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Ecology and Classification of North American Freshwater Invertebrates
Ecology and Classification of North American Freshwater Invertebrates
Ecology and Classification of North American Freshwater Invertebrates
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Ecology and Classification of North American Freshwater Invertebrates

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The Third Edition of Ecology and Classification of North American Freshwater Invertebrates continues the tradition of in-depth coverage of the biology, ecology, phylogeny, and identification of freshwater invertebrates from the USA and Canada. This edition is in color for the first time and includes greatly expanded classification of many phyla.
  • Contains extensive and detailed classification keys for identification of diverse freshwater invertebrates.
  • Many drawings and color photographs of freshwater invertebrates.
  • Single source for a broad coverage of the anatomy, physiology, ecology, and phylogeny of all major groups of invertebrates in inland waters of North America, north of Mexico.
LanguageEnglish
Release dateNov 12, 2009
ISBN9780080889818
Ecology and Classification of North American Freshwater Invertebrates

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    Ecology and Classification of North American Freshwater Invertebrates - James H. Thorp

    Ecology and Classification of North American Freshwater Invertebrates

    Third Edition

    Edited by

    James H. Thorp

    Kansas Biological Survey and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS

    Alan P. Covich

    Institute of Ecology, Odum School of Ecology, University of Georgia, Athens, GA

    Table of Contents

    Cover image

    Title page

    Front-matter

    Aquatic ecology series

    Copyright

    Dedications from the Editors

    Contributors

    Preface

    Chapter 1. Introduction to Invertebrates of Inland Waters

    I. Introduction

    II. Species and Phylogenies

    III. Taxonomy and Classification

    IV. Synopses of the Invertebrates of North American Inland Waters

    V. Taxonomic Key to Major Invertebrate Fauna of Inland Waters

    VI. Selected References

    Acknowledgments

    Chapter 2. An Overview of Inland Aquatic Habitats

    I. Introduction

    II. Lotic Environments—The Riverine Ecosystem

    III. Subterranean Habitats

    IV. Lentic Ecosystems

    V. Selected References

    Chapter 3. Protozoa

    I. Introduction

    II. Anatomy and Physiology

    III. Ecology and Evolution

    IV. Collecting, Rearing, and Preparation for Identification

    V. Identification of Protozoa

    VI. Selected References

    Chapter 4. Porifera

    I. Introduction

    II. Morphology and Physiology

    III. Ecology and Behavior

    IV. Evolution and Phylogenetics

    V. Collecting, Rearing, and Preparation for Identification

    VI. Identification of Freshwater Sponges

    VII. Key to the Freshwater Sponges of North America

    VIII. Selected References

    Acknowledgments

    Chapter 5. Cnidaria

    I. Introduction

    II. General Biology of Cnidaria

    III. Ecology of Freshwater Cnidaria

    IV. Collection and Maintenance of Freshwater Cnidaria

    V. Classification of Freshwater Cnidaria

    VI. Taxonomic Key to Genera of Freshwater Cnidaria

    VII. Selected References

    Chapter 6. Flatworms: Turbellarians and Nemertea

    I. Introduction to Turbellaria and Nemertea

    II. Turbellaria: Introduction and Status in the Animal Kingdom

    III. Anatomy and Physiology of Turbellarians

    IV. Ecology of Turbellarians

    V. Collecting, Rearing, and Identification Techniques for Turbellarians

    VI. Identification of North American Genera of Turbellarians

    VII. Nemertea: General Characteristics and External and Internal Anatomical Features

    VIII. Ecology of Nemertea

    IX. Collection, Culturing, and Preservation

    X. Taxonomic Key to Species of Freshwater Nemerteans

    XI. Selected References

    Acknowledgments

    Chapter 7. Gastrotricha

    I. Introduction

    II. Anatomy and physiology

    III. Ecology and Evolution

    IV. Collecting, Rearing, and Preparation for Identification

    V. Taxonomic Key to Gastrotricha

    VI. Selected References

    Chapter 8. Rotifera

    I. Introduction

    II. Anatomy and Physiology

    III. Ecology and Evolution

    IV. Collecting, Rearing, and Preparation for Identification

    V. Classification and Systematics

    VI. Selected References

    Acknowledgments

    Chapter 9. Nematoda and Nematomorpha

    I. Nematoda

    II. Nematomorpha

    III. Selected References

    Chapter 10. Mollusca: Gastropoda

    I. Introduction

    II. Anatomy and Physiology

    III. Ecology and Conservation

    IV. Evolution and Phylogenetics

    V. Collecting and Culturing Freshwater Gastropods

    VI. Key to Freshwater Gastropods of North America

    VII. Selected References

    Chapter 11. Mollusca: Bivalvia

    I. Introduction

    II. Morphology and Physiology

    III. Ecology

    IV. Evolution and Phylogenetics

    V. Collecting, Curation, and Rearing

    VI. Identification of North American Freshwater Bivalves

    VII. Selected References

    Chapter 12. Annelida (Clitellata): Oligochaeta, Branchiobdellida, Hirudinida, and Acanthobdellida

    I. Introduction to Oligochaeta, Branchiobdellida, Hirudinida, and Acanthobdellida

    II. Oligochaeta

    III. Branchiobdellida

    IV. Hirudinida and Acanthobdellidae

    V. Selected References

    Chapter 13. Bryozoans

    I. Introduction

    II. Morphology and Physiology

    III. Ecology and Behavior

    IV. Evolution and Phylogenetics

    V. Entoprocta

    VI. Study Methods

    VII. Taxonomic Key to North American Freshwater Bryozoans

    VIII. Selected References

    Chapter 14. Tardigrada

    I. Introduction

    II. Evolution

    III. Major Morphological Taxonomic Characters

    IV. Anatomy

    V. Latent States

    VI. Reproduction and Development

    VII. Ecology

    VIII. Techniques

    IX. Identification

    X. Taxonomic Key to Genera of Freshwater and Terrestrial Tardigrada

    XI. Selected References

    Chapter 15. Water Mites (Hydrachnidiae) and Other Arachnids

    I. Introduction to Arachnids

    II. Water Mites (Hydrachnidiae)

    III. Other Arachnids in Freshwater Habitats

    IV. Selected References

    Acknowledgments

    Chapter 16. Diversity and Classification of Insects and Collembola

    I. Introduction

    II. Aquatic Orders of Insects

    III. Partially Aquatic Orders of Insects

    IV. Semiaquatic Collembola (Springtails)

    V. Identification of Freshwater Insects and Collembola

    VI. Selected References

    Acknowledgments

    Chapter 17. Aquatic Insect Ecology

    I. Introduction

    II. Aquatic Insect Communities

    III. Aquatic Insect Life Histories

    IV. Insect-Mediated Processes

    V. Role of Disturbance and Human Impacts On Aquatic Insect Communities

    VI. Aquatic Insects in Biomonitoring Studies

    VII. Selected References

    Chapter 18. Introduction to the Subphylum Crustacea

    I. Introduction

    II. Anatomy and Physiology of Crustacea

    III. Ecology and Evolution of Selected Crustacea

    IV. Collecting, Rearing, and Preparation for Identification

    V. Classification of the Crustacea

    VI. Selected References

    Acknowledgments

    Chapter 19. Ostracoda

    I. Introduction

    II. Morphology and Physiology

    III. Ecology and Behavior

    IV. Evolution and Phylogenetics

    V. Collecting, Rearing, and Preparation for Identification

    VI. Identification of North American Nonmarine Ostracodes

    VII. Selected References

    Chapter 20. Cladocera and Other Branchiopoda

    I. Introduction and Key to Major Groups of Branchiopoda

    II. Cladoceran Anatomy and Physiology

    III. Cladoceran Distribution

    IV. Cladoceran Behavior

    V. What Cladocerans Eat and What Eats Cladocerans

    VI. Population Regulation of Cladocera

    VII. Functional Role of Cladocera in the Ecosystem

    VIII. Cladocera: Paleolimnology and the Fossil Record

    IX. Cladoceran Toxicology

    X. Collecting and Rearing Cladocera

    XI. Taxonomic Keys for North American Cladocerans

    XII. Anatomy and Physiology of Large (Noncladoceran) Branchiopods

    XIII. Ecology of Large Branchiopods

    XIV. Collecting and Rearing Techniques

    XV. Taxonomic Keys for Large Branchiopod Genera

    XVI. Selected References

    Acknowledgments

    Chapter 21. Copepoda

    I. Introduction

    II. Morphology and Physiology

    III. Ecology and Evolution

    IV. Collection, Rearing, and Preparation for Identification

    V. Identification of Copepods

    VI. Selected References

    Acknowledgments

    Chapter 22. Decapoda

    I. Introduction

    II. Morphology and Physiology

    III. Ecology and Behavior

    IV. Evolution, Phylogenetics, and Biogeography

    V. Conservation Status and Anthropogenic Impacts

    VI. Collecting, Rearing, and Preservation

    VII. Identification of Selected Shrimps, Crayfishes, and Crabs

    VIII. Selected References

    Glossary

    Subject Index

    Taxonomic Index

    Literature Citations

    Front-matter

    Aquatic ecology series

    Series Editor

    Prof. James H. Thorp

    Kansas Biological Survey, and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS

    Groundwater Ecology

    Janine Gilbert, Dan L. Danielopol, Jack A. Stanford

    Algal Ecology

    R. Jan Stevenson, Max L. Bothwell, Rex L. Lowe

    Streams and Ground Waters

    Jeremy B. Jones, Patrick J. Mulholland

    Freshwater Ecology

    Walter K. Dodds

    Aquatic Ecosystems

    Stuart E.G. Findlay, Robert L. Sinsabaugh

    Tropical Stream Ecology

    David Dudgeon

    The Riverine Ecosystem Synthesis

    James H. Thorp, Martin C. Thoms, Michael D. Delong

    Freshwater Algae of North America

    John D. Wehr and Robert G. Sheath

    Copyright

    Academic Press is an imprint of Elsevier

    32 Jamestown Road, London NW1 7BY, UK

    30 Corporate Drive, Suite 400, Burlington, MA 01803, USA

    525 B Street, Suite 1900, San Diego, CA 92101-4495, USA

    Copyright © 2010 Elsevier Inc. All rights reserved

    Chapter 15, Water Mites (Hydrachnidiae) and Other Arachnids

    Canadian Crown Copyright 2010

    No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means electronic, mechanical, photocopying, recording or otherwise without the prior written permission of the publisher

    Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone (+44) (0) 1865 843830; fax (+44) (0) 1865 853333; email: permissions@elsevier.com. Alternatively you can submit your request online by visiting the Elsevier website at (http://elsevier.com/locate/permissions), and selecting Obtaining permissions to use Elsevier material

    Notice

    No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made

    British Library Cataloguing in Publication Data

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

    Library of Congress Control Number: 2009923452

    ISBN: 978-0-12-374855-3

    For information on all Elsevier publications visit our website at www.elsevierdirect.com

    Printed and bound in Italy

    10 11 12 13 14 11 10 9 8 7 6 5 4 3 2 1

    Dedications from the Editors

    For aminals and callapiters and other natural wonders as seen through our children’s eyes.

    Prof. James H. Thorp

    To my family and to Professor G.E. Hutchinson for their inspiration and patience.

    Prof. Alan P. Covich

    Contributors

    Bonnie A. Bain, [ Chapter 12 ], Department of Biological Sciences, Southern Utah University, Cedar City, UT 84720, bain@uss.edu

    Patricia E. Bossert, [ Chapter 5 ], Long Island Group Advancing Science Education (LIGASE), State University of New York, Stony Brook, NY 11794, pbossert@optonline.net

    Ralph O. Brinkhurst, [ Chapter 12 ], 205 Cameron Court, Hermitage, TN 37076

    Kenneth M. Brown, [ Chapter 10 ], Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, kmbrown@lsu.edu

    Carla E. Cáceres, [ Chapter 20 ], School of Integrative Biology, University of Illinois at Urbana-Champaign, Urbana, IL 61801, caceres@life.uiuc.edu

    Dominic T. Chaloner, [ Chapter 17 ], Department of Biological Sciences, University of Notre Dame, Notre Dame, IN 46556, chaloner.1@nd.edu

    David R. Cook, [ Chapter 15 ], 7725 North Foothill Drive South, Paradise Valley, AZ 85253, watermites@msn.com

    Alan P. Covich, [ Chapters 1 , 2 , 18 ], Institute of Ecology, Odum School of Ecology, University of Georgia, Athens, GA 30602-2202, alanc@uga.edu

    Kevin S. Cummings, [ Chapter 11 ], Illinois Natural History Survey, Division of Biodiversity and Ecological Entomology, Champaign, IL 61820, ksc@inhs.uiuc.edu

    Ronald W. Davies, [ Chapter 12 ], Department of Biological Sciences, University of Calgary, Calgary, Alberta, Canada, ron.davies@shaw.ca

    L. Denis Delorme, [ Chapter 19 ], 621 Auburn Crescent, Burlington, Ontario, Canada

    Edward R. DeWalt, [ Chapter 16 ], Illinois Natural History Survey, Center for Biodiversity, Champaign, Il, edewalt@inhs.uiuc.edu

    Walter W. Dimmick, [ Chapter 1 ], 2612 Harper Street, Lawrence, KS 66046, U.S.A.

    Stanley L. Dodson, [ Chapter 20 ], Department of Zoology, University of Wisconsin, Madison, WI

    Stuart R. Gelder, [ Chapter 12 ], Department of Biology, University of Maine at Presque Isle, Presque Isle, ME 04769, stuart.gelder@umpi.edu

    Fredric R. Govedich, [ Chapter 12 ], Department of Biological Sciences, Southern Utah University, Cedar City, UT 84720, govedich@suu.edu

    Daniel L. Graf, [ Chapter 11 ], Department of Biological Sciences, University of Alabama, Tuscaloosa, AL 35487, dlgraf@bama.ua.edu

    Roberto Guidetti, [ Chapter 14 ], Department of the Museum of Paleobiology and Botanical Garden, University of Modena and Reggio Emilia, Via Università 4, 41100 Modena, Italy, guidetti.roberto@unimore.it

    Anne E. Hershey, [ Chapter 17 ], Department of Biology, University of North Carolina – Greensboro, Greensboro, NC 27402, anne_hershey@uncg.edu

    William L. Hilsenhoff, [ Chapter 16 ], Department of Entomology, University of Wisconsin, Madison, WI 53706

    H.H. Hobbs III., [ Chapter 22 ], Department of Biology, Wittenberg University, Springfield, OH 45501, hhobbs@wittenberg.edu

    Rick Hochberg, [ Chapter 7 ], Department of Biological Sciences, University of Massachusetts, Lowell, MA 01854, rick_hochberg@uml.edu

    William D. Hummon, [ Chapter 7 ], Department of Biological Sciences, Ohio University, Athens, OH 45701, hummon@ohio.edu

    Jurek Kolasa, [ Chapter 6 ], Department of Biology, McMaster University, Hamilton, Ontario L8S 4K1, Canada, kolasa@mcmaster.ca

    Gary A. Lamberti, [ Chapter 17 ], Department of Biological Sciences, University of Notre Dame, Notre Dame, IN 46556, gary.a.lamberti.1@nd.edu

    Charles Lydeard, [ Chapter 10 ], Smithsonian Conservation Biology Institute at NZP Center for Conservation Education and Sustainability, 1100 Jefferson Drive, Washington, DC 20013, LydeardC@si.edu

    David M. Lodge, [ Chapter 22 ], Department of Biological Sciences, University of Notre Dame, Notre Dame, IN, 46556, david.m.lodge@nd.edu

    William E. Moser, [ Chapter 13 ], Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, moserw@si.edu

    Diane R. Nelson, [ Chapter 14 ], Department of Biological Sciences, East Tennessee State University, Johnson City, TN 37614-0710, nelsond@etsu.edu

    Robert M. Northington, [ Chapter 17 ], Department of Biology, University of North Carolina – Greensboro, Greensboro, NC 27402, rmnorthi@uncg.edu

    George O. Poinar Jr., [ Chapter 9 ], Department of Zoology, Oregon State University, Corvallis, OR 97331, poinarg@science.oregonstate.edu

    Lorena Rebecchi, [ Chapter 14 ], Department of Animal Biology, University of Modena and Reggio Emilia, Via Campi 213/D, 41100 Modena, Italy, rebecchi.lorena@unimore.it

    Janet W. Reid, [ Chapter 21 ], Virginia Museum of Natural History, Martinsville, VA 24112, jwrassociates@sitestar.net

    Henry M. Reiswig, [ Chapter 4 ], Department of Biology, University of Victoria and Natural History Section, Royal British Columbia Museum, Victoria, British Columbia V8W 3N5, Canada, hmreiswig@shaw.ca

    Vincent H. Resh, [ Chapter 16 ], Department of Environmental Science, Policy, and Management, University of California, Berkeley, CA 94720, vresh@nature.berkeley.edu

    Anthony Ricciardi, [ Chapter 4 ], Redpath Museum, McGill University, Montreal, Quebec H3A 2K6, Canada, tony.ricciardi@mcgill.ca

    D. Christopher Rogers, [ Chapters 18 , 20 ], EcoAnalysts, Inc., 1307 L Street, Davis, CA 95616, crogers@ecoanalysts.com

    Robert W. Sanders, [ Chapter 3 ], Department of Biology, Temple University, Philadelphia, PA 19122-6078, robert.sanders@temple.edu

    Lawrence B. Slobodkin, [ Chapter 5 ], Deceased (2009), but formerly Department of Ecology and Evolution, State University of New York, Stony Brook, NY 11794, bzll@life.bio.sunysb.edu

    Alison J. Smith, [ Chapter 19 ], Department of Geology, Kent State University, Kent, OH 44242, alisonjs@kent.edu

    Bruce P. Smith, [ Chapter 15 ], Biology Department, Ithaca College, Ithaca, NY 14850, smithb@ithaca.edu

    Ian M. Smith, [ Chapter 15 ], Canadian National Collection of Insects and Arachnids, Invertebrate Biodiversity, Environmental Health Program, Agriculture and Agri-Food Canada, Ottawa, Onatario K1A 0C6, Canada, smithi@agr.gc.ca

    Terry W. Snell, [ Chapter 8 ], School of Biology, Environmental Science and Technology, Georgia Institute of Technology, Athens, GA 30332, terry.snell@biology.gatech.edu

    David L. Strayer, [ Chapter 7 ], Cary Institute of Ecosystem Studies, Millbrook, NY 1254, strayerd@ecostudies.org

    William D. Taylor, [ Chapter 3 ], Department of Biology, University of Waterloo, Waterloo Ontario N2L 3G1, Canada, wdtaylor@sciborg.uwaterloo.ca

    James H. Thorp, [ Chapters 1 , 2 , 18 ], Kansas Biological Survey and, Department of Ecology and Evolutionary Biology, Lawrence, KS 66047-3759, thorp@ku.edu

    Seth Tyler, [ Chapter 6 ], School of Biology and Ecology, University of Maine, Orono, ME 04469-5751, styler@maine.edu

    Robert Lee Wallace, [ Chapter 8 ], Department of Biology, Ripon College, Ripon, WI 54791, wallacer@ripon.edu

    Craig E. Williamson, [ Chapter 21 ], Department of Zoology, Miami University, Oxford, OH 45056, craig.williamson@muohio.edu

    Timothy S. Wood, [ Chapter 13 ], Department of Biological Sciences, Wright State University, Dayton, OH 45435, tim.wood@wright.edu

    Preface

    When we initially conceived the first edition of this book in the mid-1980s (subsequently published in 1991), the possibility of a second edition (2001) was only vaguely considered, much less a third (2010). While the second edition contained much new and updated information, it did not differ dramatically from the first edition, and the molecular components of taxonomy were only partially incorporated. We believe you will find this third edition significantly changed, as described below. You will immediately notice two things: (1) the book is now in four-color print instead of the previous black and white; and (2) the literature cited has been moved from the printed copy of the book to a freely downloadable website (∼15–20 references are retained at the end of each chapter). We hope you will agree that color helps convey information, especially with the addition of many new photographs of various invertebrates. While we understand that freshwater invertebrates are less colorful than their marine relatives, we hope you will better appreciate some of the variety of colors that are present in inland waters even though their biological significance is rarely clear.

    The number of references has been considerably expanded from the second edition, and these citations are available at www.Elsevierdirect.com. While we would have preferred to keep the literature cited at the end of each chapter, there are some important reasons we made this change. First, this new approach reduced the potential length of the book by at least 10% (based on the size of the second edition). This allowed us to expand the number of pages devoted to taxonomic keys by including lower taxonomic levels and more figures; it also permitted us to add more discussion of new findings in invertebrate biology, ecology, and conservation. Second, you can now download all the references from a chapter or even from the entire book if you wish. In the process of making these changes, we also shifted to a numerical citation format, again to save space but also in the hope of improving the flow of the sentences for the reader.

    Even more importantly, the third edition has been improved substantially in a variety of scientific areas. The amount of scientific expertise in the book has undergone a net increase of 13 coauthors (16 added and 3 departed). Phylogenetic information has increased throughout the book, including much greater background information on this subject in Chapter 1.

    Perhaps the most dramatic scientific change has been in the specificity and length of many taxonomic keys. Of the 19 taxonomic chapters (i.e., ignoring Chapters 1, 2, and 17), six chapters key invertebrates to species, four to genus or species (depending on the major group), seven to genus, and two to family (Protozoa and Insecta). In the last case, we employ family level keys because of the large size of the group and the presence of widely available specialty texts in those areas. In general, the increase in taxonomic specificity was greatest for the rotifer and crustacean chapters. As we stated in our two previous books, we again caution the reader to be careful to avoid exceeding your taxonomic expertise when working through the keys. Unless you have an updated, accurate list of species reported by experts in your area, you should proceed with caution when attempting to identify invertebrates to species. In some cases, a nonspecialist can reliably identify an invertebrate to the species level only when working with a particular life stage (e.g., Form I male crayfish). Most chapters provide source information on where you can locate taxonomic keys published at more detailed levels and/or for particular geographic regions.

    As in our previous two books, this edition is strongly focused on species found in inland surface waters of the United States and Canada, but several of the chapters have expanded their coverage of species dwelling in underground aquatic systems. Again, most estuarine and parasitic species are not covered in this book.

    As in our first book, many people contributed to this edition in addition to the named authors. In many chapters you will find acknowledgments to contributors who have provided information or evaluated the individual chapters. We greatly appreciate all our colleagues who have contributed information, figures, or reviews to the third edition and also thank those who provided similar services for the earlier editions, upon which the present book partially relies. In particular, we would like to thank Brian O’Neill (one of JHT’s graduate students at the University of Kansas) who helped with a number of aspects of this edition, including formatting the huge literature-cited section, and Prof. Michael Delong at Winona State University who offered advice throughout the time-consuming and sometimes exacerbating process of putting together this book. Finally, we are again grateful to the highly competent people at Academic Press/Elsevier who helped in many aspects of the book’s production from the original concept to the final marketing. In particular, we appreciate the long association with Andy Richford in London, England and newer contributions from Emily McCloskey and Candice Janco in Burlington, Massachusetts.

    James H. Thorp

    Alan P. Covich

    Chapter 1

    Introduction to Invertebrates of Inland Waters

    James H. Thorp, Kansas Biological Survey and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas

    Alan P. Covich, Institute of Ecology, Odum School of Ecology, University of Georgia, Athens, Georgia

    Walter W. Dimmick, Kansas Biological Survey, and Department of Ecology and Evolutionary Biology, University of Kansas, Larence, Kansas

    I. Introduction

    At some point in almost every biologist’s life, an invertebrate stimulated their curiosity. It might have started when they peered through a microscope at pond water and saw a protozoan, hydra, rotifer, or some other creature. Or perhaps they watched a butterfly or bee pollinating a flower, a rolly-polly pill bug curled in a ball within their palm, ants marching rapidly past while carrying heavy loads, waving anemones within a tide pool, or a crayfish scurrying from under a stream rock. For many authors contributing to this book, this childhood fascination led progressively, if not inevitably, to a life-long career in science.

    Aquatic scientists focusing on invertebrates conduct research in a wide diversity of fields. Some are fascinated by invertebrate variety and the relationships among species over evolutionary history, or they may be involved in conducting environmental assessments where knowledge of invertebrate identification is typically essential. Others seek to control human diseases by understanding and controlling invertebrate vectors of water-borne diseases, or perhaps they study the population biology or culturing of invertebrates involved in the food chains from which we derive many items, including freshwater crayfish and the zooplankton prey of trout and largemouth bass. Many more investigate the ecology of invertebrates to satisfy their scientific curiosity or to help abate environmental crises facing the natural and human world from future global climate change to more immediate and local environmental perturbations.

    Next to their more colorful, and often larger, marine relatives, freshwater invertebrates may initially seem drab and uninspiring. Yet once the curious observer penetrates beyond superficial appearances and thoroughly examines the diverse structural, physiological, behavioral, and general ecological adaptations of freshwater invertebrates, few fail to be impressed by these fascinating animals. In this book, students and professionals alike will be exposed to new aspects of the biology, ecology, and classification of freshwater and other inland aquatic invertebrates. The authors of this book have reviewed the recent literature and drawn together some of the most important highlights associated with these diverse organisms. In this edition, we have also included new images and color plates to illustrate the diversity of forms and function.

    This chapter, in particular, serves as a very brief introduction to freshwater invertebrates of Canada and the United States. Following a short discussion of procedures for naming and classifying organisms, we briefly describe the taxonomic groups covered in Chapters 3–22. We also include a taxonomic key to help the reader begin the process of classifying freshwater invertebrates. This dichotomous key will lead to the appropriate chapter for more detailed biological and taxonomic information. Additional information on freshwater invertebrates can be found in various texts on both invertebrate zoology[²,⁹,¹⁴,¹⁸] and aquatic ecology[1,22].

    II. Species and Phylogenies

    The identification of natural units is one of the major guiding principles for biologists that endeavor to study, discover, and document the world’s biodiversity. A natural unit of biodiversity is defined here as a unit that results from evolution. Examples of natural units of biodiversity are populations, species, and monophyletic groups of species. Natural units that are given a formal name (e.g., species) are called natural taxa. In contrast, subspecies is an example of a taxonomic unit that is often not a natural unit but rather a convenient partition of geographic variation. Because the number of animal species is enormous and the number of biologists studying them is minuscule in comparison, it is not surprising that our poor understanding of many genera, tribes, and families would frequently result in groups that are not natural taxa.

    The principal intellectual tools needed to discover natural taxa have emerged from the disciplines of systematic biology and population genetics. Species concepts and methods for constructing and evaluating phylogenetic trees are central to systematic biology. The discipline of population genetics is important because it permits direct study of the degree of connectivity between populations. Studies of molecular variation enable us to discover the existence of species that are undetectable when we restrict our data to morphological characters. This has resulted in the recognition of multiple species that were formerly thought to be a single wide-ranging species.

    Many biologists disagree about the best species concept and which methods of phylogenetic construction are most reliable. While this may seem to be an inconvenient situation for a new student or a researcher who is primarily interested in ecological or behavioral studies, an informed viewpoint about species concepts and phylogenetics is prerequisite for any serious student of freshwater invertebrates. Because of the extensive loss of habitat due to anthropogenic disturbances and the impending wide-scale ecological disruptions from global climate change, extinction is likely for an enormous number of species. Freshwater species could be at the leading edge of the forthcoming storm of extinction, and thus it is more important than ever to underpin ecological studies with a robust understanding of the natural units of biodiversity which we label, or sometimes mislabel, with taxonomic names. The following two sections present a brief introduction to some of the important implications of using different species concepts and the critical importance of phylogenetic trees.

    A. Species and Concepts of Species

    Students of invertebrate zoology are occasionally confronted with the daunting prospect of identifying specimens to the level of species. Unfortunately, the intellectual basis for taxonomic decisions at the species level is sometimes questionable. Freshwater invertebrates are an interesting and difficult group of organisms because of their great biological diversity and a dearth of genetic studies of wide-ranging species. Informed decisions about the validity of a species requires an understanding of which species concept was used to label the group of organisms collected at a particular place and time.

    Species are the result of historical natural processes[⁶]. Anagenesis is the modification of lineages through mutation, gene flow, natural selection, and random genetic drift. Speciation is the creation of a new lineage by splitting a preexisting lineage or the result of hybridization between two preexisting lineages. Clearly, evolution is a phenomenon of lineages, and these lineages are called species by evolutionary biologists. Therefore, the first logical step to understanding current and past biodiversity is the identification of species and other natural units that result from evolution[¹²].

    Because species are the products of evolution, their importance as units of biodiversity is paramount. Scientists often need to identify units of biodiversity below the species level, but ultimately definitions of these units must be cast in the light of an appropriate species concept because species serve as the ultimate frame of reference for studies of population variation. Species concepts play a critical role in the interpretation of intraspecific variation because species as natural taxa provide an upper boundary for comparisons among different populations. Whether a particular population or group of populations is distinctive enough to be recognized as a species has been a key issue in many taxonomic and conservation disputes. An understanding of what species are and how they can be identified is required to interpret studies of intraspecific variation and formulate biologically meaningful policies for identifying natural units of biodiversity.

    The scientific literature about the nature of species and the best method of identifying species is enormous and contentious. Perhaps, the large quantity and legitimate controversies can even be considered a barrier to understanding for students, professional biologists, and policymakers. As an intellectual framework for understanding and organizing the many different concepts of species, it is useful to consider that they almost always fall into one of two different categories: operational species concepts or ontological species concepts. Operational species concepts provide a researcher with specific criteria, that is, an algorithm to implement in order to determine whether or not two different populations belong to the same species. The biological species concept of Mayr[¹³] is a classic example of an operational species concept. The operation used by Mayr’s biological species concept is the discovery of barriers to sexual reproduction that are assumed to define the boundaries of species. Ontological species concepts provide a theoretical definition of what a species is but do not specify a methodology for the identification of species. The evolutionary species concept of Wiley[²³] is an example of an ontological species concept. In this case, species are defined as lineages expected to persist through time, but no method (operation) is prescribed for the discovery of these lineages. In more common parlance, ontological concepts can be understood as theoretical definitions of what species are. Operational species concepts provide only a method of how to discover species and are, by definition, limited by their particular methodology.

    Below you will find brief definitions of the biological, phylogenetic, cohesion, and evolutionary species concepts along with comments on their strengths and limitations. These concepts have all been influential and provide a good basis for an entry-level understanding of the many important issues regarding concepts of species. For a more thorough discussion and evaluation of species concepts, we recommend a careful reading of an article by Mayden and Wood[¹²].

    1. Biological Species Concept

    Definition

    Species are groups of actually or potentially interbreeding populations which are reproductively isolated from other such groups[¹³].

    The biological species concept has been prevalent in the evolutionary literature for the last several decades and is emphasized in many college-level biology courses. It is probably the species concept most familiar to biologists in diverse fields, such as conservation biology, forestry, fisheries, and wildlife management. Species defined by the biological species concept have also been championed as units of conservation[17].

    Theodosius Dobzhansky, a prominent evolutionary geneticist and an important contributor to the modern evolutionary synthesis, characterized the concept of a biological species as a system of populations[7,8]. The gene exchange between these systems (species) is limited or prevented by reproductive isolating mechanisms, such as species-specific breeding behaviors, hybrid sterility, and gametic incompatibility. Thus, under the biological species concept, species are simultaneously a reproductive community, a gene pool, and a genetic system. The study of reproductive isolating mechanisms is central to the biological species concept because these mechanisms provide barriers to gene flow that define the boundaries of the reproductive community and gene pool, and preserve the integrity of the genetic system of the species. In practice, however, isolating mechanisms are rarely studied and species are usually diagnosed by differences in phenotypic (morphological) features.

    Despite the long historical acceptance of the biological species concept, it has become controversial because a growing number of evolutionary biologists have found the biological species concept unworkable in a wide variety of situations. Critics of the biological species concept come from the fields of both botany and zoology. A fundamental drawback to this concept is that it is exclusively defined in terms of sexual reproduction. Asexual taxa are obviously excluded from this concept, but it is also true that many species capable of sexual reproduction cannot be easily accommodated within the framework of the biological species concept. From the viewpoint of population genetics, species capable of self-fertilization (e.g., parasitic tapeworms and some plants) and those with mandatory sibling mating are more similar to asexual than to sexually outcrossing species[²⁰]. Species that freely hybridize (open mating systems) with one or more other species yet maintain their evolutionary identity as species also provide a serious challenge to the validity of the biological species concept. Groups of freely hybridizing species are known from plants, insects, and vertebrates[²⁰].

    Another important limitation of the biological species concept concerns speciation. The most widely accepted model of speciation is the allopatric model. Generally speaking, the allopatric model entails the geographic subdivision of a single population followed by the differentiation of the isolated subpopulations into new species. Historically, the notion of a correlation between geographic subdivision of populations and speciation grew out of the observation that the closest relatives tend to occupy separate but contiguous geographic areas. Thus, in allopatric speciation lineage independence is achieved when two or more lineages are geographically disjunct. Therefore, isolating mechanisms which are fundamental to the biological species concept have very little, if anything, to do with the process of speciation because the populations undergoing speciation are geographically disjunct from one another. Hence, the evolutionary forces responsible for allopatric speciation have nothing to do with the isolating mechanisms that are a fundamental aspect of the biological species concept. A species concept that fundamentally fails to illuminate the process of speciation cannot provide the intellectual framework for the identification of units of biodiversity. Because it is impossible to study gene flow and reproductive behavior of species known only from fossil remains, the biological species concept cannot be applied to the thousands of species known only from their fossils.

    In summary, the major limitations of the biological species concept are that it is irrelevant to allopatric speciation and is inapplicable to: (1) fossil species; (2) organisms reproducing asexually or with extensive self-fertilization; and (3) sexual organisms with open mating systems (species that freely hybridize). On the other hand, the biological species concept is intuitively appealing to people because Homo sapiens fits in the narrow range of sexual reproduction that is the domain of the biological species concept.

    2. Phylogenetic Species Concept

    Definition

    A species is an irreducible cluster of organisms diagnostically distinct from other such clusters and within which there is a parental pattern of ancestry and descent[⁵].

    Phylogenetic methods have emerged as the most prominent approach for estimating the evolutionary relationships among species. Several species concepts have been proposed that are based on the methods and philosophy of phylogenetic systematics[⁵,15,16]. The exemplar definition presented above was proposed by Cracraft[⁵] and it illustrates the historical viewpoint of phylogenetic species concepts. The identification of phylogenetic species is based on the discovery of characters unique to a population or group of populations. Morphological features are the most commonly used characters to diagnose different species. Behavioral attributes and molecular genetic features (i.e., allozymes, mtDNA haplotypes, nucleic acid sequences, amino acid sequences) are also accepted as valid characters. These characters are the result of the unique evolutionary history of the population(s) (e.g., species) and serve to distinguish them from other species.

    Phylogenetic species concepts are attractive to many biologists because they use a popular methodology (cladistics). Furthermore, the methodology for discovering phylogenetic species is clearly articulated in several of the proposed phylogenetic species concepts. In contrast to the biological species concept, phylogenetic species concepts can accommodate both asexual and sexual species, as well as species that may undergo occasional or extensive hybridization. Another advantage of the phylogenetic species concept is that species are considered to be units of evolution that are equivalent in comparative biological studies[4].

    Even though phylogenetic species concepts have several positive features, several drawbacks are inherent to phylogenetic species concepts [¹²]. In some circumstances, they are prone to either underestimate or seriously overestimate the number of species present.

    3. Evolutionary Species Concept

    Definition

    A species is a single lineage of ancestor–descendant populations which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate[²³].

    The evolutionary species concept was refined and popularized in Wiley’s book[²³] on the principles of phylogenetic systematics published in 1981. The evolutionary and phylogenetic concepts are related but very different. The evolutionary species concept is ontological and simply states that species exist as lineages. It does not specify a methodology for the identification of species. However, the phylogenetic species concept is operational, not ontological, since it specifies an explicit methodology for the identification of species.

    The evolutionary concept recognizes species as a single lineage of ancestor–descendant populations which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate. This concept is advantageous because it is applicable to living, extinct, sexual, and asexual groups. It also emphasizes that species can be held together through gene flow and both developmental and ecological constraints.

    Interestingly enough, this is the concept that most biologists actually use whether or not they realize it. Decisions regarding species status are usually based on patterns of phenotypic cohesion within a group compared to phenotypic discontinuity with other groups. Although scientists rarely articulate which species concept they used when describing new species, the vast majority of all species described have been based on morphological comparisons. Even taxonomists that are ardent supporters of the biological species concept are essentially using the evolutionary species concept because their decisions are based on phenotypic continuity/discontinuity and not comparisons of gene flow among populations.

    A complaint about the evolutionary species concept concerns the predictive element (…own evolutionary tendencies and historical fate) of the definition. Many biologists do not like the requirement that you must judge or predict the future of a population when naming a species. This predictive element is in fact a logical expectation of operating within the modern framework of the hypothetico–deductive model; thus, naming a species is generically similar to most scientific activities. A hypothesis is formulated that makes a prediction about the nature of the physical world that will be borne out or rejected by the acquisition of future empirical data.

    The phylogenetic species concept does not contribute anything to the evolutionary concept, but the phylogenetic species concept has the drawback that there is an operational aspect to the definition. The operational aspect is that the definition is limited to diagnosable species which are predicated on human abilities. Because biological diversity is enormous, it seems highly unlikely that any single operational concept could be devised to encompass all species. The many limitations of the biological species concept are a strong testimony to this assertion. Multiple operational concepts are not an option because this would result in different kinds of species, and the list of kinds of species would grow every time we encounter a new group of organisms that did not fit into our preexisting concepts.

    4. Cohesion Species Concept

    Definition

    A species is the most inclusive group of organisms having the potential for demographic and genetic exchangeability[²⁰].

    The cohesion species concept was proposed and described in detail by Templeton[²⁰]. His concept emphasizes the evolutionary processes that hold evolutionary lineages (species) together through time. Genetic exchangeability is the exchange of genes between individuals through sexual reproduction. Since biological species are defined by gene flow, the evolutionary processes (e.g., isolating mechanisms) that curtail or promote gene flow are paramount in defining the boundaries of sexually reproducing species. However, Templeton pointed out that gene flow is only one of the microevolutionary forces acting on populations and that random genetic drift and natural selection must have a more universal role in defining evolutionary lineages because they can act on both sexual and asexual populations.

    B. Demographic Exchangeability

    Since genetic exchangeability can counter the subdividing effects of natural selection and genetic drift for sexually reproducing populations, it is crucial to know what kinds of mechanisms prevent or promote subdivision of asexual populations. Another explanation is required for the cohesion of asexual taxa, and this explanation is termed demographic exchangeability. The fundamental niche is defined by the genetic tolerances of individuals to some set of ecological conditions. If two individual members of an asexual population occupy the same fundamental niche, then as individuals they are demographically exchangeable. Thus, complete demographic exchangeability occurs when all individuals in a population display exactly the same range of tolerances for all relevant ecological variables. Another way of thinking about this is in the historical perspective of ancestor–descendant relationships. Consider that in a hypothetical sexual population with complete genetic exchangeability, any single individual organism could become the ancestor to all members of the population at some point in the future. In an asexual population with complete demographic exchangeability, the same result could occur. Any one individual could become the ancestor to all members of the future population. Demographic exchangeability is an important mechanism for maintaining cohesion among asexual taxa.

    To better understand how demographic exchangeability works, we can compare it with genetic exchangeability in sexual taxa. Suppose a mutation arises in an individual of a sexual population, which decreases the individual’s ability to mate with some subset of the members of its population. This means it would no longer have complete genetic exchangeability. This is obviously a weakening of the cohesion to the population for this individual and any of its descendants. Now further suppose that an asexual individual has a mutation that changes its tolerances to the ecological factors that define its fundamental niche. It will then pass on these changes to its descendants. Given that the parameters of their fundamental niche are no longer completely demographically exchangeable, their cohesion to the overall population has consequently been diminished. Templeton’s[²⁰] concept of demographic exchangeability readily allows the incorporation of microevolutionary forces other than gene flow. Random genetic drift and natural selection are forces that operate as readily on asexual as sexual reproductive systems, and they act as cohesive forces for asexual species.

    The cohesion, evolutionary, and phylogenetic concepts each have in common their recognition of the importance of cohesion and ancestor–descendant relationships. Templeton, the author of the cohesion concept, criticized the evolutionary concept as dealing with the manifestation of cohesion and not its mechanics. Templeton has greatly illuminated the mechanisms of cohesion, but his cohesion concept is really contained within the framework of the evolutionary concept.

    A good intellectual benchmark to start from, or return to, when considering any aspect of species or speciation is Darwin’s basic proposition that new species originate by the splitting of preexisting species. Under this premise we accept the idea that species are new lineages, but we are not constrained in any way on how to identify them. This leads us to an ontological species concept rather than an operational species concept. Arguably, the evolutionary species concept is the best available concept that can be used to identify species level units of biodiversity because this concept is consistent with what is known about naturally occurring units that are the result of evolutionary history[¹²]. Nevertheless, the cohesion concept articulated by Templeton[²⁰] complements the evolutionary species concept by identifying the mechanisms that are responsible for the speciation process. Templeton’s cohesion mechanisms[²⁰] deserve careful consideration because they play a critical role in the process of speciation and generation of biodiversity. Students of invertebrate zoology are encouraged to carefully read and consider Templeton’s paper[²⁰] because it is especially relevant to the diverse reproductive strategies of aquatic species of invertebrates.

    C. The Role of Phylogenies in Studies of Ecology and Behavior

    The arrangement of the present volume is based on the current classification of invertebrate phyla, and that classification rests upon an understanding of each group’s phylogenetic relationships. A stable scheme of classification is of inestimable value for organizing our knowledge of biodiversity. However, phylogenies play an even more critical role because they provide a crucial intellectual framework for comparative studies of not only morphology but also behavior and ecology at the organismal and population levels. In the preface to their book, Phylogeny, Ecology and Behavior, Brooks and McLennan[³] noted that three of the most eminent invertebrate zoologists in the modern history of biology, Charles Darwin, Willi Hennig, and Herbert H. Ross, consistently stressed the importance of integrating studies of phylogeny, ecology, and behavior. Although some of the reasons that a phylogenetic basis for comparative studies is needed are summarized below, the landmark book by Brooks and McLennan should be consulted by researchers engaged in studies that are predicated upon comparisons among species.

    Perhaps the best way to discover biologically meaningful groups of species that are statistically comparable is to analyze their phylogeny. This is very difficult to accomplish effectively or accurately above the species level. (Many ecological questions can, of course, be studied at community and greater organizational levels using genera or even higher taxonomic levels.) Statistical techniques routinely assume that samples are independently drawn from a model distribution of samples. However, species are not independent but are instead connected through history. Because some species are more closely related than others or arose at different times, species cannot be blindly treated as independent samples because of their historical, phylogenetic connection. The pattern of phylogeny must be accounted for and that is why comparative studies of morphology, behavior, and organismal and population ecology are best examined in the context of a phylogenetic framework. Moreover, because all species are a mosaic of characters that have evolved over time, not only the pattern of speciation but also the timing of speciation and character evolution must be considered. A character that evolved two million years ago (mya) cannot necessarily be evaluated in the context of current ecological conditions. A phylogeny is also needed to ensure that a feature is evaluated in the correct historical context. The chief difficulty of integrating a phylogenetic approach to comparative studies is that the demand for accurate phylogenies greatly exceeds the supply.

    D. Phylogenetic Trees

    From the time of Darwin until the 1950s, phylogenies were, for the most part, the result of personal opinion based on expertise and familiarity with a particular group. There simply were no repeatable methods widely used to formulate phylogenetic hypotheses. The development of numerical taxonomy by Charles Michener and Robert Sokal catapulted the enterprise of formulating phylogenetic hypotheses from the era of informed opinion into the age of empirical statistical analysis. Their basic premise was that to escape the subjectivity of the previous approach, the analysis of a large number of characters and a measure of the overall degree of similarity between pairs of species would provide a more reliable phylogenetic tree and thus a more stable scheme of classification. Their approach became known as phenetics.

    The idea of basing phylogenetic hypotheses on overall similarity was soon challenged by the emerging discipline of cladistics. Over a decade of rancorous debate ensued between the disciples of phenetics and cladistics following the 1966 English translation of Hennig’s book, Phylogenetic Systematics[¹¹]. Mark Twain once wrote, A scientist will never show a kindness for a theory which he did not start himself. Perhaps at no other time have these words rung truer than during the contest between cladists and pheneticists.

    In his book, Willi Hennig asserted that only one of three types of similarity could be used to formulate phylogenetic hypotheses. A character or attribute of two or more species may be similar due to convergent or parallel evolution. This type of similarity is not only devoid of phylogenetic information, it is likely to be positively misleading because a character or an attribute of two or more species may be similar due to common descent. This type of character is termed a homologous character and exists in two forms: shared derived and shared primitive. The phenetic approach includes all three of these types of similarity. In contrast, Hennig’s cladistic methodology is based on the premise that only shared derived characters can be used to support phylogenetic relationships. Shared derived characters are also called synapomorphies and must meet the criterion of homology. Shared primitive characters (symplesiorphies) also must meet the criterion of homology, but they cannot be used to establish phylogenetic relationships. Whether a character is synapomorphic or symplesiomorphic is contingent upon its distribution on a phylogenetic tree in reference to the group being analyzed. For example, the character state presence of hair is a synapomorphy supporting the monophyly of mammals. However, the character state presence of hair is a symplesiomorphy for the family Hominidae because it evolved before the group Hominidae and thus cannot inform us about relationships among members of the Hominidae.

    The widespread use of allozyme markers and mtDNA haplotypes stimulated the development of sophisticated computer algorithms to analyze these data. During the late 1980s, the development of polymerase chain reaction technology revolutionized molecular biology and enabled systematists to gather DNA sequence data for relatively large number of species. The general approach used by many computer programs that are capable of analyzing large data sets can be modestly described as a two-step process. In the first step all possible tree topologies are generated. In the second step, the characters are optimized along the tree branch and a criterion such as maximum likelihood or maximum parsimony is used to choose the best tree. An alternative approach is to generate a measure of genetic distance from the sequence data and fit the pairwise distances between all taxa to a branching diagram (tree). Because the number of trees scales up dramatically as the number of taxa under study is increased, it is not usually possible to optimize the characters for every possible tree topology.

    The currently available computer packages for conducting phylogenetic studies of morphological and molecular data present the user with a very large range of analytical choices. Each choice is burdened with a particular assumption about the data or how to analyze the data. The choices made can have a dramatic impact on the resulting phylogeny and the consumers of these phylogenies bear the burden of deciding whether or not the published phylogeny meets their standard of acceptability. For example, the choice of whether to analyze morphological data simultaneously with molecular data or to analyze the data separately and present the resulting phylogenies as a consensus tree remains a disputed point, but the two different approaches will likely produce dramatically different phylogenies. A long recognized but unresolved problem concerns the difficulty in determining the robustness of a phylogenetic tree or a subsection of a tree. A comprehensive review of the methods and controversies regarding the inference of phylogenetic trees can be found in Felsenstien[¹⁰]. Our best general advice for prospective users of phylogenetic trees who are unfamiliar with the methods and controversies is caveat emptor.

    III. Taxonomy and Classification

    Even though phylogenies and species are the result of evolution, the rules of taxonomy and hierarchical classification of groups of species are arbitrary human endeavors. Although it is vitally important for our classifications and named species to reflect biological reality, it is unrealistic to expect a perfect coherence between our classifications and the material results of more than three billion years of evolution. Thus, no phylum is exempt from an occasional reshuffling of species, renaming of constituent taxa, and recalculation of total species richness (number of species). This dynamic feature of the discipline is the expected response to the acquisition of new knowledge, but it also results from the ubiquitous and almost inherent disagreements that usually typify groups of two or more scientists. In this book, you will often encounter references to current taxonomic debates. The final classification of taxa has been left to the authors of individual chapters. As editors, Thorp and Covich have allowed the chapter authors a great amount of leeway in this area, requesting only that they inform the reader about the principal areas of dispute.

    Species are named using a system of binominal nomenclature (=names composed of two names) based on the 18th century proposal of the brilliant Swedish naturalist Carl von Linné (or Carolus Linnaeus, the latinized form he preferred to use). This is the genus and species designation familiar to most biology students (e.g., Homo sapiens for the human species). You will occasionally see a species designated with three names (a trinomen), which is permissible under the guidelines of an international code described below. For example, Bosmina (Sinobosmina) freyi is the genus, subgenus, and species name of a water flea, or cladoceran, found in some rivers (it is one of a species complex formerly grouped and named Bosmina longirostris). In contrast, Pheidole xerophila tucsonica is the genus, species, and subspecies name of a harvester ant living in the desert near Tucson, Arizona. Note the difference in capitalization of these two taxa. Scientific names are either in Latin or in a form that has been latinized [e.g., B.(S.) freyi was formed by latinizing the last name of Dr. David G. Frey, to honor this now-deceased coauthor of the branchiopod chapters of previous editions of this book].

    The final arbiter for naming taxa according to this system is the International Commission on Zoological Nomenclature, a judicial body elected periodically by the International Congress of Zoology to evaluate taxonomic names proposed by scientists. The guiding principles of these rules are priority and stability. Priority is an important principle because it helps to ensure that the first scientist to name a species will receive credit for their work. Stability is absolutely vital for communication and understanding among colleagues world-wide in the past, present, and future. The spelling of a species name remains constant among disparate languages.

    The International Commission on Zoological Nomen-clature uses a set of agreed-upon rules, termed the International Code of Zoological Nomenclature (ICZN). The commission bases its decisions on rules covering the priority of names and designation of type taxa. As the name implies, priority specifies that the first name assigned in a publication is generally the official name of that taxon. The commission has plenary powers to set aside the rightful name of a species if by doing so the cause of science would be further advanced. For example, alteration of a familiar, long-standing name could cause a major confusion in the scientific literature. Occasionally, two distinct taxa are inadvertently assigned the same binominal name by scientists working in different institutions. In that case, the species described last loses its name as a junior homonym. To lessen the state of confusion and diminish possibilities for future arguments, one individual of each species is designated as the prime example of that taxon. Such type specimens are deposited in one of the many internationally recognized museums and made available for study by responsible scientists. Type specimens can only be replaced if the original is inadvertently lost or destroyed, even if the type specimen turns out not to be the best representative of that species’ form.

    Information on systematic zoology is published in various journals, such as the Zoological Record, Zeitschrift f ;r Wissenschaftliche Zoologie (Abteilung B), and Bulletin Signaletique (CNRS), as well as in numerous society publications and journals dealing with a broad range of subjects including classification. Readers interested in knowing where best to search for papers on a given taxon should consult the Selected References section in the appropriate chapter of this book.

    The International Code of Zoological Nomenclature passes judgment on names of genera and species but does not strictly control higher classification. For that reason, some disagreements exist about the numbers and names of higher classification tiers, especially at the ordinal level and above (e.g., the phyla Cnidaria vs. Coelenterata, Bryozoa vs. Ectoprocta, and disagreements on whether Crustacea is a phylum, subphylum, or class). The system for adding suffixes to taxonomic names as a means of designating classification level has been confusing in the past but is beginning to change for the better. A nonexclusive list of the principal zoological ranks and suffixes is shown in Table 1.1.

    Table 1.1

    Principal zoological ranks and suffixes where designated.

    aAn ending recommended but not mandatory according to the International Code of Zoological Nomenclature.

    bAn ending customary but not cited in the code.

    The importance of correct identification and consistent approaches to classification soon becomes apparent to all people interested in learning about general biotic relationships. Vast amounts of information are cataloged in research libraries and museums that use the Linnean system of binominal classification. Once a specimen is correctly identified, a large number of pertinent references in specialized journals can be accessed, not only in systematics but also in ecology, evolution, animal behavior, and medical sciences. Furthermore, development of the worldwide web and the Internet has opened a tremendous resource for taxonomists and other biologists. By using computer-based information retrieval systems present in most libraries or your personal access to the worldwide web, you can rapidly track down biological and taxonomic research on an incredible number of species. Be cautious as you search the web, however, because the age and reliability of the information vary greatly among sites. In general, information that has been published in a refereed journal or book is much more reliable if not overly dated. However, websites sponsored by scientific societies or large museums are usually good sources of information and are more easily accessed by most students.

    IV. Synopses of the Invertebrates of North American Inland Waters

    All major phyla of invertebrates, other than Echinodermata, have some freshwater representatives, but only a few are more diverse in freshwaters than in the world’s oceans (e.g., insects and rotifers). Some salient features of these freshwater taxa, as discussed in Chapters 3–22, are summarized in the remainder of Section IV.

    A. Protozoa (Chapter 3)

    Protozoa are unicellular (or acellular) eukaryotes existing either as individuals or as members of a loose-knit colony (Fig. 1.1). The term protozoa is a taxon of convenience, having no significance in classification schemes within the kingdom Protista (or Protoctista) other than as a traditional functional grouping of all heterotrophic and motile species. For the purpose of this chapter, the authors have defined protozoa as unicellular or colonial eukaryotes that are heterotrophic, and they restrict their coverage to free-living, phagotrophic forms in freshwater habitats.

    Figure 1.1 (A) Dinobryon , a colonial flagellated protozoan (from Chapter 3, Fig. 3.13n);

    (B) Chaos, a non-testate amoeba (from Fig. 3.15j);

    (C) Strongylidium, a free-swimming ciliate (from Fig. 3.20p); and

    (D) Vorticella, an attached ciliate (from Fig. 3.20k).

    (Many of the figures in this chapter were taken directly from figures used in other chapters; see those chapters for original sources.)

    Modern taxonomic treatments divide the kingdom Protista into 13–27 phyla (more or less), of which two contain significant numbers of free-living, phagotrophic species. These two major groups, the ciliated protozoa and most amoeboid protozoa, are the primary subjects of Chapter 3, by William D. Taylor and Robert W. Sanders.

    Protozoa play important, though often overlooked, roles in freshwater ecosystems. For example, flagellate protozoans are the predominant predators of picoplankton (primarily bacteria and small cyanobacteria), and thus are essential components of the microbial loop (see Chapter 2) within lakes and rivers. Many genera consume bacteria within the surficial layer of sediments and in the water column, and larger taxa capture and eat algae or other protozoa. They, in turn, are preyed upon by some species of oligochaetes, chironomids, and rotifers.

    B. Porifera (Chapter 4)

    About 32 species of sponges colonize freshwater habitats in the continental North America, with a general decline in species richness as you move south and west. Although less diverse than protozoa, freshwater members of the phylum Porifera—the sponges—nonetheless are commonly found on hard substrates within many lakes and streams (Fig. 1.2). Except for the marine mesozoa and placozoa, sponges have the simplest structure of all multicellular phyla, showing little variation in external form and having an incipient, tissue-level organization. The most prominent feature of their anatomy is the progressively finer series of canals permeating the entire sponge body. Water is propelled through this canal system both passively, as a result of its basic hydrodynamic structure, and actively by choanocytes (collar cells). These cells also extract bacteria and algae from the water column by suspension (filter) feeding. Many species also contain symbiotic green algae (zoochlorellae), and for one genus shelter a yellow–green algal symbiont; both types of algae often provide more than 50% of the sponge’s nutrition.

    Figure 1.2 (A) Photograph of the freshwater sponge Ephydatia muelleri with a gradient of associated zoochlorellae (green color); (B) colony of the common sponge Spongilla lacustris exhibiting a characteristic, finger-like growth form. (A–B from Chapter 4, Figs. 4.1, 4.4, respectively)

    The annual life cycle of a freshwater sponge is characterized by shifts between periods of active growth and dormancy in the gemmule stage. Availability of suitable hard substrate and adequate food seem to be the critical factors regulating the spread and growth of sponges, as described by Henry M. Reiswig, Thomas M. Frost¹, and Anthony Ricciardi in Chapter 4. Few animals prey on sponges, and most of those graze only limited components of a sponge, leaving the remaining organisms alive to grow and reproduce.

    C. Cnidaria (Chapter 5)

    Cnidarians, or coelenterates, have been relatively unsuccessful in adapting to freshwater, being represented only by one class (Hydrozoa) with 4–5 purely freshwater genera: the common brown (Hydra) and green hydras (Hydra or Chlorohydra) with four species groups (Fig. 1.3a), and the rare medusoid Craspedacusta (Fig. 1.3b) and polypoid Calposoma and Polypodium. Cordylophora, a mostly estuarine taxon, occasionally penetrates relatively fresh waters, where it lives in a colonial, sessile form. Despite their low diversity, cnidarians—or at least hydras—are routinely present on almost any reasonably hard substrate in permanent ponds and lakes, and in streams ranging from headwater streams to large rivers.

    Figure 1.3 (A) Green hydra with an attached, budding hydra; (B) Craspedacusta medusoid hydrozoan. (A–B from Chapter 5, Figs. 5.12, 5.15, respectively)

    The cnidarian body consists of a generally thin, acellular layer of mesoglea sandwiched between two thicker cellular layers (ecto- and endoderm), all surrounding a central body cavity (the coelenteron). Coelenterates dispatch

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