Freshwater Algae of North America: Ecology and Classification

Freshwater Algae of North America

Ecology and Classification

Second Edition

John D. Wehr

Louis Calder Center—Biological Station, Fordham University, Armonk, New York, USA

Robert G. Sheath

Department of Biological Sciences, California State University San Marcos, San Marcos, California, USA

J. Patrick Kociolek

Department of Ecology and Evolutionary Biology and Museum of Natural History, University of Colorado, Boulder, Colorado, USA

University of Michigan Biological Station, Pellston, Michigan, USA

Table of Contents

Cover image

Title page

Copyright

Dedication

Contributors

Preface

First Edition

Second Edition

Chapter 1: Introduction to the Freshwater Algae

Abstract

I Introduction

II Classification

III Groups of Freshwater Algae

Chapter 2: Habitats of Freshwater Algae

Abstract

Acknowledgments

I What are Freshwater Habitats?

II Lentic Habitats

III Lotic Habitats

IV Wetland Habitats

V Spring Habitats

VI Subaerial Habitats

Chapter 3: Coccoid Cyanobacteria

Abstract

I Introduction

II Morphology and Diversity

III Ecology and Distribution

IV Collection, Preparation, and Culture

V Key and Description of Genera

VI Guide to the Literature for Species Identification

Chapter 4: Filamentous Cyanobacteria

Abstract

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key to North American Genera

VI Guide to Literature for Species Identification

Chapter 5: Red Algae

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key and Descriptions of Genera

VI Guide to the Literature for Species Identification

Chapter 6: Flagellate Green Algae

Abstract

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key to Genera

VI Description of North American Genera

VII A Guide to the Literature for Species Identification

Chapter 7: Nonmotile Coccoid and Colonial Green Algae

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Investigation and Identification

V Key and Descriptions of Genera

VI A Guide to the Literature for Species Identification

Chapter 8: Filamentous (Nonconjugating) and Plantlike Green Algae

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation of Samples

V Key and Descriptions of North American Genera

VI Guide to Literature for Species Identification

Chapter 9: Conjugating Green Algae Including Desmids

Abstract

I Introduction

II Biodiversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key to North American Genera

VI Descriptions of Genera

VII Guide to the Literature for Species Identification

Chapter 10: Photosynthetic Euglenoids

Abstract

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key to North American Genera

VI A Guide to the Literature for Species Identification

Chapter 11: Xanthophyte, Eustigmatophyte, and Raphidophyte Algae

Abstract

Acknowledgments

I General Introduction

II Xanthophytes

III Eustigmatophyceae

IV Keys and Descriptions of Genera of Xanthophytes and Eustigmatophytes

V Raphidophytes

VI Collection and Preparation for Identification

VII Guide to the Literature for Species Identification

Chapter 12: Chrysophyceae and Phaeothamniophyceae

Abstract

Acknowledgments

I Introduction¹

II Diversity and Morphology

III Ecology

IV Collection and Preparation for Identification

V Key and Descriptions of Genera

VI Guide to Literature for Species Identification

Chapter 13: Haptophyte Algae

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key and Descriptions of Genera

VI Guide to the Literature for Species Identification

Chapter 14: Synurophyte Algae*

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Keys to Genera and Common Species Found in North America

VI A Guide to the Literature for Species Identification & Museum Collections

Chapter 15: Centric and Araphid Diatoms

Abstract

Acknowledgments

I General Introduction to the Diatoms

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key and Descriptions of Genera

VI Descriptions of Genera

VII Genera

VIII Freshwater Araphid Diatoms

IX Guide to Literature for Species Identification

Chapter 16: Bacillariophyceae: The Raphid Diatoms

Abstract

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Keys and Description of Genera

VI Description of Genera

VII Guide to Literature for Species Identification

VIII Guide to the Literature of Species of Bacillariales, Rhopalodiales, and Surirellales

Chapter 17: Dinoflagellates

Abstract

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Collection and Preparation for Identification

V Key to North American Genera

VI A Guide to the Literature for Species Identification

Chapter 18: Cryptomonads

Abstract

I Introduction

II Ultrastructure and Morphology (Figures 1 and 2)

III Origin of Cryptomonads

IV Ecology and Distribution

V Collection, Isolation, and Culturing

VI Classification and Key (Figures 3–17)

VII Descriptions of Genera

VIII Availability of Cryptomonads

IX Phylum Kathablepharida (Figures 18–20)

Chapter 19: Brown Algae

Abstract

Acknowledgments

I Introduction

II Diversity and Morphology

III Ecology and Distribution

IV Methods for Collection and Identification

V Key and Description of Genera

VI Guide to Literature for Species Identification

Chapter 20: Harmful Algal Blooms

Abstract

Acknowledgments

I Introduction and Overview

II Planktonic Blooms

III Benthic HABs

IV Chemical Ecology of HABs (Semiochemicals)

V Quantifying, Monitoring, Modeling, and Managing HABs

Chapter 21: Use of Algae in Ecological Assessments

Abstract

Acknowledgments

I Introduction

II A Framework for Ecological Assessment

III Sampling Algae in Freshwater Habitats

IV Characterizing Attributes of Algal Assemblages

V Characterizing Condition

VI Diagnosing Stressors

VII Management Decisions

VIII Conclusions

Glossary

Author Index

Subject Index

Taxonomic Index

Copyright

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Notices

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Dedication

We dedicate this volume to the phycologists and aquatic scientists who have passed away since the publication of our first edition in 2003. They were our teachers, colleagues, and friends—remarkable people who guided and inspired us. They are missed:

Kostas Anagnostidis, Tony Bailey-Watts, Hilda Canter-Lund, Kathleen M. Cole, Lebaron C. Colt, Jr., David B. Czarnecki, Thamarapu V. Desikachary, Michael R. Droop, Gordon E. Fogg, John C. Kingston, Paul Kugrens, John W. G. Lund, Francis Magne, Jack L. McLachlan, Richard E. Norris, Murray J. Parsons, Ruth Patrick, Louisa P. Perestenko, Charles W. Reimer, Frank E. Round, Paul C. Silva, Eugene F. Stoermer, Francis R. Trainor, Richard A. Vollenweider, Robert G. Wetzel, and Hugh B.S. Womersley.

We hope that their contributions, as well as these pages, will inspire new teachers, new colleagues, and new friends.

John D. Wehr, Robert G. Sheath, and J. Patrick Kociolek

Contributors

Numbers in parenthesis indicate the pages on which the authors’ contributions begin.

Bryan W. Brooks   873  Department of Environmental Science, Baylor University, Waco, Texas, USA

Susan Carty   773  Department of Biological and Environmental Sciences, Heidelberg University, Tiffin, Ohio, USA

Brec L. Clay   809  CH Diagnostic & Consulting Service, Inc., Berthoud, Colorado, USA

Thomas Friedl   485  Experimentelle Phykologie und Sammlung von Algenkulturen, Georg-August-Universität Göttingen, Göttingen, Germany

Georg Gärtner   315  Institute of Botany, University of Innsbruck, Innsbruck, Austria

John D. Hall   429  University of Maryland, College Park, Maryland, USA

Scott N. Higgins   873  International Institute for Sustainable Development, Winnipeg, Manitoba, Canada

Jeffrey R. Johansen   75, 135  Department of Biology, John Carroll University, University Heights, Ohio, USA

Department of Botany, Faculty of Science, University of South Bohemia, Branišovská 31, 370 05, České Budějovice, Czech Republic

David M. John   375  Life Sciences Department, The Natural History Museum, London, United Kingdom

Matthew Julius   653  Department of Biological Sciences, St. Cloud State University, St. Cloud, Minnesota, USA

John C. Kingston†   653  Center for Water and the Environment, Natural Resources Research Institute, University of Minnesota Duluth, Ely, Minnesota, USA

† Deceased.

Jiří Komárek   375, 135  Institute of Botany AS CR, Třeboň, and Department of Botany, Faculty of Science, University of South Bohemia, Branišovská 31, 370 05, České Budějovice, Czech Republic

R.L. Lowe 709 University of Michigan Biological Station, Pellston, Michigan, USA

Department of Biological Sciences, Bowling Green State University, Bowling Green, Ohio, USA

Richard M. McCourt   429  Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA

Takashi Nakada   265  Institute for Advanced Biosciences, Keio University, Tsuruoka, Yamagata, Japan

Systems Biology Program, Graduate School of Media and Governance, Keio University, Fujisawa, Kanagawa, Japan

Kenneth H. Nicholls   537, 587  RR #1, Sunderland, Ontario, Canada

Hisayoshi Nozaki   265  Graduate School of Science, University of Tokyo, Bunkyo, Tokyo, Japan

Carla K. Oldham-Ott   485  School for Professional Studies, Walsh University, Akron, Ohio, USA

Donald W. Ott   485  Department of Biology, The University of Akron, Akron, Ohio, USA

Hans W. Paerl   873  University of North Carolina at Chapel Hill, Morehead City, North Carolina, USA

Matthew W. Parrow   773  Department of Biological Sciences, University of North Carolina at Charlotte, Charlotte, North Carolina, USA

J. Patrick Kociolek   653, 709  Department of Ecology and Evolutionary Biology and Museum of Natural History, University of Colorado, Boulder, Colorado, USA

University of Michigan Biological Station, Pellston, Michigan, USA

Fabio Rindi   375  Dipartimento di Scienze della Vita e dell'Ambiente, Università Politecnica delle Marche, Ancona, Italy

Nataliya Rybalka   485  Institut für Mikrobiologie und Genetik, Georg-August-Universität Göttingen, Göttingen, Germany

Experimentelle Phykologie und Sammlung von Algenkulturen, Georg-August-Universität Göttingen, Göttingen, Germany

Robert G. Sheath   1, 13, 237  Department of Biological Sciences, California State University San Marcos, San Marcos, California, USA

Elliot Shubert   315  Faculty of Science and Technology, The University of Westminster, London, UK

Peter A. Siver   607  Botany Department, Connecticut College, New London, Connecticut, USA

John P. Smol   921  Paleoecological Environmental Assessment and Research Laboratory (PEARL), Department of Biology, Queen’s University, Kingston, Ontario, Canada

S.A. Spaulding   709  United States Geological Survey, University of Colorado, Boulder, Colorado, USA

INSTAAR, University of Colorado, Boulder, Colorado, USA

R. Jan Stevenson   921  Department of Zoology, Michigan State University, East Lansing, Michigan, USA

Eugene F. Stoermer†   653  Center for Great Lakes and Aquatic Sciences, University of Michigan, Ann Arbor, Michigan, USA

Edward C. Theriot   653  Department of Integrative Biology, and Texas Memorial Museum, University of Texas at Austin, Austin, Texas, USA

Richard E. Triemer   459  Department of Plant Biology, Michigan State University, East Lansing, Michigan, USA

Morgan L. Vis   237  Department of Environmental and Plant Biology, Ohio University, Athens, Ohio, USA

Sue B. Watson   873  Environment Canada, Canadian Centre for Inland Waters, Burlington, Ontario, Canada

John D. Wehr   1, 13, 851, 873  Louis Calder Center—Biological Station, Fordham University, Armonk, New York, USA

Brian A. Whitton   873  School of Biological and Biomedical Sciences, University of Durham, Durham, UK

David M. Williams   653  Life Sciences Department, Natural History Museum, London, United Kingdom

Daniel E. Wujek   537  epartment of Biology, Central Michigan University, Mt. Pleasant, Michigan, USA

Bożena Zakryś   459  Department of Plant Systematics and Geography, Faculty of Biology, University of Warsaw, Warsaw, Poland

Preface

John D. Wehr; Robert G. Sheath; J. Patrick Kociolek

First Edition

The study of freshwater algae in North America has a long and rich history, with some of the early monographic works dating back to the late 1800s. In recent years, there has been an enormous and remarkable level of research on this very diverse and heterogeneous collection of organisms, making any definitive taxonomic or ecological treatise always out of date. Nonetheless, it is our goal with this book to synthesize and update much of this vast knowledge and to provide a practical and comprehensive guide to all of the genera of freshwater algae known from throughout the continent, in one volume. Chapters also provide guides to other publications and specialized works for the identification and ecological information at the species level. Our intent is to combine the necessary ecological and taxonomic information in a practical book that can be used by all scientists working in aquatic environments, whether their specialty is in environmental monitoring, ecology, evolution, systematics, biodiversity, or molecular biology. This is the first book of its sort covering the entire continent. We also hope that this book will serve to encourage new generations of aquatic biologists to explore freshwater algae carefully, rather than regarding phytoplankton or benthic algae as simply quantities of chlorophyll or carbon. The enormous variety of algae in lakes, rivers, and other aquatic habitats is part of the ecological content of aquatic communities, and their ecosystem functions vary with the species that occur there.

Many of the previous monographs dealing with a broad geographic region are still useful, such as Smith’s (1950) Freshwater Algae of the United States, but most are decades old and do not contain recent taxonomic changes. Our approach is to include chapters authored by experts who have specialized in the study of specific groups of freshwater algae. Given the great quantity of research that has been produced on all of the major algal taxa, it is no longer possible for one or two authors to produce an authoritative book of this kind and one that will span the entire range of taxonomic and ecological detail that is now known about all the organisms termed algae. This volume is modeled closely after the book by Thorp and Covich on freshwater invertebrates (Ecology and Classification of North American Freshwater Invertebrates), also published by Academic Press.

The organization of this book includes an introduction to the freshwater algae (with a guide to the taxonomic chapters that follow), an overview of freshwater habitats, 20 taxonomic chapters, and finally chapters on the use of algae in environmental assessments and control of nuisance algae. More than 770 genera are described and illustrated in this book, and each taxonomic chapter includes an introduction to the key terms and characteristics of the group, ecological distribution, and a guide to the taxonomic literature to distinguish species within each genus. While we have undoubtedly omitted some less common or yet unrecorded freshwater genera, this compilation represents an increase in the taxonomic scope and geographic coverage of the freshwater algae of North America. This compares with roughly 490 genera recorded from the United States by Smith (1950), about 335 in Prescott’s (1962) coverage of the Western Great Lakes region, and nearly 380 genera from the southeastern United States reported by Whitford and Schumacher (1984). Because not all algal groups are equally well studied, coverage in the present volume varies among taxa and chapters. We hope that students, scientists working in water management agencies, and experienced phycologists will use this book thoroughly and provide us with feedback, such as missing taxa or incomplete geographic information. We will endeavor to incorporate this information into a future edition.

Second Edition

The second edition of Freshwater Algae of North America occurs at a time of rapid and exciting change in the fields of phycology and aquatic ecology. The development and widespread use of molecular tools in the intervening years have provided remarkable insights into the evolution, classification, and ecology of algae. Genetic data have been used to clarify phylogenetic relationships among algal groups, to discern how marine species may have evolved to colonize freshwater environments, to understand gene function as cells respond to their environment, and to uncover far greater diversity than was imagined. The challenge for algal taxonomists is this: How do we reconcile new genetic data with traditional species concepts based on morphology and reproduction? Fortunately, some studies have found agreements between gene-based and older species concepts. Notably, this includes names dating back to Linnaeus (Hayden et al., 2003). In other cases, the discovery of cryptic and, in some cases, endemic species not previously recognized using microscopy has both expanded our knowledge of the freshwater algal flora and created challenges for morpho-taxonomic work (Boo et al., 2010; Guiry, 2012; Komárek and Mares, 2012; Mann and Vanormelingen, 2013). We should also note that through these studies a few genera reported in the first edition are no longer valid and have either been renamed or merged with other taxa. However, many, perhaps most, genera have yet to be studied in any detail by molecular methods.

It has been estimated that more than half of the cyanobacteria species on earth remain to be described by any method (Nabout et al., 2013) and that only 10% of all diatoms species have yet been named (Guiry, 2012). Discoveries of new species and genera will continue. There are some that have been reported from just a few lakes or a single pool, or their true identity has not yet been confirmed. Other taxa are regarded as uncertain or doubtful and await further study and molecular analysis. We contend that these are healthy developments for our science. This volume, while still based largely on morpho-species, has nonetheless expanded from the first edition in part due to the inclusion of taxa that have been recognized through these new methods. Also, several authors have described genera (and species) that have been observed on other continents, but expected to occur in North America, leaving the door open for further additions to our flora. Many of new genera are described in this volume, some the result of reclassifications, others from discoveries of taxa not previously observed or known. We predict that this growth will continue and that further links between traditional and molecular approaches will be created. The present volume now describes 922 genera, an increase of more than 150 from the first edition.

On the whole, readers of our first edition have been very complimentary and generous in their feedback and advice. A student carrying a well-worn book is always a welcome sign. One frequent suggestion was to include color images. This is an understandable request we are pleased to provide, given that pigments are important in characterizing different algal groups and that these colors highlight the beauty and diversity that encompass the algae. The organization of this book is similar to that of the first edition, but has been reorganized in recognition of some of the new classification schemes that have emerged in recent years. The result is 17 taxonomic chapters, preceded by Chapter 1, a general introduction to these chapters, and Chapter 2, an overview of freshwater algal habitats. We still employ common names for the chapters, but also provide current phylogenetic information. The final two chapters focus on two important applied topics in algal research; bioassessment methods and harmful algal blooms in fresh waters. Feedback from all users of this edition is warmly encouraged.

We are again exceedingly indebted to the contributors to this volume. Their care and expertise are evident in these pages. We also offer our sincere thanks to all of the external reviewers who have provided invaluable comments for our authors: Robert A. Andersen, António J. Calado, Peter F. M. Coesel, Gertrud Cronberg, Walter K. Dodds, Karolina Fučíková, Thomas Friedl, Jennifer L. Graham, Theodore D. Harris, David M. John, Ric W. Jordan, J. Patrick Kociolek, Lothar Krienitz, Jørgen Kristiansen, Brian S. Leander, Robert E. Lee, Louise A. Lewis, Eric W. Linton, Rex L. Lowe, Richard M. McCourt, Kirsten M. Müller, Orlando Necchi, Jr., Phil M. Novis, Karin Rengefors, Samuel R. Rushforth, Robert G Sheath, Alison R. Sherwood, and Alan D. Steinman.

Literature Cited

Boo SM, Kim HS, Shin W, Boo GH, Cho SM, Jo BY, Kim J-H, Kim JH, Yang EC, Siver PA, et al. Complex phylogeographic patterns in the freshwater alga Synura provide new insights into ubiquity vs. endemism in microbial eukaryotes. Mol. Ecol. 2010;19:4328–4338.

Guiry MD. How many species of algae are there? J. Phycol. 2012;48:1057–1063.

Hayden HS, Blomster J, Maggs CA, Silva PC, Stanhope MJ, Waaland JR. Linnaeus was right all along: Ulva and Enteromorpha are not distinct genera. Eur. J. Phycol. 2003;38:277–294.

Komárek J, Mares J. An update to modern taxonomy (2011) of freshwater planktic heterocytous cyanobacteria. Hydrobiologia. 2012;698:327–351.

Mann DG, Vanormelingen F. An inordinate fondness? The number, distributions, and origins of diatom species. J. Eukaryot. Microbiol. 2013;60:414–420.

Nabout JC, da Silva Rocha B., Carneiro FM, Sant’Anna CL. How many species of Cyanobacteria are there? Using a discovery curve to predict the species number. Biodivers. Conserv. 2013;22:2907–2918.

Prescott GW. Algae of the Western Great Lakes Area. second ed. Dubuque, Iowa: W.C. Brown; 1962.

Smith GM. The Fresh-water Algae of the United States. second ed. New York: McGraw-Hill; 1950.

Whitford LA, Schumacher GJ. A Manual of Fresh-Water Algae. Revised ed. Raleigh, NC: Sparks Press; 1984.

Chapter 1

Introduction to the Freshwater Algae

Robert G. Sheath¹; John D. Wehr²    ¹ Department of Biological Sciences, California State University San Marcos, San Marcos, California, USA

² Louis Calder Center—Biological Station, Fordham University, Armonk, New York, USA

Abstract

This chapter briefly introduces the diverse morphologies of freshwater algae, explaining seven basic types, which range from unicells to colonies (which may or may not be flagellated), pseudofilaments to true filaments (with or without branching), pseudoparenchymatous forms to true tissues, and coenocytic structures. Photographs of each type of morphology are provided. The chapter then provides a key to the taxonomic chapters in the book with an accompanying table of distinguishing features. The groups of freshwater algae are briefly described based on the taxonomic organization of the book.

Keywords

algae

classification

diversity

freshwater

identification

morphology

Chapter Contents

I. Introduction   1

II. Classification   5

A. Key to the Taxonomic Chapters in This Book   5

III. Groups of Freshwater Algae   7

A. Cyanobacteria   7

B. Red Algae   7

C. Green Algae   8

D. Euglenoids   8

E. Eustigmatophyte, Raphidiophyte, and Xanthophyte Algae   8

F. Chrysophycean Algae   8

G. Haptophyte Algae   9

H. Synurophyte Algae   9

I. Diatoms   9

J. Dinoflagellates   9

K. Cryptomonads   10

L. Brown Algae   10

Literature Cited   10

I Introduction

Freshwater algae are globally ubiquitous and highly diverse, with tens or perhaps hundreds of thousands of species, in a myriad of forms and sizes (Andersen, 1992; Norton et al.; 2004; Mann and Vanormelingen, 2013; Guiry and Guiry, 2014). The algae represent between eight and 12 major evolutionary lineages (Graham et al., 2008; Cock et al., 2010), and all have representatives in inland waters. Current classifications consider most algae to be protists with chloroplasts, but there are also photosynthetic prokaryotes (the cyanobacteria) and a subset of the land plants, the Charales, which have been considered to be algae in previous texts (Patterson, 2014). With new molecular tools being applied to understanding algal taxonomy, systematics, and evolution, our understanding of this diversity is rapidly changing and expanding. Efforts to characterize this biological diversity (such as the Tree of Life Project: Maddison et al., 2007) have contributed to a better understanding of many groups of freshwater algae (e.g., Lane and Archibald, 2008; Entwisle et al., 2009; Hall et al., 2010; Ashworth et al., 2013; Stancheva et al., 2013a; Fučíková et al., 2014).

For simplicity, algae are treated in this book in the same sense as they are in recent phycology texts (e.g., Graham et al., 2008; Lee, 2008); that is, they are considered to be a loose (polyphyletic) group of organisms that have all or most of the following characteristics: aquatic, photosynthetic (possessing chlorophyll a), simple vegetative structures without a vascular system, and reproductive bodies that lack a sterile layer of protecting cells. There are important exceptions, however, particularly with regard to their habitat (Chapter 2), colorless relatives (lacking photosynthetic pigments; Chapters 10, 12, and 18), and species that switch between photosynthesis and predation on bacteria (mixotrophy; Chapters 10, 12, and 17). Both prokaryotic (those with cells lacking membrane-bound organelles) and eukaryotic taxa (cells with organelles) are considered in this book. Within the algae, there is an enormous range of vegetative morphologies, including the following:

1. Unicells: Species that occur as solitary cells that may be non-motile or motile, the latter of which move by using one or more flagella or gliding along surfaces via mucilage or other means. A wide variety of forms exist among unicells, including those contained within a gelatinous sheath (Fig. 1A); cells with intricate shapes, walls, and markings (Figs. 1B and C); cells having flexible cell shapes (Fig. 1D); and cells with two flagella of equal length (Fig. 1E) or flagella with unequal length (Fig. 1F). Cells can be drawn out into hornlike projections (Fig. 1G) or are contained in a hardened case or lorica (Fig. 1H).

Figure 1 Unicellular and colonial forms of freshwater algae. A. Chroococcus giganteus (coccoid cyanobacterium), a unicell to small grouping of cells (regarded as colonial) contained within concentrically layered gelatinous sheaths. B. Micrasterias americana (conjugating green alga, desmid), a unicell with a deep incision (isthmus) that divides two semicells (clear central area is the nucleus), each of which has several deep incisions that may be subdivided into a series of lobes and lobules. C. Didymosphenia geminata (pennate diatom), a large unicell whose walls are constructed from silica, with a central raphe slit, and marked by a series of intricate pores arranged in lines (striae). D. Euglena sp. (photosynthetic euglenoid), a uniflagellate unicell with multiple green chloroplasts, red eyespot, and exterior covering composed of a series of proteinaceous strips (pellicle) that can readily change shape. E. Dunaliella sp. (flagellated green alga) a unicell with two equal flagella. F. Ochromonas sp. (chrysophycean alga), a golden unicellular alga with two apically inserted flagella, one long (tinsel) and one short (smooth). G. Ceratium hirundinella (dinoflagellate), a unicell with a theca composed of cellulose plates with cellular extensions or horns and a transverse flagellum and trailing longitudinal flagellum. H. Lagynion sp. (chrysophycean alga) a golden rhizopodial unicell within a rigid lorica. I. Woronichinia naegeliana (coccoid cyanobacterium), a colony with spherical or ovoid cells arranged at the periphery of a gelatinous matrix. J. Tetraspora lubrica (nonmotile colonial green alga), a sac-like colony of irregularly arranged special cells within a gelatinous matrix. K. Dictyosphaerium granulatum (non-motile colonial green alga), a colony with spherical cells attached together by gelatinous strands. L. Oocystis lacustris (non-motile colonial green alga), a colony with groups of four to eight cells produced inside the walls of parent cells. Scale bars = 25 μm, except E, F, H = 10 μm; B, C, G = 50 μm. (Photos A, D, K, L courtesy of Chris Carter, with permission, photo F courtesy of Peter A. Siver, Chrysophytes LLC, with permission).

2. Colonies: An aggregation of cells that are held together either in a loose (Fig. 1I and J) or tight well-organized pattern (Figs. 1K-L, 2A-C). Depending on the algal taxon, colonies may contain a variable number of cells, or they may be constant throughout their development (coenobium). Colonies may be composed of flagellated or non-flagellated cells. The basis for cellular connection varies among colonies, including a surrounding gelatinous matrix (Fig. 1I and J), gelatinous stalks (Fig. 1K), or a common parental wall (Fig. 1L). Cells may be arranged in a highly organized pattern (Fig. 2A), directly attached at their cellular edges (Fig. 2B), or at the middle portion of each cell (Fig. 2C). Alternatively, cells may be connected to others by their loricas (Fig. 2E).

Figure 2 Colonial, pseudofilamentous, and filamentous forms of freshwater algae. A. Merismopedia sp. (coccoid cyanobacterium), a colony of numerous cells arranged in rectangular rows within a flattened colony. B. Tabellaria flocculosa (pennate diatom), a colony with rectangular cells seen in side (girdle) view, attached at their edges by mucilage pads in a zigzag fashion. C. Fragilaria crotonensis (pennate diatom), a ribbon-like colony of linear cells attached only at the central region by interlocking spines. D. Dinobryon divergens (chrysophycean alga), a colony of golden flagellates contained within individual funnel-shaped loricas. E. Stichosiphon willei (coccoid cyanobacterium), a series of individual cells contained within a common mucilaginous sheath, forming a pseudofilament. F. Chroodactylon sp. (red alga), a pseudofilament with cells distantly arranged in an end-to-end pattern in a common gelatinous matrix, but not directly connected to each other. G. Chamaesiphon sp. (coccoid cyanobacterium) forming patches of brown crusts on a rock from a fast-flowing stream. H. Zygnema sp. (conjugating green alga), an unbranched filament without a gelatinous matrix. I. Limnoraphis sp. (filamentous cyanobacterium), an unbranched filament composed of short, discoid cells contained within a gelatinous sheath. J. Scytonema sp. (filamentous cyanobacterium), a filament within a common sheath that produces double false branches that result from breakage and further growth of each fragment. K. Bangia sp. (red alga), an unbranched, multiseriate filament with two to many cells across embedded within a firm gelatinous matrix. Scale bars = 25 μm, except C, D, E = 10 μm; G = 1 cm.

3. Pseudofilaments: An aggregation of cells in an end-to-end fashion. The cells are not directly connected to each other; rather, they are spaced apart and may be contained within a common gelatinous matrix or sheath (Figs. 2E and F). A few pseudofilamentous cyanobacteria form macroscopically recognizable crusts on rocks (Fig. 2G).

4. Filaments: A chain or series of cells in which the cells are arranged end to end, with adjacent cells sharing a common cross wall (Figs. 2H-J, 3B and C). Traditionally, linear colonies can be distinguished from true filaments by the fact that abutting colonial cells each possess their own entire walls (e.g., Fig. 2F), while true filaments do not (Fig. 2H). However, some chain-forming diatoms, such as Melosira, may also be referred to as filaments (Chapter 15). Filaments may be arranged in a single series (uniseriate or uniaxial) (Fig. 2H-J) or in more than one series of cells (termed multiseriate or multiaxial) (Fig. 2K). Filaments may be unbranched (Figs. 2H and I) or may produce false branches (Fig. 2J). False branches, which occur in some cyanobacteria, such as Scytonema (Fig. 2J), are formed from breaks in the trichome within a common sheath and differ from true branches because they do not form via direct cell divisions. In some filaments, the main axes produce branches in a new plane that are morphologically similar to the main axis (Fig. 3A), short side branches (Fig. 3B), or branches that are heterotrichous; that is, they have a distinct, often larger primary axis with smaller branches (Figs. 3C, D). Branching may be dichotomous (Fig. 3A), alternate (Fig. 3B), opposite (Fig. 3C), or whorled (Fig. 3C). Differentiated filaments may also have reproductive cells, or in some cyanobacteria, physiologically specialized cells specialized for N-fixation, termed heterocytes (Figs. 2I, 3E; Chapter 4).

Figure 3 Filamentous, saclike, crustose, pseudoparenchymatous, and siphonous forms of freshwater algae. A. Cladophora glomerata (filamentous green alga), a filament with dichotomous (forked), equal-diameter branches, and cells containing parietal net-like chloroplasts. B. Batrachospermum macrosporum (red alga), the alternately branched chantransia stage with equal-diameter, but shorter side branches. C. Batrachospermum helminthosum (red alga), a filament with a corticated central axis and whorled lateral branches (flattened by coverslip). D. Draparnaldia acuta (filamentous green alga), a filament that has tuft-like lateral branches with cells that are considerably smaller than those of the main axis and taper to fine points. E. Stigonema mammilosum (filamentous cyanobacterium), a multiseriate filament with cells diving in all three planes, including specialized cells for nitrogen fixation (heterocytes). F.  Lemanea sp. (red alga), a pseudoparenchymatous thallus of closely arranged cells forming a cartilaginous tube surrounding a central uniseriate axis. G.  Gongrosira fluminensis (filamentous green alga), a pseudoparenchymatous cushion or crust formed from a series of erect and prostrate sparsely branched filaments. H.  Heribaudiella sp. (brown alga), a pseudoparenchymatous crust consisting of a densely arranged basal system and tightly packed, erect filaments adherent to rock substrata. I.  Ulva flexuosa (filamentous and plant-like alga), a parenchymatous alga forming large macroscopic tubes or sacs. J.  Ulva flexuosa detail of cortical cells on outer surface. K.  Vaucheria geminata (xanthophyte, yellow-green alga), a siphonous thallus without cross walls separating the nuclei and multiple chloroplasts. Scale bars = 25 μm, except D, G, K = 50 μm; C, F = 100 μm, I = 5 cm. (Photos J, K courtesy of Chris Carter, with permission).

5. Pseudoparenchymatous structures: Tissue-like thalli that consist of closely appressed branches of a uniseriate or multiseriate filament (Figs. 3F-I). In the red alga Lemanea, the main axis has a surrounding layer of small cells termed cortication (Fig. 3F). Crustose morphologies develop from a series of short, compacted filaments, such as the green alga Gongrosira (Fig. 3G) and the brown alga Heribaudiella (Fig. 3H).

6. Parenchymatous forms: True tissues composed of a solid mass of cells that is three dimensional, variously shaped, and not filamentous or colonial in construction. The cells are often differentiated into an outer photosynthetic layer (cortex) and an inner non-photosynthetic region (medulla). Most tissue-like forms in freshwater habitats are simple tubes or blades, like the tubular or saccate green alga Ulva (formerly Enteromorpha), which consists of a single layer of cells (Fig. 3I-J).

7. Coenocytic or siphonous forms: Large multinucleate forms of various shapes without cross walls to separate the nuclei or other organelles. An example is the yellow-green alga Vaucheria (Fig. 3K). A cross wall does form during formation of the oogonium, which separates it from the parent filament.

Freshwater algae exhibit all of these morphologies, but the macroscopic pseudoparenchymatous and parenchymatous forms tend to be smaller than those found in marine systems (e.g., Sheath and Hambrook, 1990). In addition, planktonic (floating) forms are typically small and often microscopic and consist mostly of the simpler forms. In contrast, the benthic (attached) algae include the entire range of sizes and morphologies, although flagellated taxa are less common than in the plankton.

II Classification

The algae do not represent a formal taxonomic group of organisms, but rather constitute a heterogeneous collection of phyla or divisions with representatives in several kingdoms and having most of the characteristics described above. The divisions are distinguished from each other based on a combination of characteristics, including photosynthetic pigments, starch-like (or other) reserve products, cell covering, and other aspects of cellular organization (e.g., Graham et al., 2008; Lee, 2008). There is little consensus among phycologists as to the exact number of algal divisions, and the modern methods are likely to change higher classification categories for some years to come.

The key characteristics of algae in the 17 major groups recognized in this book are compared in Table 1. The approach of this book is to break the algal major groups into manageable taxonomic units. Thus, for divisions with many freshwater representatives, there are multiple chapters, while some chapters cover more than one phylum. The following key gives the major characteristics to allow the reader to immediately proceed to the appropriate chapter to determine an unknown algal sample (Table 1).

Table 1

Major Distinguishing Features of the Major Algal Groups Presented in This Booka

a From various phycology textbooks (e.g., Graham et al., 2008; Lee, 2008).

b chl = chlorophyll (green), PE = phycoerythrin (red), PC = phycocyanin (blue), APC = allophycocyanin (blue), fucoxanthin and peridinin (golden to brown).

c All of the reserves are polymers of glucose; they differ by their linkages; cyanophycean and floridean α-1,4 and α-1,6 branches; true with amylose α-1, 4 and amylopectin α-1,4 and α-1,6 branches; paramylon β-1,3; chrysolaminarin and laminarin β-1,3 and β-1,6 branches. Only true starch stains positively with iodine (purple to black).

d Pellicle and periplast within plasma membrane; the rest are external to it.

A Key to the Taxonomic Chapters in This Book

1. Cells with no chloroplast or other membrane-bound organelles; typically blue-green colored throughout (occasionally reddish, purple, brown or olive)....................................................................................................................................2

1. Cells with variously colored pigments localized in one or more chloroplasts....................................................................................................................................3

2. Organisms unicellular or colonial, or occasionally forming short chains (coccoid or colonial cyanobacteria)....................................................................................................................................................................Chapter 3

2. Organisms filamentous (filamentous cyanobacteria...............................................................................................................Chapter 4

3. Cells stain positively (purple to black) with iodine for true starch...........................................4

3. Cells do not stain positively (orange or yellow) with iodine for starch.............................................................................................................9

4. Green-colored chloroplasts with chlorophylls a and b as predominant photosynthetic pigments.............................................................................................................5

4. Chloroplasts with other colors and predominant photosynthetic pigments........................................................................................................................................8

5. Organisms flagellated in the vegetative state (flagellated green algae)........................................................................................................................................Chapter 6

5. Organisms non-flagellated in the vegetative stage6

6. Organisms coccoid or colonial, not reproducing via conjugation (coccoid and colonial non-motile green algae)........................................................................................................................................Chapter 7

6. Organisms filamentous, plant-like or with sexual reproduction by conjugation........................................................................7

7. Organisms with filamentous or plant-like morphologies without conjugation (filamentous and plant-like green algae........................................................................................................................................Chapter 8

7. Organisms with unicellular (various shapes) or filamentous morphologies with conjugation (conjugating green algae filaments and desmids........................................................................................................................................Chapter 9

8. Cells with golden or brown chloroplasts with peridinin as the predominant photosynthetic pigment; two flagella, one transverse, the other posterior (dinoflagellates)........................................................................................................................................Chapter 17

8. Cells with blue-, brown-or red-colored chloroplasts with either phycocyanin or phycoerythrin as the predominant photosynthetic pigment; flagella subapical (cryptophyte algae)........................................................................................................................................Chapter 18

9. Cells with blue-colored or red-colored chloroplasts with phycocyanin or phycoerythrin as the predominant photosynthetic pigment (red algae)........................................................................................................................................Chapter 5

9. Cells green or golden-colored with other predominant photosynthetic pigments........................................................................................................................................10

10. Cells green-colored; with a pellicle (layer below the plasma membrane that is proteinaceous, interlocking, often spiral strips) on motile cells (euglenoids)........................................................................................................................................Chapter 10

10. Cells yellow-green or golden colored; naked or walled cells without a pellicle........................................................................................................................................11

11. Yellow-green colored chloroplasts with chlorophylls a and c as the predominant photosynthetic pigments (xanthophytes and related groups)........................................................................................................................................Chapter 11

11. Golden-colored chloroplasts with fucoxanthin as the predominant photosynthetic pigment....................................................................12

12. Cells with a silica frustule covering (diatoms)........................................................................................................................................13

12. Cells with no covering or with one that is not a siliceous frustule (may have siliceous scales)........................................................................................................................................14

13. Diatom frustules bilaterally symmetrical with a raphe system; many times evident along the central axis in valve view, but sometimes occurring along the periphery of the valve face or only on the edge or mantle (raphid diatoms)........................................................................................................................................Chapter 16

13. Diatom frustules bilaterally symmetrical, but lacking a raphe system, or radially symmetrical (araphid pennate and centric diatoms)........................................................................................................................................Chapter 15

14. Cells with two flagella plus a central, specialized appendage, the haptonema, in vegetative and/or reproductive stages (haptophyte algae)Chapter 13

14. Cells without a haptonema in any stage........................................................................................................................................15

15. Vegetative cells with siliceous scales and/or spines (synurophyte algae)........................................................................................................................................Chapter 14

15. Vegetative cells without siliceous scales........................................................................................................................................16

16. Cell walls with alginates; benthic, often macroscopic thalli with no unicellular or colonial representatives; brown filaments or crusts (brown algae)........................................................................................................................................Chapter 19

16. Cell walls without alginates; unicellular or colonial representatives, mostly microscopic thalli (chrysophyte algae)........................................................................................................................................Chapter 12

III Groups of Freshwater Algae

A Cyanobacteria

The cyanobacteria or blue-green algae are prokaryotes without membrane-bound organelles (Table 1; Chapter 3). Other characteristics of this division include unstacked thylakoids, phycobiliprotein pigments, cyanophycean starch, and peptidoglycan matrices or walls. There are 174 genera reported from inland habitats in North America, of which 66 are unicellular, colonial, or pseudofilamentous (Chapter 3) and 108 are filamentous (Chapter 4). However, the taxonomy of this division is currently being revised (Komárek, 2013, Chapter 3), and the number of genera should be considered tentative.

Cyanobacteria inhabit the widest variety of freshwater habitats on Earth and can become important in surface blooms in nutrient-rich standing waters (Chapters 2-4 and 20). However, nitrogen-fixing taxa have been shown to be good indicators of N-limited sites, such as streams in California (Stancheva et al., 2013b), as well as in many lakes and reservoirs (Howarth et al., 1988). Some of the cyanobacterial blooms can be toxic to zooplankton and fish, as well as livestock that drink water containing these organisms. Inland cyanobacteria also occur in extreme environments, such as hot springs, saline lakes, and endolithic desert soils and rocks (Chapters 2 and 20).

B Red Algae

Rhodophyta or red algae represent a division that is characterized by chloroplasts that have no external endoplasmic reticulum and unstacked thylakoids, phycobiliprotein pigments, floridean starch, and lack of flagella in all stages (Table 1; Chapter 5). They are predominantly marine in distribution with fewer than 3% of more than 6500 species occurring in truly freshwater habitats (Guiry and Guiry, 2014; Guiry et al., 2014). In North America, 26 genera are recognized in inland habitats (Chapter 5).

Freshwater red algae are largely restricted to streams and rivers but also can occur in other inland habitats, such as lakes, hot springs, soils, caves, and even sloth hair (Chapter 5). While many species are indicators of good water quality, a small number, like the chantransia stage of Batrachospermum (formerly Chantransia) macrospora, which can be invasive in relatively polluted waters, due to human activities such as dumping aquarium contents (Kato et al., 2009).

C Green Algae

Chlorophyta and Charophyta, the green algae constitute divisions that have the following set of attributes: chloroplasts with no external endoplasmic reticulum, thylakoids typically in stacks of two to six, chlorophylls a and b as photosynthetic pigments, true starch, and cellulosic walls or scales (Table 1). This is a diverse group in inland habitats of North America that includes 48 flagellated genera (Chapter 6), at least 129 coccoid and non-motile colonies (Chapter 7), 81 filamentous and plantlike genera (Chapter 8), and 49 conjugating genera and desmids (Chapter 9). Some members of the green algae (Charophyeae) are part of a lineage that is ancestral to higher plants.

Green algae are widespread in inland habitats, but certain groups may have specific ecological requirements. For example, flagellated chlorophytes tend to be more abundant in standing waters that are nutrient rich (Chapter 6). Coccoid unicells and colonies are common in the plankton of standing waters and slowly moving rivers when nutrients, light, and temperature are reasonably high (Chapters 2, 7). The majority of filamentous and plantlike Chlorophyta are attached to hard surfaces in standing or flowing waters, but some are free-floating or colonize soils or other subaerial habitats (Chapter 8). Some of these species are quite tolerant of desiccation stress (Holzinger and Karsten, 2013). Filamentous conjugating green algae are most frequent in stagnant waters of roadside ditches and ponds and in the littoral zones of lakes, where they can form free-floating mats or intermingle with other algae in attached or floating masses (Chapter 9). Desmids are more common in ponds and streams that have low conductance and moderate nutrient levels and often intermingle with macrophytes and filamentous algae.

D Euglenoids

Photosynthetic Euglenophyta or euglenoids have chloroplasts surrounded by three membranes, thylakoids in stacks of three, chlorophyll a and b as photosynthetic pigments, the ability to store paramylon rather than starch, and a proteinaceous (often spirally arranged) covering (termed a pellicle) (Table 1; Chapter 10). Twelve genera are reported from North American freshwater habitats (Chapter 10). Euglenoids are particularly abundant in the plankton of standing waters rich in nutrients and organic matter, and they can be associated with sediments, fringing higher plants, and leaf litter, although some may dominate in highly acidic environments (Chapters 2 and 10). Some species, like Euglena mutabilis, are pioneering species in severely acid mine drainage-impacted sites with very acidic waters (pH < 3) and elevated metals (Al, Cd, Cu, Fe, Ni, Zn) (Olaveson and Nalewajko, 2000).

E Eustigmatophyte, Raphidiophyte, and Xanthophyte Algae

Eustigmatophyte, raphidiophyte, and xanthophyte algae comprise a loose group of algae that share the following characteristics: chloroplasts with four surrounding membranes, thylakoids in stacks of three, chlorophyll a and c as the typical photosynthetic pigments, and chrysolaminarin as the photosynthetic reserve product (where known) (Table 1; Chapter 11). The yellow-green algae are quite diverse in freshwater habitats of North America: At least 90 genera have been reported. The widespread coenocyte genus Vaucheria can form dense mats in streams when nitrogen and phosphorus concentrations exceed a threshold level (Stevenson et al., 2007), but they also are common on moist soils. In contrast, the eustigmatophytes and raphidiophytes are relatively small groups that comprise eight and three genera, respectively (Chapter 11). Many of these genera seldom have been reported, although the flagellate Gonyostomum is a common component of acidic bog ponds and has created blooms in some boreal lakes (Trigal et al., 2013). Members of this group of algae have been collected from a wide variety of habitats, but many are collected primarily in northern habitats (Chapter 11). They are both planktonic and associated with a variety of substrata.

F Chrysophycean Algae

Chrysophyceae or chrysomonads are distinguished by chloroplasts that have four surrounding membranes, thylakoids in stacks of three, fucoxanthin that typically masks chlorophyll a and c, and chrysolaminarin as the photosynthetic reserve. At least 80 genera are reported from inland habitats of North America (Chapter 12). Chrysophycean algae are typically associated with standing bodies of water that have low or moderate nutrients, low alkalinity and specific conductance, and a pH that is slightly acidic to neutral (Chapter 12). The majority of genera tend to be planktonic. Notably, some species of the widespread planktonic genus Dinobryon are mixotrophic, supplementing photoautotrophy with phagotrophy of bacteria (Heinze et al., 2013). A few benthic forms, such as Hydrurus, form mucilaginous macroscopic thalli (Chapter 12).

G Haptophyte Algae

Members of the Haptophyceae are characterized by chloroplasts that have four surrounding membranes, thylakoids in stacks of three, fucoxanthin that masks chlorophyll a and c, chrysolaminarin as the photosynthetic reserve, and a unique appendage associated with the flagellar apparatus, the haptonema (Table 1; Chapter 13). Currently, nine genera have been reported from freshwater or other inland water bodies in North America (Chapter 13).

The most common freshwater genus, Chrysochromulina, is fairly frequently reported in samples of phytoplankton from lakes and ponds, and it occasionally forms predominant blooms (Chapter 13). For example, Chrysochromulina parva was frequently encountered in oligotrophic softwater lakes in the Ontario Precambrian Shield region (Nicholls et al., 1992) and is a common member of the nanoplankton in Grasmere, a mesotrophic system in the English Lake District (Reynolds et al., 2012).

H Synurophyte Algae

The Synurophyceae is characterized by chloroplasts that have four surrounding membranes, thylakoids in stacks of three, fucoxanthin that masks chlorophyll a and c, chrysolaminarin as the photosynthetic reserve product, and siliceous scales (Table 1; Chapter 14). Three genera are found in North American freshwater habitats (a fourth is known only from Australia), but the genera Mallomonas and Synura are species-rich (Chapter 14).

Synurophytes are exclusively freshwater phytoplankters in lakes, ponds, and slowly flowing rivers (Chapter 14). Habitats that support the largest flora are slightly acidic, low in conductance, alkalinity, and nutrients, and containing moderate amounts of humic substances. The ornamentation of the scales is diagnostic for species identifications, but their overall shape can change as a consequence of rising temperature (Pichrtová and Němcová, 2013).

I Diatoms

Bacillariophyceae or diatoms are distinguished by chloroplasts that have four surrounding membranes, thylakoids in stacks of three, fucoxanthin that masks chlorophyll a and c, chrysolaminarin as the photosynthetic reserve product, and a siliceous frustule (or theca) that comprises the external covering (Table 1; Chapter 15). The diatoms are a complex and diverse group in terms of frustule morphology, with centric forms symmetrical about a point, and most pennate forms symmetrical about a line. The North American freshwater genera currently consist of 25 centric and araphid genera (Chapter 15), and 105 raphid, pennate genera (Chapter 16).

Diatoms are found in all freshwater habitats, including standing and flowing waters and planktonic and benthic habitats. They can often dominate both the biomass and biological diversity of the microscopic flora in many aquatic ecosystems. Because diatoms inhabit a broad array of habitats (but many have specific habitat requirements), they have been used in freshwater environment bioassessment and to monitor long-term changes in ecological conditions (Chapter 21). Some species occasionally become nuisance organisms, such as the stream periphyton diatom Didymosphenia geminata, which may impact freshwater fish, aquatic plants, and insects, causing disturbances to benthic in food webs (Blanco and Ector, 2009, Chapter 20).

J Dinoflagellates

The Pyrrhophyta or dinoflagellates are characterized by chloroplasts that have three surrounding membranes, thylakoids in stacks of three, peridinin that masks chlorophyll a and c, true starch, a nucleus that has condensed chromosomes in cell cycle phases, a cellulosic thecal covering, and frequently a transverse and posterior flagellum. (Table 1; Chapter 17). There are 37 recognized genera in North American freshwater habitats (Chapter 17).

The dinoflagellates are not often major components of the phytoplankton of lakes and ponds but sometimes form dense blooms, particularly in the presence of high levels of nitrates and phosphates (Chapter 17). Some species are capable of forming toxic freshwater red tides (Lee et al., 2006).

K Cryptomonads

The Cryptophyta, cryptomonads or cryptophyte algae, have chloroplasts that have four surrounding membranes in which a nucleomorph occurs between the outer and inner pairs of membranes, thylakoids in loose pairs, phycocyanin or phycoerythrin that masks chlorophyll a and c, true starch as the photosynthetic reserve, a periplast, and two subapical flagella (Table 1; Chapter 18). There are nine genera reported from the inland waters of North America (Chapter 18).

Cryptomonads are typically planktonic in lakes and ponds and are particularly diverse in temperate regions (Chapter 18). These populations can undergo vertical migration patterns in response to conditions such as temperature changes and grazing pressures (Bicudo et al., 2009).

L Brown Algae

Phaeophyceae or brown algae are distinguished by chloroplasts that have four surrounding membranes, thylakoids in stacks of three, fucoxanthin that masks chlorophyll a and c, laminarin as the photosynthetic reserve, and alginates as the wall matrix component. There are six recognized genera of freshwater brown algae, four of which have been collected from freshwater habitats in North America (Chapter 19).

Brown algae are predominantly marine in distribution; fewer than 1% of the species are from freshwater. The inland species are benthic, either in lakes or streams, and distribution is quite scattered (Chapter 19). Some species are considered to be possible invaders from brackish or marine habitats, such as Pleurocladia lacustris (Wehr et al., 2013), although others, such as Heribaudiella, appear to be strict freshwater taxa.

The identification of freshwater algae based on these broad traits requires patience and persistence. Even experts find this task challenging at times! The keys provided here and in subsequent chapters are aimed to help guide and train students and professionals toward that end. Nonetheless, the great diversity of algae that can be observed in a sample of lake water, or in a collection of periphyton from a stream, will offer the individual researcher a fascinating array of forms and species to examine.

Literature Cited

Andersen RA. Diversity of eukaryotic algae. Biodivers. Conserv. 1992;1:267–292.

Ashworth MP, Nakov T, Theriot EC. Revisiting Ross and Sims (1971): toward a molecular phylogeny of the Biddulphiaceae and Eupodiscaceae (Bacillariophyceae). J. Phycol. 2013;49:1207–1222.

Bicudo CEM, Ferragut C, Massagardi MR. Cryptophyceae population dynamics in an oligo-mesotrophic reservoir (Ninféias pond) in São Paulo, southeast Brazil. Hoehnea. 2009;36:99–111.

Blanco S, Ector L. Distribution, ecology and nuisance effects of the freshwater invasive diatom Didymosphenia geminata (Lyngbye) M. Schmidt: a literature review. Nova Hedwiga. 2009;88:347–422.

Cock JM, Sterck L, Rouzé P, Scornet D, Allen AE, Amoutzias G, Anthouard V, et al. The Ectocarpus genome and the independent evolution of multicellularity in brown algae. Nature. 2010;465:617–621.

Entwisle TJ, Vis ML, Chiasson WB, Necchi Jr. O, Sherwood AR. Systematics of the Batrachospermales – a synthesis. J. Phycol. 2009;45:704–715.

Fučíková K, Lewis PO, Lewis LA. Putting incertae sedis taxa in their place: a proposal for nine new families and three new genera in Sphaeropleales (Chlorophyceae, Chlorophyta). J. Phycol. 2014;50:14–25.

Graham LE, Graham JM, Wilcox LW. Algae. Second ed. San Francisco: Benjamin Cummings; 2008 616 pp plus supplemental material.

Guiry MD, Guiry GM. AlgaeBase. Galway: World-wide electronic publication, National University of Ireland; 2014. http://www.algaebase.org Accessed on June 15, 2014.

Guiry MD, Guiry GM, Morrison L, Rindi L, Miranda SV, Mathieson AC, Parker BC, et al. AlgaeBase: an on-line resource for algae. Cryptogam. Algol. 2014;35:105–115.

Hall JD, Fučíková K, Lo C, Lewis LA, Karol KG. Assessing proposed DNA barcodes in the green algae. Cryptogam. Algol. 2010;31:529–555.

Heinze AW, Truesdale CL, DeVaul SB, Swinden J, Sanders RW. Role of temperature in growth, feeding, and vertical distribution of the mixotrophic chrysophyte Dinobryon. Aquat. Microb. Ecol. 2013;71:155–163.

Holzinger A, Karsten U. Desiccation stress and tolerance in green algae: consequences for ultrastructure, physiological, and molecular mechanisms. Frontiers in Plant Science; 2013.doi:10.3389/fpls.2013.00327.4 22 August 2013,

Howarth RW, Marino R, Cole JJ. Nitrogen fixation in freshwater, estuarine, and marine ecosystems. 2. Biogeochemical controls. Limnol. Oceanogr. 1988;33:688–701.

Kato A, Morita N, Hiratsuka T, Suda S. Recent introduction of a freshwater red alga Chantransia macrospora (Batrachospermales, Rhodophyta) to Okinawa, Japan. Aquat. Invasions. 2009;4:567–574.

Komárek J. 2013. Cyanoprokaryota II. Teil/Part 3: Heterocytous Genera. In: Süßwasserflora von Mitteleuropa, Band 19/31. Springer-Spektrum; 1131 pp.

Lane CE, Archibald JM. The eukaryotic tree of life: endosymbiosis takes its TOL. Trends Ecol. Evol. 2008;23:268–275.

Lee RE. Phycology. Fourth ed. Cambridge, UK: Cambridge University Press; 2008 614 pp.

Lee JJ, Chang SH, Lee JH, Lee JH. Morphology and ecology of Peridinium bipes var. occultatum Lindem. (Dinophyceae) forming freshwater red tides in Korean dam reservoirs. Algae. 2006;21:433–443.

Maddison DR, Schulz K-S, Maddison WP. The Tree of Life web project. Zootaxa. 2007;1668:19–40.

Mann DG, Vanormelingen P. An inordinate fondness? The number, distributions, and origins of diatom species. J. Eukaryot. Microbiol. 2013;60:414–420.

Nicholls KH, Nakamoto L, Keller W. Phytoplankton of Sudbury area lakes (Ontario) and relationships with acidification status. Can. J. Fish. Aquat. Sci. 1992;49:40–51.

Norton TA, Melkonian M, Andersen RA. Algal biodiversity. Phycologia. 2004;35:308–326.

Olaveson MM, Nalewajko C. Effects of acidity on the growth of two Euglena species. Hydrobiologia. 2000;433:39–56.

Patterson DJ. Algae: protists with chloroplasts. Tree of Life web project; 2014. http://tolweb.org/tree/phylogeny.html Accessed on June 14, 2014.

Pichrtová M, Němcová Y. Effect of temperature on size and shape of silica scales in Synura petersenii and Mallomonas tonsurata (Stramenopiles). Hydrobiologia. 2013;673:1–11.

Reynolds CS, Maberly SC, Parker JE, DeVille MM. Forty years of monitoring water quality in Grasmere (English Lake District): separating the effects of enrichment by treated sewage and hydraulic flushing on phytoplankton ecology. Freshw. Ecol. 2012;57:384–399.

Sheath RG, Hambrook JA. Freshwater ecology. In: Cole KM, Sheath RG, eds. Biology of the Red Algae. Cambridge University Press, Cambridge, UK: Cambridge University Press; 1990:423–453.

Stancheva R, Hall JD, McCourt RM, Sheath RG. Identity and phylogenetic placement of Spirogyra species (Zygnematophyceae, Charophyta) from California streams and elsewhere. J. Phycol. 2013a;49:588–607.

Stancheva R, Sheath RG, Read BA, McArthur KD, Schroepfer C, Kociolek JP, Fetscher AE. Nitrogen-fixing cyanobacteria (free-living and diatom endosymbionts): their use in southern California stream bioassessment. Hydrobiologia. 2013b;720:111–127.

Stevenson RJ, Pinowska A, Albertin A, Sickman JO. Ecological condition of algae and nutrients in Florida springs: the synthesis report. Florida: Department of Environmental Protection; 2007.

Trigal C, Hallstana S, Johansson KSL, Johnson RK. Factors affecting occurrence and bloom formation of the nuisance flagellate Gonyostomum semen in boreal lakes. Harmful Algae. 2013;27:60–67.

Wehr JD, Stancheva R, Truhn K, Sheath RG. Discovery of the rare freshwater brown alga Pleurocladia lacustris (Ectocarpales, Phaeophyceae) in California streams. West. N. Am. Nat. 2013;73:148–157.

Chapter 2

Habitats of Freshwater Algae

John D. Wehr¹; Robert G. Sheath²    ¹ Louis Calder Center—Biological Station, Fordham University, Armonk, New York, USA.

² Department of Biological Sciences, California State University San Marcos, San Marcos, California, USA

Abstract

Algae occur in every biome across the globe, and they colonize water bodies on every continent. Freshwater ecosystems encompass a very wide range of chemistry, temperature, and physical conditions, which select for a particular assemblage of species. Some inland waters are quite saline; others are temporary or have very little water at all. One reason for such a broad scope is that algal biologists who examine more traditional freshwater environments also study inland saline lakes, wetlands, hot springs, snow and ice, damp soils, and even the insides of plant tissues. Algal habitats span spatial scales of hundreds of square kilometers down to just a few cubic micrometers. This chapter examines the enormous range of inland aquatic habitats in which algae can be found in North America and offers a sampling of the diversity of algal taxa that occur in these systems.

Keywords

freshwater

algae

habitats

rivers

lakes

streams

wetlands

diversity

Chapter Contents

I. What are Freshwater Habitats?   13

II. Lentic Habitats   14

A. Major Lakes of North America   14

B. Lakes and Ponds   17

C. Small-Scale Lentic Habitats and Microhabitats   20

D. Mutualistic, Commensal, and Symbiotic Habitats in Lakes   21

E. Planktonic Habitats and Assemblages in Lakes   23

F. Benthic Habitats and Assemblages in Lakes   25

G. Wastewater Systems   29

III. Lotic Habitats   30

A. Brief Overview of the Geomorphology of Rivers   30

B. Headwaters and Stony Streams   30

C. Major Rivers of North America   33

D. The River Continuum and Other Models   35

E. Benthic Habitats and Assemblages in Rivers   37

F. Planktonic Habitats and Assemblages in Rivers   40

IV. Wetland Habitats   42

A. Functional Importance of Algae in Wetlands   42

B. Algal Diversity and Production in Freshwater Wetlands   43

V. Spring Habitats   45

A. Thermal Springs   45

B. Acid Springs   46

C. Karst Springs   47

VI. Subaerial Habitats   48

A. Soils   48

B. Epilithic and Endolithic Habitats   49

C. Plants and Animals   50

D. Snow and Ice   52

Acknowledgments   53

Literature Cited   53

Acknowledgments

Thanks are due to Dr. Alan Steinman (Grand Valley State University) and Dr. Walter Dodds (Kansas State University) for very helpful reviews and advice on relevant literature. We also thank Robert Bell, Dean W. Blinn, Chris Carter, Andrew F. Casper, Don Chamberlain, Todd A. Crowl, Deborah A. Donaldson, Ronald W. Hoham, Todd Huspeni, Louise A. Lewis, Juan M. Lopez-Bautista, Louis J. Maher Jr, Craig W. Schneider, Janet R. Stein, James H. Thorp, Kam Truhn, Yuuji Tsukii, and Warwick F. Vincent for the use of photos used in this chapter.

I What are Freshwater Habitats?

Algae occur in nearly all surface water bodies in every biome across the globe. Many of these environments are freshwater ecosystems. So what is fresh water? The distinction between freshwater and marine systems is not clear-cut. Oceans are clearly saline (~ 35 g of salts L− 1) and most lakes relatively dilute (< 100 mg L− 1) (Wetzel, 2001), yet there is enormous variation in physicochemical conditions that freshwater algae occupy. Lakes and rivers vary widely in pH, nutrients, trace elements, dissolved oxygen, buffering capacity, redox, and salinity, with some lakes having greater total salts than the open ocean. Limnologists also use the term inland waters to encompass a greater range of aquatic ecosystems. Yet algae occupy many other habitats, such as soils, cave walls, and symbiotic associations, many of which are examined in this book.

While the marine-freshwater divide is not always clear, the distinction is revealed in the variety of algae that occur in these environments. There are no exclusively freshwater groups of algae, but Cyanobacteria, Chlorophyta, and Charophyta exhibit much greater abundance and diversity in fresh waters