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Freshwater Algae
Identification and Use as Bioindicators
Edward G. Bellinger
Environmental Science and Policy, Central European University, Hungary
and
David C. Sigee
Manchester University, UK
A John Wiley & Sons, Ltd, Publication
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Freshwater Algae
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Freshwater Algae
Identification and Use as Bioindicators
Edward G. Bellinger
Environmental Science and Policy, Central European University, Hungary
and
David C. Sigee
Manchester University, UK
A John Wiley & Sons, Ltd, Publication
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C 2010 by John Wiley & Sons, Ltd
This edition first published 2010, Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific, Technical and Medical
business with Blackwell Publishing.
Registered Office:
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All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by
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competent professional should be sought.
Library of Congress Cataloguing-in-Publication Data
Bellinger, E. G.
Freshwater algae : identification and use as bioindicators / Edward G. Bellinger and David C. Sigee.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-470-05814-5 (cloth)
1. Freshwater algae. 2. Indicators (Biology). 3. Environmental monitoring. I. Sigee, D. C. II. Title.
QK570.25.B45 2010
579.8–dc22
2009052145
A catalogue record for this book is available from the British Library.
ISBN 978-0-470-05814-5
Set in 10/12pt Times by Aptara Inc., New Delhi, India
Printed in Great Britain by Antony Rowe, Ltd. Chippenham, Wilts
First impression 2010
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Contents
Preface
ix
Copyright Acknowledgements
xi
1 Introduction to Freshwater Algae
1
1.1
1.2
1.3
1.4
1.5
1.6
General introduction
1.1.1 Algae – an overview
1.1.2 Algae as primary producers
1.1.3 Freshwater environments
1.1.4 Planktonic and benthic algae
1.1.5 Size and shape
Taxonomic variation – the major groups of algae
1.2.1 Microscopical appearance
1.2.2 Biochemistry and cell structure
1.2.3 Molecular characterization and identification
Blue-green algae
1.3.1 Cytology
1.3.2 Morphological and taxonomic diversity
1.3.3 Ecology
1.3.4 Blue-green algae as bio-indicators
Green algae
1.4.1 Cytology
1.4.2 Morphological diversity
1.4.3 Ecology
1.4.4 Green algae as bioindicators
Euglenoids
1.5.1 Cytology
1.5.2 Morphological diversity
1.5.3 Ecology
1.5.4 Euglenoids as bioindicators
Yellow-green algae
1.6.1 Cytology
1.6.2 Morphological diversity
1.6.3 Ecology
1.6.4 Yellow-green algae as bioindicators
v
1
1
2
3
3
4
5
8
9
9
12
12
13
15
17
17
17
17
18
21
21
22
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CONTENTS
1.7
1.8
1.9
1.10
1.11
1.12
2
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Dinoflagellates
1.7.1
Cytology
1.7.2
Morphological diversity
1.7.3
Ecology
Cryptomonads
1.8.1
Cytology
1.8.2
Comparison with euglenoid algae
1.8.3
Biodiversity
1.8.4
Ecology
1.8.5
Cryptomonads as bioindicators
Chrysophytes
1.9.1
Cytology
1.9.2
Morphological diversity
1.9.3
Ecology
1.9.4
Chrysophytes as bioindicators
Diatoms
1.10.1 Cytology
1.10.2 Morphological diversity
1.10.3 Ecology
1.10.4 Diatoms as bioindicators
Red algae
Brown algae
Sampling, Biomass Estimation and Counts of Freshwater Algae
A. PLANKTONIC ALGAE
2.1
2.2
2.3
2.4
2.5
2.6
Protocol for collection
2.1.1
Standing water phytoplankton
2.1.2
River phytoplankton
Mode of collection
2.2.1
Phytoplankton trawl net
2.2.2
Volume samplers
2.2.3
Integrated sampling
2.2.4
Sediment traps
Phytoplankton biomass
2.3.1
Turbidity
2.3.2
Dry weight and ash-free dry weight
2.3.3
Pigment concentrations
Flow cytometry: automated analysis of phytoplankton populations
Microscope counts of species populations
2.5.1
Sample preservation and processing
2.5.2
Species counts
2.5.3
Conversion of species counts to biovolumes
2.5.4
Chemical cleaning of diatoms
Diversity within single-species populations
2.6.1
Molecular analysis
2.6.2
Analytical microscopical techniques
25
25
26
27
28
28
29
29
30
30
31
31
31
32
32
33
33
35
38
39
40
40
41
41
41
42
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50
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CONTENTS
B. NON-PLANKTONIC ALGAE
2.7
2.8
2.9
2.10
3
82
82
83
84
84
84
84
87
89
89
89
91
93
93
93
95
Algae as Bioindicators
99
3.1
99
99
100
101
102
105
105
106
110
119
121
121
121
122
123
126
128
133
133
134
3.2
3.3
3.4
3.5
4
Deep water benthic algae
2.7.1
Benthic–pelagic coupling
2.7.2
Benthic algae and sediment stability
2.7.3
Invertebrate grazing of benthic algae
Shallow water communities
2.8.1
Substrate
2.8.2
Algal communities
Algal biofilms
2.9.1
Mucilaginous biofilms
2.9.2
Biomass
2.9.3
Taxonomic composition
2.9.4
Matrix structure
Periphyton – algal mats
2.10.1 Inorganic substratum
2.10.2 Plant surfaces
vii
Bioindicators and water quality
3.1.1
Biomarkers and bioindicators
3.1.2
Characteristics of bioindicators
3.1.3
Biological monitoring versus chemical measurements
3.1.4
Monitoring water quality: objectives
Lakes
3.2.1
Contemporary planktonic and attached algae as bioindicators
3.2.2
Fossil algae as bioindicators: lake sediment analysis
3.2.3
Water quality parameters: inorganic and organic nutrients, acidity and heavy metals
Wetlands
Rivers
3.4.1
The periphyton community
3.4.2
River diatoms
3.4.3
Evaluation of the diatom community
3.4.4
Human impacts and diatom indices
3.4.5
Calculation of diatom indices
3.4.6
Practical applications of diatom indices
Estuaries
3.5.1
Ecosystem complexity
3.5.2
Algae as estuarine bioindicators
A Key to the More Frequently Occurring Freshwater Algae
4.1
4.2
Introduction to the key
4.1.1
Using the key
4.1.2
Morphological groupings
Key to the main genera and species
137
137
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CONTENTS
4.3
4.4
List of algae included and their occurrence in the key
Algal identification: bibliography
241
243
Glossary
245
References
255
Index
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Preface
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 scums.
Freshwater algae constitute a very diverse group
of organisms. Their range of shapes and beauty when
viewed through a microscope have 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 in the list of copyright 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.
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Copyright Acknowledgements
We are very grateful to the following individuals for
allowing us to publish previously unpublished matter: Andrew Dean (Tables 2.3, 2.4), Matt Capstick
(Figs. 4.10–4.12, 4.42, 4.62, 4.64, 4.66, 4.68–4.70a,
4.73a) and Huda Qari (Fig. 2.8).
We also thank the following copyright holders for
giving us permission to use previously published
material – Academic Press, American Health Association, Cambridge University Press, Journal of
Plankton Research, McGraw-Hill, Phycologia, Prentice Hall.
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1
Introduction to Freshwater Algae
1.1
General introduction
Algae are widely present in freshwater environments,
such as lakes and rivers, where they are typically
present as micro-organisms – visible only with the aid
of a light microscope. Although relatively inconspicuous, they have a major importance in the freshwater
environment, both in terms of fundamental ecology
and in relation to human use of natural resources.
This book considers the diversity of algae in freshwater environments and gives a general overview of
the major groups of these organisms (Chapter 1),
methods of collection and enumeration (Chapter 2)
and keys to algal groups and major genera (Chapter 4). Algae are considered as indicators of environmental conditions (bioindicators) in terms of
individual species (Chapter 1) and as communities
(Chapter 3).
1.1.1
Algae – an overview
The word ‘algae’ originates from the Latin word for
seaweed and is now applied to a broad assemblage
of organisms that can be defined both in terms of
morphology and general physiology. They are simple
organisms, without differentiation into roots, stems
and leaves – and their sexual organs are not enclosed within protective coverings. In terms of physiology, they are fundamentally autotrophic (obtaining
all their materials from inorganic sources) and phoFreshwater Algae: Identification and Use as Bioindicators
C 2010 John Wiley & Sons, Ltd
tosynthetic – generating complex carbon compounds
from carbon dioxide and light energy. Some algae
have become secondarily heterotrophic, taking up
complex organic molecules by organotrophy or heterotrophy (Tuchman, 1996), but still retaining fundamental genetic affinities with their photosynthetic
relatives (Pfandl et al., 2009).
The term ‘algae’ (singular alga) is not strictly a
taxonomic term but is used as an inclusive label for
a number of different phyla that fit the broad description noted above. These organisms include both
prokaryotes (Section 1.3, cells lacking a membranebound nucleus) and eukaryotes (cells with a nucleus
plus typical membrane-bound organelles).
Humans have long made use of algal species, both
living and dead. Fossil algal diatomite deposits, for
example, in the form of light but strong rocks, have
been used as building materials and filtration media in
water purification and swimming pools. Some fossil
algae, such as Botryococcus, can give rise to oil-rich
deposits. Certain species of green algae are cultivated
for the purpose of extracting key biochemicals for
use in medicine and cosmetics. Even blue-green algae, often regarded as nuisance organisms, may have
beneficial uses. This is particularly the case for Spirulina, which was harvested by the Aztecs of Mexico
and is still used by the people around Lake Chad as
a dietary supplement. Spirulina tablets may still be
obtained in some health food shops. Blue-green algae are, however, are better known in the freshwater
environment as nuisance organisms, forming dense
blooms having adverse effects on human activities
Edward G. Bellinger and David C. Sigee