THE ALGAE

SOME OTHER ELBS LOW-PRICED EDITIONS

Ashby AN INTRODUCTION TO PLANT ECOLOGY Macmillan Bell and THE DIVERSITY OF GREEN PLANTS Arnold Woodcock

Burnett FUNDAMENTALSOF~YCOLOGY Arnold Fogg PHOTOSYNTHESIS English

Universities Press Heywood PLANT TAXONOMY Arnold Ingold BIOLOGY OF FUNGI Hutchinson

Educational Kershaw QUANTITATIVE AND DYNAMIC PLANT ECOLOGY Arnold Purseglove TROPICAL CROPS DICOTYLEDONS Longman Richardson TRANSLOCATION IN PLANTS Arnold Street and PLANT ~ETABOLISM Pergamon Cockburn

Street and THE PHYSIOLOGY OF FLOWERING PLANTS Arnold Opik

Sutcliffe PLANTS AND WATER Arnold Wareing and THE CoNTROL OF GROWTH AND Pergamon Phillips DIFFERENTIATION IN PLANTS

Wheeler AN INTRODUCTION TO PLANT DISEASES Wiley

THE ALGAE

V. J. CHAPMAN, M.A., PH.D. (Cantab.) Professor of Botany, University of Auckland

New Zealand

and

D. J. CHAPMAN, B.Sc. (Auck.), PH.D. (Cal.) Assistant Professor of Botany, University of Chicago

SECOND EDITION

ENGLISH LANGUAGE BOOK SOCIETY and

Macmillan Education

©D. J. Chapman 1962, 1973

All rights reserved. No part of this publication may be reproduced or transmitted, in any form or by any

means, without permission

First published January 1962 Reprinted June 1962, 1964, 1968, 1969

Second edition 1973 ELBS edition first published 1975

Reprinted 1975

Published by THE MACMILLAN PRESS LTD

London and Basingstoke Associated companies in New York Dublin

Melbourne Johannesburg and Madras

SBN 333 14270 5 (hard cover) 333 I 7439 9 (ELBS paper cover)

ISBN 978-1-349-27912-8 ISBN 978-1-349-27910-4 (eBook) DOI 10.1007/978-1-349-27910-4

The paperback edition of this book is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, resold, hired out, or otherwise circulated without the publisher's prior consent, in any form of binding or cover other than that in which it is published and without a similar condition in­cluding this condition being imposed on the

subsequent purchaser.

PREFACE TO SECOND EDITION

In preparing this new edition substantial changes have been made in both the arrangement and the material. The underlying system in the taxonomic portion is the same as in the first edition as we are convinced that this is the best way to introduce students to different plants. A great deal of work has been carried out on the flagellates and allied organisms in recent years and this has thrown new light on algal phylogeny. The original chapters on these groups have therefore been greatly expanded and we have also included the Euglenophyta in this new edition. In view of the importance of phyto­plankton in the sea and in fresh waters an entirely new chapter on this aspect of the algae has been added.

The book is intended as a brief introductory text only. Because of this the choice of material and references for inclusion must be highly selective. This is especially true of certain chapters, for example those on physiology and symbiosis, for which very extensive works are now available. The references are intended as a guide to further detailed study.

In preparing this new edition we want to express our thanks to the follow­ing experts who have very kindly read certain chapters and made very useful and valuable comments: Professor G. F. Papenfuss (Chlorophyta and Phaeophyta); Professor R. E. Norris (Rhodophyta); Professor I. Manton (Prasinophyceae and Haptophyceae); Professor Harlan Johnson (Fossil Algae); Professor J. E. Morton (Rocky coast Ecology); Professor H. Clemenc;on (Xanthophyceae); Dr. J. R. Taylor (Phytoplankton); Dr. U. V. Cassie (Bacillariophyta); Dr. A. R. Loeblich III (Pyrrhophyta). Whilst we are greatly indebted to the above specialists we accept full responsibility for the contents of this book and the way in which the material is set out.

We would like to thank the authorities at the Cranbrook Institute of Science who made Professor Chapman's visit to the United States possible and consequent personal collaboration on this book.

V. J. CHAPMAN Cranbrook Institute of Science and Auck­land University

D. J. CHAPMAN University of Chicago v

PREFACE TO FIRST EDITION

This book has been prepared in response to many requests that I have received from colleagues. Although the present volume follows the general lay-out of my first book An Introduction to the Study of Algae, there has been so much rearrangement, the complete rewriting of many chapters and the addition of new ones, that it cannot be considered a second edition of the first book. I am particularly grateful to the Syndics of the Cambridge University Press for releasing the original copyright so that it has been possible to use some portions of the first book. In recent years considerable advances have been made in the study of the algae and this has affected, not only our knowledge of their life histories, but also our views on phylogeny. These new results have necessitated much rearrangement of material. There has also been extensive new work on the ecology of the algae, so that the chapters on marine algal ecology and ecological factors are more or less completely new. The same is true also of algal physiology. In order that the survey should be more complete there is a chapter briefly describing the historical aspect of the subject, and also a chapter on the economic uses of algae.

The purpose of the book remains the same as my first volume, namely to provide a general survey of all aspects of the algae such as may be required by a University undergraduate, with selected portions (marked by an asterisk) that can be used by first year students or which are suitable for use in schools at the sixth form level. The type method of instruction has again been em­ployed because continued experience has not changed my belief in it. I still feel, however, that no study of the algae is complete without reference to the other aspects which have been included. I am aware that not everyone will agree with what has been included and material that has been excluded, nor with all the views expressed, but where there are divergencies of opinion, I have tried to see that the other view is also presented or made known. This is perhaps particularly true in the chapter on Evolution.

vii

viii PREFACE TO FIRST EDITION

There are a number of excellent works available to advanced students and research workers, and the present volume is not intended to compete with them in any way. Apart from those works to which I expressed my indebted­ness in the preface to An Introduction to the Study of Algae, there are some new books that have appeared since, and which have provided valuable reference material. I include here the second volume of Fritsch's Structure and Reproduction of the Algae, the Manual of Phychology, edited by the late Gilbert Smith, and the recent extensive work on the Rhodophyceae by the late Prof. Kylin. The sources of the new illustrations are acknowledged in the legends.

At the end of each chapter certain references are provided which can be consulted for further specific information. No attempt has been made to provide anything approaching a complete reference list and the choice of the various references is entirely my own. I am aware that this choice will not necessarily please everyone, but I hope it will prove useful.

I am grateful to my colleagues, Dr. J. A. Rattenbury for critical reading of the manuscript and to Dr. J. M.A. Brown for his reading and criticism of the two chapters on ecological factors and algal physiology. Dr. A. B. Cribb of the University of Brisbane has also read the proofs and made valuable suggestions for which I am most appreciative. My thanks go to Mrs. J. Rutherford for assistance in proof-reading.

V. J. CHAPMAN London 1960

CONTENTS

CHAPTER 1. CLASSIFICATION

2. CYANOPHYTA Cyanophyceae

3. CHLOROPHYTA Prasinophyceae:

Pyramimonadales Halosphaerales

Chlorophyceae: Volvocales Chlorococcales Ulotrichales

4. CHLOROPHYTA Chlorophyceae:

Oedogoniales Chaetophorales Siphonocladales Dasycladales

5. CHLOROPHYTA Chlorophyceae:

Derbesiales Siphonales Dichotomosiphonales Zygnematales

Charophyceae: Charales

6. EUGLENOPHYTA Euglenophyceae:

Euglenales ix

PAGE

1

13

28 30 31 32 35 50 59

77 77 81 90 96

101 101 101 108 109 117 117

124 125

X CONTENTS CHAPTER PAGE

XANTHOPHYTA 127 Xanthophyceae: 127

Chloramoebales 128 Rhizochlori dales 130 Heterogloea1es 130 Mischococca1es 130 Tribonematales 134 Vaucheriales 135

CHLOROMONADOPHYTA 140 Chloromonadophyceae: 140

Chloromonadaceae 140 7. CRYPTOPHYTA 144

Cryptophyceae: 144 Cryptomonadales 145 Cryptococcales 146

PYRRHOPHYTA 147 Desmophyceae: 148

Prorocentrales 148 Dinophyceae: 149

Gymnodiniales 150 Peridiniales 153 Dinophysidales 155 Dinocapsales 156 Dinotrichales 156 Dinamoebidiales 156 Phytodiniales 156

8. CHRYSOPHYTA 159 Chrysophyceae: 160

Chrysomonadales 161 Rhizochrysidales 165 Chrysocapsales 167 Chrysosphaerales 168 Phaeothamniales 169

Haptophyceae: 170 Isochrysidales 171 Prymnesiales 172

BACILLARIOPHYTA: 176 Bacillariophyceae 176

9. PHAEOPHYTA Phaeophyceae: 183

Ectocarpales 186 Tilopteridales 189 Sphacelariales 190 Cutleriales 193 Dictyotales 196

CHAPTER

Chordariales Sporochnales Desmarestiales Dictyosiphona1es

10. PHAEOPHYTA Phaeophyceae:

Laminariales Ascoseirales Durvilleales

CONTENTS

Fucales Fuco-Ascophylleae Cystoseiro-Sargassaceae Anomalae

11. RHODOPHYTA Rhodophyceae Bangiophycidae:

Bangiales Florideophycidae:

Nemalionales Bonnemaisoniales Gelidiales Cryptonemiales

12. RHODOPHYTA Florideophycideae:

Gigartinales Rhodymeniales Ceramiales

13. FOSSIL ALGAE

14. REPRODUCTION AND EVOLUTION

15. EcoLOGY OF RocKY CoAsTs

16. ECOLOGY OF SALT MARSHES AND MANGROVE SWAMPS

17. SOIL ALGAE AND SYMBIOSIS

18. FREsH WATER ECOLOGY

19. PHYTOPLANKTON

20. ECOLOGICAL FACTORS

21. BIOGEOGRAPHY AND LIFE FORM

22. ALGAL PHYSIOLOGY

INDEX

xi PAGE

198 209 209 211

220 236 237 239 241 249 251 256 256 262 265 269 270 273 276 278

288 298 301 309 328 351 372 381 388 404 421 453 465 481

GLOSSARY

Acronematic: Smooth flagellum with terminal hair. Akinete: Thick walled resting spore, generally derived from vegetative cell. Aplanospore: Non-motile spore. Autospore: Aplanospore resembling a mature cell. Auxospore: Cell resulting from syngamy (Bacillariophyceae ). Carpogonium: Female gametangium (Rhodophyceae ). Carposporangium: Sporangium resulting from division of zygote nucleus·

(Rhodophyceae). Carpospore: Spore produced by carposporangium (Rhodophyceae). Carposporophyte: Product of fertilized carpogonium (Rhodophyceae ). Coenocyte: Multinucleate cell lacking cross walls. Coenobium: A group of independent cells enclosed in a common envelope or

united together so that the colony functions as a single individual. Consocies: An ecological term used to denote a community in a succession in

which only a single species is dominant. Cryptostomata: Thallus cavities from which hairs arise (Phaeophyceae). Ecad: A term used to designate a morphological form of a species that is

related to a specific ecological niche. Endospore: Thin walled spore within the cell (Cyanophyceae). -etum: Termination used in ecology to distinguish an Association. Exospore: Spore formed, one at a time, from apex downwards (Cyano-

phyceae). Flimmergeissel: Flagellum with lateral flagellar hairs ('tinsel flagellum'). Frustule: One half of a cell (Bacillariophyceae ). Girdle: Transverse groove (Dinophyceae). Region where frustules overlap

(Bacillariophyceae). Gonimoblast: Gonimoblast filaments are the cells bearing the carposporangia

(Rhodophyceae ). xii

GLOSSARY xiii Haptonema: Flagella-like organ used for attachment to substrate (Hapto-

phyceae). Heterocyst: Spore-like structure (Cyanophyceae). Heterokont: Condition in which two flagella are of dissimilar morphology. Heteromorphic: Morphologically dissimilar. Heterotrichous: Combination of erect filament and prostrate thallus portion. Hormogone: Multicellular segment capable of gliding (Cyanophyceae). Hypnospore: A resistant spore of the same size and dimensions as the cell

from which it is formed. Isokont: Condition in which two flagella are of similar morphology. Isomorphic: Morphologically similar. Intercalary: Applied to describe a meristem in the middle of a filament or

thallus. Mastigoneme: Hair-like thread occurring on flagella. Meiospore: Spore formed at meiosis. Monosporangium: Vegetative cell that changes to produce a single spore

called monospore (Rhodophyceae and Phaeophyceae). Oospore: Zygospore resulting from oogamous fusion. Pantonematic: Arrangement describing flagellar hairs on both sides of

flagellum. Paraphyses: Sterile hairs or threads within sporangium or gametangium

(Phaeophyceae ). Peitschengeissel: Flagellum lacking lateral hairs: 'whiplash' or 'smooth'

flagellum. Pellicle: Thin membrane around the protoplast. Pericarp: Gametophyte tissue surrounding the carposporophyte (Rhodo-

phyceae). Pleuronematic: Flagellum with hairs= Flimmergeissel. Plurilocular: Multichambered gametangia or sporangia (Phaeophyceae ). Polyspore: Spore of a group in which more than four spores are formed from

mother cell. Pneumatocyst: Gas bladders or enlarged hollow area of stipe (Phaeophyceae ). Socies: A term used to designate a small community in a consocies dominated

by a subordinate species. Sorus: Cluster of sporangia (Phaeophyceae ). Statospore: Resting cell or spore formed within vegetative cell, usually with

ornamented cell wall. Stichonematic: Flagellum with lateral hairs on one side only. Synzoospore: Compound zoospore. Tetraspore: Spore from a group of four formed at meiosis. Tetrasporophyte: Plant producing tetras pores (Rhodophyceae ). Trichoblast: Colorless hair-like branch (Rhodophyceae). Trichocyst: Cytoplasmic organelle, often dart or thread-like and often

ejected by cell (Dinophyceae, Cryptophyceae and Chloromonadophy­ceae).

Trichogyne: Hair-like extremity of female gametangium (Rhodophyceae). Trichome: Filament (Cyanophyceae).

xiv GLOSSARY

Trichothal/ic: Describes growth at the base of uniseriate filament (Phaeo-phyceae).

Unilocular: Describing a single-chambered sporangium (Phaeophyceae). Zoochlorellae: Chlorophyceae endosymbiotic with invertebrates. Zooxanthe/lae: Dinophyceae endosymbiotic with invertebrates. Zoospore: Motile flagellated spore. Zygospore: Thick walled resting stage resulting when zygote from isogamy

ceases motility. Zygote: Diploid cell resulting from syngamy.