon Series ...................................................... 42 Mastophoroideae and Melobesioideae ........................................ 44 Thincrusts .............................................................. 44 Ribbon Corallines ......................................................... 46 Thickcrusts ............................................................. 47 Unattached CorallineA lgae ................................................ 48 Epiphytic CorallineA lgae .................................................. 48 Parasitic CorallineA lgae ................................................... 52 Summary ................................................................ 55 Chapter 4 erutcurtS of enillaroC detalucitrA Algae ...................................... 57 Introduction ............................................................. 57 Corallinoideae ............................................................ 57 Intergenicula ............................................................. 57 Genicula ................................................................. 60 Branching ............................................................... 63 FrondInitiation .......................................................... 65 Amphiroideae ............................................................ 66 Amphiroa ............................................................... 67 Genicula ................................................................. 68 Lithothrix ............................................................... 72 ...................................................... Metagoniolithoideae 74 Summary ................................................................ 76 Chapter 5 Reproduction ............................................................ 79 Introduction ............................................................. 79 Vegetative Reproduction ................................................... 79 Conceptacles ............................................................. 79 TetrasporangialC onceptacles ............................................... 84 Tetrasporangia and Bisporangia ............................................. 19 MaleConceptacles ........................................................ 92 Spermatangia ............................................................ 97 Carpogonia .............................................................. 98 Carposporophytes ................................................... 101 LifeHistories ........................................................... 106 Summary ............................................................... 108 Chapter 6 Calcification ............................................................ 111 lntroduction ............................................................ 111 Cell Wall composition ................................................... 111 The Calcification Process ................................................. 112 Summary ............................................................... 117 Chapter 7 Phytogeography ......................................................... 119 Introduction ............................................................ 119 Cold Northwestern Atlantic ............................................... 119 Cold Northeastern Atlantic ............................................... 124 Cold Northwestern Pacific ................................................ 127 Cold Northeastern Pacific ................................................ 129 TropicalRegions ........................................................ 129 Below 30" South Latitude ................................................. 133 Summary ............................................................... 133 Chapter 8 Growth and Environment ................................................. 135 Introduction ............................................................ 135 Sporeasn d Substrates .................................................... 135 Seasonality ............................................................. 136 Temperature ............................................................ 137 Light .................................................................. 138 Desiccation ............................................................. 139 WaterMotion ........................................................... 140 Seawater Ingredients ..................................................... 142 BioticInteractions ....................................................... 143 Summary ............................................................... 147 Chapter 9 Production ............................................................. 149 Introduction ............................................................ 14 9 Growth Rates ........................................................... 149 Colonization and Succession .............................................. 150 OrganicProduction ...................................................... 151 InorganicProduction .................................................... 152 Unattached Coralline Algae ............................................... 153 FateoftheCalcite ....................................................... 154 Summary ............................................................... 157 Chapter 10 feeR Building ........................................................... 159 Introduction ............................................................ 159 Basic Reef Structure ..................................................... 159 Grazing and Coralline Development ........................................ 161 Indo-Pacific Reefs ....................................................... 162 Porolithon in the Pacific .................................................. 163 CaribbeanReefs ......................................................... 164 Coralline Ridge-Formers in theC aribbean ................................... 168 Summary ............................................................... 170 Chapter 1 1 Coralline Fossil Algae .................................................... 173 Introduction ............................................................ 173 Solenoporaceae ......................................................... 173 Ancestral Coralline Algae ................................................. 175 Appearance of theC orallinaceae ........................................... 176 Summary ............................................................... 177 Chapter 12 Taxonomy .............................................................. 179 Introduction ............................................................ 179 History ................................................................ 179 Recognizing the Genera ................................................... 183 Current Schemes of Classification .......................................... 187 Adaptations in Coralline Algae ............................................ 188 Summary ............................................................... 191 References .............................................................. 193 Glossary ............................................................... 209 Appendix1 ............................................................. 215 Appendix2 ............................................................. 225 Index .................................................................. 227

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