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Freshwater Algae: Identification, Enumeration and Use as Bioindicators
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FRESHWATER
ALGAE
Identification, Enumeration
and Use as Bioindicators
SECOND EDITION
Edward G. Bellinger
David C. Sigee
JWST514-fm JWST514-Bellinger Printer: Yet to Come November 22, 2014 6:56 246mm×189mm
ii
JWST514-fm JWST514-Bellinger Printer: Yet to Come November 22, 2014 6:56 246mm×189mm
Freshwater Algae
i
Freshwater Algae
Identification, Enumeration and Use as Bioindicators
Second Edition
Edward G. Bellinger
Department of Environmental Sciences and Policy,
Central European University, Hungary
and
David C. Sigee
School of Earth, Atmospheric and Environmental Sciences,
University of Manchester, UK
This edition first published 2015 © 2015 by John Wiley & Sons, Ltd
Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester,
West Sussex, PO19 8SQ, UK
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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 cm
Includes 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 2015
579.8′
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 electronic
books.
Cover image: David Sigee
Cover 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 1
1.1.1 Algae – an overview 1
1.1.2 Algae as primary producers 2
1.1.3 Freshwater environments 2
1.1.4 Planktonic and benthic algae 2
1.1.5 Size and shape 4
1.2 Taxonomic variation – the major groups of algae 5
1.2.1 Microscopical appearance 8
1.2.2 Biochemistry and cell structure 8
1.2.3 Molecular characterisation and identification 10
1.3 Blue-green algae 13
1.3.1 Cytology 13
1.3.2 Morphological and taxonomic diversity 14
1.3.3 Ecology 15
1.3.4 Blue-green algae as bioindicators 16
1.4 Green algae 17
1.4.1 Cytology 18
1.4.2 Morphological diversity 18
1.4.3 Ecology 19
1.4.4 Green algae as bioindicators 22
1.5 Euglenoids 22
1.5.1 Cytology 23
1.5.2 Morphological diversity 23
1.5.3 Ecology 24
1.5.4 Euglenoids as bioindicators 24
1.6 Yellow-green algae 24
1.6.1 Cytology 24
1.6.2 Morphological diversity 25
1.6.3 Ecology 25
1.6.4 Yellow-green algae as bioindicators 26
vi CONTENTS
1.7 Dinoflagellates 26
1.7.1 Cytology 26
1.7.2 Morphological diversity 28
1.7.3 Ecology 28
1.8 Cryptomonads 29
1.8.1 Cytology 29
1.8.2 Comparison with euglenoid algae 30
1.8.3 Biodiversity 31
1.8.4 Ecology 31
1.8.5 Cryptomonads as bioindicators 32
1.9 Chrysophytes 32
1.9.1 Cytology 32
1.9.2 Morphological diversity 32
1.9.3 Ecology 32
1.9.4 Chrysophytes as bioindicators 33
1.10 Diatoms 34
1.10.1 Cytology 35
1.10.2 Morphological diversity 38
1.10.3 Ecology 40
1.10.4 Diatoms as bioindicators 41
1.11 Red algae 41
1.12 Brown algae 42
2 Sampling, Biomass Estimation and Counts of Freshwater Algae 43
A. PLANKTONIC ALGAE 43
2.1 Protocol for collection 43
2.1.1 Standing water phytoplankton 44
2.1.2 River phytoplankton 47
2.2 Mode of collection 48
2.2.1 Phytoplankton trawl net 48
2.2.2 Volume samplers 50
2.2.3 Integrated sampling 51
2.2.4 Sediment traps 52
2.3 Phytoplankton biomass 53
2.3.1 Turbidity 53
2.3.2 Dry weight and ash-free dry weight 54
2.3.3 Pigment concentrations 55
2.4 Flow cytometry: automated analysis of phytoplankton populations 60
2.5 Biodiversity of mixed-species populations: microscope counts and biovolumes 63
2.5.1 Sample preservation and processing 63
2.5.2 Chemical cleaning of diatoms 64
2.5.3 Species counts 65
2.5.4 Conversion of species counts to biovolumes 72
2.5.5 Indices of biodiversity 77
2.6 Biodiversity within single-species populations 78
2.6.1 Molecular analysis 78
2.6.2 Analytical microscopical techniques 79
CONTENTS vii
B. NON-PLANKTONIC ALGAE 83
2.7 Deep-water benthic algae 84
2.7.1 Benthic-pelagic coupling 84
2.7.2 Benthic algae and sediment stability 85
2.7.3 Invertebrate grazing of benthic algae 86
2.8 Shallow-water communities 86
2.8.1 Substrate 86
2.8.2 Algal communities 89
2.9 Algal biofilms 90
2.9.1 Mucilaginous biofilms 91
2.9.2 Biomass 91
2.9.3 Taxonomic composition 92
2.9.4 Matrix structure 94
2.10 Periphyton – algal mats 95
2.10.1 Inorganic substratum 95
2.10.2 Plant surfaces 95
3 Algae as Bioindicators 101
3.1 Bioindicators and water quality 101
3.1.1 Biomarkers and bioindicators 101
3.1.2 Characteristics of bioindicators 102
3.1.3 Biological monitoring versus chemical measurements 103
3.1.4 Monitoring water quality: objectives 104
3.2 Lakes 107
3.2.1 Contemporary planktonic and attached algae as bioindicators 107
3.2.2 Fossil algae as bioindicators: lake sediment analysis 108
3.2.3 Water quality parameters: inorganic and organic nutrients, acidity and
heavy metals 111
3.3 Wetlands 121
3.3.1 Marshes 121
3.3.2 Peatlands 123
3.4 Rivers 123
3.4.1 The periphyton community 123
3.4.2 River diatoms 123
3.4.3 Evaluation of the diatom community 124
3.4.4 Human impacts and diatom indices 126
3.4.5 Calculation of diatom indices 128
3.4.6 Practical applications of diatom indices 130
3.4.7 Nitrogen-fixing blue-green algae 135
3.5 Estuaries 135
3.5.1 Ecosystem complexity 136
3.5.2 Algae as estuarine bioindicators 137
4 A Key to the More Frequently Occurring Freshwater Algae 141
4.1 Introduction to the key 141
4.1.1 Using the key 141
4.1.2 Morphological groupings 142
viii CONTENTS
4.2 Key to the main genera and species 142
4.3 List of algae included and their occurrence in the key 249
4.4 Algal identification: bibliography 251
Glossary 253
References 259
Index 269
Preface to the First Edition
Almost any freshwater or brackish water site will
contain one or many species of algae. Although they
are mainly microscopic and therefore not as visually
apparent as larger aquatic organisms, such as higher
plants or fish, algae play an equally important role
in the ecology of these water bodies. Their presence
can sometimes be noticed when they occur as dense
populations, colouring the water and in some cases
forming massive surface scum.
Freshwater algae constitute a very diverse group of
organisms. Their range of shapes and beauty, when
viewed through a microscope, has delighted biologists for more than a hundred years. They have an
enormous range of size from less than one micrometre
to several centimetres (for the stoneworts) – equalling
the size span (104) for higher plants seen in a tropical rainforest. Algal morphology is diverse, ranging from single cells to complex colonies and filaments. Some species are capable of active movement. The term ‘algae’ embraces a number of phyla
(e.g. Cyanophyta, Bacillariophyta and Chlorophyta)
of chlorophyll-containing organisms with different
growth forms and cytologies. Algae are important
primary producers in both freshwater and marine systems. In many lakes and rivers, they generate biomass
which is the foundation of diverse food chains.
Although algae have beneficial impacts on aquatic
ecosystems, they can also have adverse effects. When
present in very large numbers they can produce
‘blooms’ that, on decomposition, deoxygenate the
water – causing fish death and other ecological problems. Some algae produce toxins that are lethal to
both aquatic and terrestrial organisms. It is important
to be aware of these impacts and to monitor waters
for the presence of these potentially harmful organisms. Algae can be used to flag up and assess a range
of human and natural impacts in aquatic systems
because of their often rapid response to changes in the
environment. Examples include nutrient enrichment
(eutrophication), industrial pollution and changes to
the hydrological regime of the water body. Some
groups of algae preserve well as fossils in geological
deposits such as lake sediments, analysis of which
gives us information on past environmental changes.
This book comes at a time of increasing concern
over the widespread effects of human activities on
the general environment of this planet. Monitoring
shifts in algal population gives us an insight into these
changes. We need to be able to assess the ‘health’
of aquatic systems such as lakes and rivers, since
water is vital to both human and general ecosystem
survival. Knowledge of algal population dynamics
can help us develop effective management strategies
for those systems. Included in this book are sections on the general features of the main freshwater
algal groups with notes on their ecology, methods
of sample collection and enumeration, using algae
as indictors of environmental conditions and, finally,
a key to the identification of the more frequently
occurring genera. The authors have tried to combine
descriptive material with original colour photographs
and line drawings, where possible, to help the reader.
We would also like to gratefully acknowledge the
help and encouragement of colleagues and students,
and particularly appreciate the direct contributions
of postdoctoral workers and research students mentioned under Acknowledgements. We would also like
to thank our families for their understanding and
patience during the preparation of this text.
We hope that all those using the book will find
it useful, and will enjoy the numerous colour photographs of these very beautiful organisms.
Preface to the Second Edition
Revisions of the first edition have been carried out
to give a general update and to broaden the global
perspective. In Chapter 4, particularly, a substantial
number of new photographs have been contributed
from the United States and China (see Acknowledgements), and various plates have been redrawn
to provide greater detail. The key has been extensively modified to give greater clarity and to provide
additional information on several genera.
Acknowledgements
We are very grateful to Andrew Dean (Tables 2.3
and 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 Huda
Qari (Fig. 2.8) for allowing us to present previously
unpublished data.
We also thank Academic Press, American Health
Association, Cambridge University Press, Journal
of Plankton Research, McGraw-Hill, Phycologia
and Prentice Hall for giving us permission to use
previously published data.
With the incorporation of a substantial amount of
new material into the second edition of the book, we
would particularly like to thank two colleagues from
the United States and the Republic of China for their
contributions:
Dr. Robin A. Matthews (Western Washington University, 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, Shanghai, China) – Figs. 4.8, 4.31, 4.42, 4.47, 4.48, 4.53
and 4.63.