FRESHWATER ALGAEIdentification, Enumeration and Use as Bioindicators

SECOND EDITION

Edward G. BellingerDavid C. Sigee

JWST514-fm JWST514-Bellinger Printer: Yet to Come November 22, 2014 6:56 246mm×189mm

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JWST514-fm JWST514-Bellinger Printer: Yet to Come November 22, 2014 6:56 246mm×189mm

Freshwater Algae

i

Freshwater AlgaeIdentification, Enumeration and Use as Bioindicators

Second Edition

Edward G. BellingerDepartment of Environmental Sciences and Policy,

Central European University, Hungary

and

David C. SigeeSchool of Earth, Atmospheric and Environmental Sciences,

University of Manchester, UK

This edition first published 2015 © 2015 by John Wiley & Sons, Ltd

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Library of Congress Cataloging-in-Publication Data

Bellinger, Edward G., author.Freshwater algae : identification, enumeration and use as bioindicators / Edward G. Bellinger and David C. Sigee. – 2e.

pages cmIncludes index.ISBN 978-1-118-91716-9 (hardback)

1. Freshwater algae. 2. Indicators (Biology) 3. Environmental monitoring. I. Sigee, David C., author. II. Title.QK570.25.B45 2015579.8′176–dc23

2014031375

A catalogue record for this book is available from the British Library.

Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronicbooks.

Cover image: David SigeeCover design by Soephian Zainal

Set in 10/12pt Times by Aptara Inc., New Delhi, India.

2 2015

Contents

Preface to the First Edition ix

Preface to the Second Edition xi

Acknowledgements xiii

1 Introduction to Freshwater Algae 1

1.1 General introduction 11.1.1 Algae – an overview 11.1.2 Algae as primary producers 21.1.3 Freshwater environments 21.1.4 Planktonic and benthic algae 21.1.5 Size and shape 4

1.2 Taxonomic variation – the major groups of algae 51.2.1 Microscopical appearance 81.2.2 Biochemistry and cell structure 81.2.3 Molecular characterisation and identification 10

1.3 Blue-green algae 131.3.1 Cytology 131.3.2 Morphological and taxonomic diversity 141.3.3 Ecology 151.3.4 Blue-green algae as bioindicators 16

1.4 Green algae 171.4.1 Cytology 181.4.2 Morphological diversity 181.4.3 Ecology 191.4.4 Green algae as bioindicators 22

1.5 Euglenoids 221.5.1 Cytology 231.5.2 Morphological diversity 231.5.3 Ecology 241.5.4 Euglenoids as bioindicators 24

1.6 Yellow-green algae 241.6.1 Cytology 241.6.2 Morphological diversity 251.6.3 Ecology 251.6.4 Yellow-green algae as bioindicators 26

vi CONTENTS

1.7 Dinoflagellates 261.7.1 Cytology 261.7.2 Morphological diversity 281.7.3 Ecology 28

1.8 Cryptomonads 291.8.1 Cytology 291.8.2 Comparison with euglenoid algae 301.8.3 Biodiversity 311.8.4 Ecology 311.8.5 Cryptomonads as bioindicators 32

1.9 Chrysophytes 321.9.1 Cytology 321.9.2 Morphological diversity 321.9.3 Ecology 321.9.4 Chrysophytes as bioindicators 33

1.10 Diatoms 341.10.1 Cytology 351.10.2 Morphological diversity 381.10.3 Ecology 401.10.4 Diatoms as bioindicators 41

1.11 Red algae 411.12 Brown algae 42

2 Sampling, Biomass Estimation and Counts of Freshwater Algae 43

A. PLANKTONIC ALGAE 43

2.1 Protocol for collection 432.1.1 Standing water phytoplankton 442.1.2 River phytoplankton 47

2.2 Mode of collection 482.2.1 Phytoplankton trawl net 482.2.2 Volume samplers 502.2.3 Integrated sampling 512.2.4 Sediment traps 52

2.3 Phytoplankton biomass 532.3.1 Turbidity 532.3.2 Dry weight and ash-free dry weight 542.3.3 Pigment concentrations 55

2.4 Flow cytometry: automated analysis of phytoplankton populations 602.5 Biodiversity of mixed-species populations: microscope counts and biovolumes 63

2.5.1 Sample preservation and processing 632.5.2 Chemical cleaning of diatoms 642.5.3 Species counts 652.5.4 Conversion of species counts to biovolumes 722.5.5 Indices of biodiversity 77

2.6 Biodiversity within single-species populations 782.6.1 Molecular analysis 782.6.2 Analytical microscopical techniques 79

CONTENTS vii

B. NON-PLANKTONIC ALGAE 83

2.7 Deep-water benthic algae 842.7.1 Benthic-pelagic coupling 842.7.2 Benthic algae and sediment stability 852.7.3 Invertebrate grazing of benthic algae 86

2.8 Shallow-water communities 862.8.1 Substrate 862.8.2 Algal communities 89

2.9 Algal biofilms 902.9.1 Mucilaginous biofilms 912.9.2 Biomass 912.9.3 Taxonomic composition 922.9.4 Matrix structure 94

2.10 Periphyton – algal mats 952.10.1 Inorganic substratum 952.10.2 Plant surfaces 95

3 Algae as Bioindicators 101

3.1 Bioindicators and water quality 1013.1.1 Biomarkers and bioindicators 1013.1.2 Characteristics of bioindicators 1023.1.3 Biological monitoring versus chemical measurements 1033.1.4 Monitoring water quality: objectives 104

3.2 Lakes 1073.2.1 Contemporary planktonic and attached algae as bioindicators 1073.2.2 Fossil algae as bioindicators: lake sediment analysis 1083.2.3 Water quality parameters: inorganic and organic nutrients, acidity and

heavy metals 1113.3 Wetlands 121

3.3.1 Marshes 1213.3.2 Peatlands 123

3.4 Rivers 1233.4.1 The periphyton community 1233.4.2 River diatoms 1233.4.3 Evaluation of the diatom community 1243.4.4 Human impacts and diatom indices 1263.4.5 Calculation of diatom indices 1283.4.6 Practical applications of diatom indices 1303.4.7 Nitrogen-fixing blue-green algae 135

3.5 Estuaries 1353.5.1 Ecosystem complexity 1363.5.2 Algae as estuarine bioindicators 137

4 A Key to the More Frequently Occurring Freshwater Algae 141

4.1 Introduction to the key 1414.1.1 Using the key 1414.1.2 Morphological groupings 142

viii CONTENTS

4.2 Key to the main genera and species 1424.3 List of algae included and their occurrence in the key 2494.4 Algal identification: bibliography 251

Glossary 253

References 259

Index 269

Preface to the First Edition

Almost any freshwater or brackish water site willcontain one or many species of algae. Although theyare mainly microscopic and therefore not as visuallyapparent as larger aquatic organisms, such as higherplants or fish, algae play an equally important rolein the ecology of these water bodies. Their presencecan sometimes be noticed when they occur as densepopulations, colouring the water and in some casesforming massive surface scum.

Freshwater algae constitute a very diverse group oforganisms. Their range of shapes and beauty, whenviewed through a microscope, has delighted biolo-gists for more than a hundred years. They have anenormous range of size from less than one micrometreto several centimetres (for the stoneworts) – equallingthe size span (104) for higher plants seen in a trop-ical rainforest. Algal morphology is diverse, rang-ing from single cells to complex colonies and fil-aments. Some species are capable of active move-ment. The term ‘algae’ embraces a number of phyla(e.g. Cyanophyta, Bacillariophyta and Chlorophyta)of chlorophyll-containing organisms with differentgrowth forms and cytologies. Algae are importantprimary producers in both freshwater and marine sys-tems. In many lakes and rivers, they generate biomasswhich is the foundation of diverse food chains.Although algae have beneficial impacts on aquaticecosystems, they can also have adverse effects. Whenpresent in very large numbers they can produce‘blooms’ that, on decomposition, deoxygenate thewater – causing fish death and other ecological prob-lems. Some algae produce toxins that are lethal toboth aquatic and terrestrial organisms. It is importantto be aware of these impacts and to monitor watersfor the presence of these potentially harmful organ-isms. Algae can be used to flag up and assess a range

of human and natural impacts in aquatic systemsbecause of their often rapid response to changes in theenvironment. Examples include nutrient enrichment(eutrophication), industrial pollution and changes tothe hydrological regime of the water body. Somegroups of algae preserve well as fossils in geologicaldeposits such as lake sediments, analysis of whichgives us information on past environmental changes.

This book comes at a time of increasing concernover the widespread effects of human activities onthe general environment of this planet. Monitoringshifts in algal population gives us an insight into thesechanges. We need to be able to assess the ‘health’of aquatic systems such as lakes and rivers, sincewater is vital to both human and general ecosystemsurvival. Knowledge of algal population dynamicscan help us develop effective management strategiesfor those systems. Included in this book are sec-tions on the general features of the main freshwateralgal groups with notes on their ecology, methodsof sample collection and enumeration, using algaeas indictors of environmental conditions and, finally,a key to the identification of the more frequentlyoccurring genera. The authors have tried to combinedescriptive material with original colour photographsand line drawings, where possible, to help the reader.We would also like to gratefully acknowledge thehelp and encouragement of colleagues and students,and particularly appreciate the direct contributionsof postdoctoral workers and research students men-tioned under Acknowledgements. We would also liketo thank our families for their understanding andpatience during the preparation of this text.

We hope that all those using the book will findit useful, and will enjoy the numerous colour pho-tographs of these very beautiful organisms.

Preface to the Second Edition

Revisions of the first edition have been carried outto give a general update and to broaden the globalperspective. In Chapter 4, particularly, a substantialnumber of new photographs have been contributedfrom the United States and China (see Acknowl-

edgements), and various plates have been redrawnto provide greater detail. The key has been exten-sively modified to give greater clarity and to provideadditional information on several genera.

Acknowledgements

We are very grateful to Andrew Dean (Tables 2.3and 2.4), Matt Capstick (Figs. 4.8, 4.10–4.12, 4.42,4.62, 4.64, 4.66, 4.68–4.70a and 4.73a) and HudaQari (Fig. 2.8) for allowing us to present previouslyunpublished data.

We also thank Academic Press, American HealthAssociation, Cambridge University Press, Journalof Plankton Research, McGraw-Hill, Phycologiaand Prentice Hall for giving us permission to usepreviously published data.

With the incorporation of a substantial amount ofnew material into the second edition of the book, we

would particularly like to thank two colleagues fromthe United States and the Republic of China for theircontributions:

Dr. Robin A. Matthews (Western Washington Uni-versity, Bellingham, WA) – Figs. 4.2a, 4.2b, 4.4b,4.18, 4.19, 4.24d, 4.29, 4.36, 4.45, 4.49, 4.51a,4.51b, 4.52a, 4.53, 4.54 and 4.55.

Dr. Gaohua Ji (Shanghai Ocean University, Shang-hai, China) – Figs. 4.8, 4.31, 4.42, 4.47, 4.48, 4.53and 4.63.