Thorp and Covich's Freshwater Invertebrates: Ecology and General Biology

Ecology and General Biology

Thorp and Covich's Freshwater Invertebrates

Fourth Edition

Volume I

Editors

James H. Thorp

D. Christopher Rogers

Table of Contents

Cover image

Title page

Thorp and Covich's Freshwater Invertebrates

Copyright

Dedications from the Editors

Contributors to Volume I

Acknowledgments for Volume I

About the Editors

Preface to the Fourth Edition

Preface to Volume I

Section I. Introduction

Chapter 1. Introduction to Invertebrates of Inland Waters

Introduction

Species and Phylogenies

Using Systematics and Taxonomy to Classify Organisms

Taxonomic Keys to Invertebrates of Inland Waters

Chapter 2. Overview of Inland Water Habitats

Introduction

Global Variations in Aquatic Habitats

Lotic Ecosystems: Rivers, Streams, and Springbrooks

Subterranean Habitats

Lentic Ecosystems: Lakes, Ponds, Wetlands, and Phytotelmata

Chapter 3. Collecting, Preserving, and Culturing Invertebrates

Introduction

Collecting and Sampling

Record-Keeping

Preserving and Fixing Specimens

Culturing Invertebrates

Section II. General Ecology and Human Impacts

Chapter 4. Functional Relationships of Freshwater Invertebrates

Introduction

Distribution in Space

Acquiring Energy

Regulating Populations and Communities

Chapter 5. Ecology of Invasive Alien Invertebrates

Rates and Global Extent of Freshwater Invertebrate Invasions

Human Vectors of Dispersal

Traits Conferring Invasion Success

Ecological Impacts

Can the Impacts of Alien Invertebrates Be Predicted?

Management of Invasive Aquatic Invertebrates

Chapter 6. Economic Aspects of Freshwater Invertebrates

Freshwater Invertebrates in Commerce

Biomonitoring

Nuisance Aquatic Insects

Benefits or Damages Caused by Introduced or Invasive Species

Medicinal Leeches

Stresses to Livestock and Wildlife from Biting Flies

Diseases Vectored by Freshwater Invertebrates

Section III. Protozoa to Tardigrada

Chapter 7. Free-Living Protozoa

Introduction

Structure and Function of the Protozoan Cell

Biological Diversity of Free-Living Protozoa

General Ecology of Protozoa

Collecting and Culturing Protozoa

Chapter 8. Phylum Porifera

Introduction

General Biology

General Ecology and Behavior

Collecting, Rearing, and Preparation for Identification

Chapter 9. Phylum Cnidaria

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 10. Phylum Platyhelminthes

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 11. Phylum Nemertea

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 12. Phylum Gastrotricha

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preservation

Chapter 13. Phylum Rotifera

Introduction to Rotifera

General Biology

Ecology and Evolution

Collecting, Culturing, and Preparation for Identification

Chapter 14. Phylum Nemata

Introduction

General Biology

General Ecology and Behavior

Collecting, Rearing, and Preparing Specimens

Chapter 15. Phylum Nematomorpha

Introduction to Nematomorpha

General Biology

General Ecology and Behavior

Collecting, Culturing, and Preparing Specimens

Chapter 16. Phyla Ectoprocta and Entoprocta (Bryozoans)

Introduction

General Biology of Ectoprocta

General Ecology and Behavior of Ectoprocta

Entoprocta

Collecting, Culturing, and Specimen Preparation

Chapter 17. Phylum Tardigrada

Introduction

General Biology

General Ecology and Behavior

Collection, Culturing, and Specimen Preparation

Section IV. Phylum Mollusca

Chapter 18. Introduction to Mollusca and the Class Gastropoda

Introduction to Freshwater Members of the Phylum Mollusca

Freshwater Members of the Class Gastropoda

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 19. Class Bivalvia

Introduction

General Biology

General Ecology

Collecting, Curation, and Rearing

Section V. Phylum Annelida

Chapter 20. Introduction to Annelida and the Class Polychaeta

Introduction to Inland Water Annelida

General Biology

General Ecology and Behavior

Collecting Annelids and Specimen Preparation

Introduction to the Inland Water Polychaeta

General Biology of Polychaeta

General Ecology and Behavior of Polychaeta

Collecting Polychaetes and Specimen Preparation

Chapter 21. Clitellata: Oligochaeta

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 22. Clitellata: Branchiobdellida

Introduction to the Branchiobdellida

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 23. Clitellata: Hirudinida and Acanthobdellida

Introduction to Hirudinida and Acanthobdellida

General Biology

General Ecology and Behavior

Collecting, Rearing, and Preparation for Identification

Section VI. Phylum Arthropoda

Chapter 24. Introduction to the Phylum Arthropoda

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Preparing Specimens

Chapter 25. Subphylum Chelicerata, Class Arachnida

Introduction to Arachnids

Spiders (Araneae)

Acari: Parasitiformes

Acariformes: Freshwater Sarcoptiformes

Acariformes: Prostigmata: Overview and Freshwater Taxa Excluding Hydrachnidiae

Water Mites (Prostigmata: Parasitengonina: Hydrachnidiae)

Collecting, Rearing, and Preparation for Study

Dedication

Chapter 26. Subphylum Myriapoda, Class Diplopoda

Introduction to the Subphylum

General Biology

Ecology and Behavior of Freshwater Millipedes

Collecting, Rearing, and Preparation for Identification

Chapter 27. Introduction to Crustacea

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 28. Class Branchiopoda

Introduction

General Biology

Life History and Ecology

Collecting, Culturing, and Specimen Preparation

Chapter 29. Class Maxillopoda

Introduction

General Biology of Copepoda

General Ecology of Copepoda

General Biology and Ecology of Branchiura

General Biology and Ecology of Cirripedia

Collecting, Culturing, and Specimen Preparation

Chapter 30. Class Ostracoda

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 31. Class Malacostraca, Superorders Peracarida and Syncarida

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 32. Class Malacostraca, Order Decapoda

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 33. Hexapoda—Introduction to Insects and Collembola

Introduction to Hexapoda

Introduction to the Class Insecta

General Biology of Aquatic Insects

General Ecology and Behavior of Aquatic Insects

Introduction to Collembola

Collecting, Culturing, and Specimen Preparation of Insects and Springtails

Chapter 34. Order Ephemeroptera

Introduction To Mayflies (Ephemeroptera)

General Biology

General Ecology and Behavior

Collecting, Rearing, and Specimen Preparation

Chapter 35. Order Odonata

Introduction

General Biology

General Ecology and Behavior

Collecting, Culturing, and Specimen Preparation

Chapter 36. Order Plecoptera

Introduction

General Biology

General Ecology and Behavior

Collecting and Rearing Stoneflies

Chapter 37. Order Hemiptera

Introduction

General Biology

Ecology and Behavior

Collecting, Culturing, and Preparing Specimens

Chapter 38. Order Trichoptera

Introduction

General Biology

General Ecology and Behavior

Collecting, Rearing, and Specimen Preparation

Chapter 39. Order Coleoptera

Introduction to Aquatic Coleoptera

General Biology

Ecology and Behavior

Collecting, Preservation, and Culturing

Appendix 1: Family Descriptions by Suborders

Chapter 40. Order Diptera

Introduction

General Biology

General Ecology And Behavior

Collecting, Culturing, And Specimen Preparation

Chapter 41. Minor Insect Orders

Introduction

Megaloptera

Neuroptera

Blattodea

Hymenoptera

Lepidoptera

Mecoptera

Orthoptera

Subject Index

Taxonomy Index

Thorp and Covich’s Freshwater Invertebrates

A Global Series of Books on the Identification,

Ecology, and General Biology of Inland Water Invertebrates

by Experts from Around the World

Fourth Edition

Edited by James H. Thorp

Volume I : Ecology and General Biology

Edited by James H. Thorp and D. Christopher Rogers

Published 2015

Volume II: Keys to Nearctic Fauna

Edited by James H. Thorp and D. Christopher Rogers

Expected Publication Date: 2015

Volume III: Keys to Palaearctic Fauna

Edited by D. Christopher Rogers and James H. Thorp

Expected Publication Date: 2016

Planned Future Volumes of the Fourth Edition

Keys to Palaearctic Insects and Aquatic Collembola

Keys to Oriental Fauna

Keys to Australasian and Oceana Fauna

Keys to Neotropical and Antarctica Fauna

Keys to Afrotropical Fauna

Related Publications in This Series

Ecology and Classification of North American Freshwater Invertebrates

Edited by J.H. Thorp and A.P. Covich

First (1991), Second (2001), and Third (2010) Editions

Copyright

Academic Press is an imprint of Elsevier

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This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein).

Notices

Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary.

Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility.

To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability 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.

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

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For information on all Academic Press publications visit our website at http://store.elsevier.com/

ISBN: 978-0-12-385026-3

Printed and bound in China

Dedications from the Editors

"To the many dedicated authors and both old and new friends who over the years have contributed their efforts and hard-won knowledge to these books and research on invertebrate ecology and taxonomy."

James H. Thorp

"To my family and anyone else who has wondered what was going on under the water’s surface."

D. Christopher Rogers

Contributors to Volume I

Michael T. Bogan ,     Department of Environmental Science, Policy, and Management, University of California Berkeley, Berkeley, California, USA, email: michaelthomasbogan@gmail.com

[Chapter 41]

Matthew G. Bolek ,     Department of Zoology, Oklahoma State University, 415 Life Sciences West, Stillwater, Oklahoma 74078 USA, email: bolek@okstate.edu

[Chapter 15]

John E. Brittain ,     Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway, email: j.e.brittain@nhm.uio.no

[Chapter 34]

Kenneth M. Brown ,     Department of Biological Sciences, Louisiana State University, A343 Life Sciences Annex, Baton Rouge, Louisiana 70803, USA, email: kmbrown@lsu.edu

[Chapter 18]

Francisco Brusa ,     División Zoología Invertebrados, Museo de La Plata, FCNyM-UNLP, 1900 La Plata, Argentina, , email: fbrusa@fcnym.unlp.edu.ar

[Chapter 10]

Carla E. Cáceres ,     School of Integrative Biology, Program in Ecology, Evolution and Conservation Biology, Morrill Hall, University of Illinois at Urbana-Champaign, 505 South Goodwin, Urbana, Illinois 61801, USA, email: caceres@life.illinois.edu

[Chapter 28]

David R. Cook,     7725 North Foothill Drive South, Paradise Valley, Arizona, USA

[Chapter 25]

Rickey D. Cothran ,     Department of Biological Sciences, University of Pittsburgh, 4249 Fifth Avenue, Pittsburgh, Pennsylvania 15260 USA, email: rdc28@pitt.edu

[Chapter 31]

Gregory W. Courtney ,     Department of Entomology, Iowa State University, 432 Science II, Ames, Iowa 50011 USA, email: gwcourt@iastate.edu

[Chapter 40]

Matthew R. Cover ,     Department of Biological Sciences, California State University Stanislaus, One University Circle, Turlock, California 95382 USA, , email: mcover@csustan.edu

[Chapter 41]

Alan P. Covich ,     Odum School of Ecology, Ecology Building, University of Georgia, Athens, Georgia 30602-2202 USA, , email: alanc@uga.edu

[Chapter 2, 27]

Peter S. Cranston,     Evolution, Ecology, and Genetics, Research School of Biology, The Australian National University, Canberra ACT 0200 Australia; Also an Emeritus Professor at Entomology Department, University of California Davis, Davis, California 95616, USA, email: pscranston@ucdavis

[Chapter 40]

Neil Cumberlidge ,     Department of Biology, 2009 New Science Facility, Northern Michigan University, 1401 Presque Isle Avenue, Marquette, Michigan 49855 USA, email: ncumberl@nmu.edu

[Chapter 32]

Kevin S. Cummings ,     Illinois Natural History Survey, Division of Biodiversity and Ecological Entomology, University of Illinois, 607 E. Peabody Drive, MC-652, Champaign, Illinois 61820 USA, email: ksc@inhs.illinois.edu

[Chapter 19]

Cristina Damborenea ,     División Zoología Invertebrados, Museo de La Plata, FCNyM-UNLP, 1900 La Plata, Argentina, email: cdambor@fcnym.unlp.edu.ar

[Chapter 10]

L. Cristina De Villalobos ,     Museo de Ciencias Naturales, Paseo del Bosque FCNyM-UNLP, 1900 La Plata, Argentina, email: villalo@fcnym.unlp.edu.ar

[Chapter 15]

R. Edward DeWalt ,     Prairie Research Institute, Illinois Natural History Survey, University of Illinois, 607 East Peabody Drive, Champaign, Illinois 61820 USA, email: edewalt@inhs.illinois.edu

[Chapter 36]

Klaas-Dowe B. Dijkstra ,     Nederlands Centrum voor Biodiversiteit Naturalis, Leiden, Darwinweg 2, 2333 CR Leiden, The Netherlands, , email: kd.dijkstra@ncbnaturalis.nl

[Chapter 35]

Walter W. Dimmick ,     2612 Harper Street, Lawrence, Kansas 66046 USA, email: dimmick@sunflower.com

[Chapter 1]

Genoveva Esteban ,     Conservation Ecology and Environmental Sciences Group, School of Applied Sciences, Bournemouth University, Talbot Campus Poole, Dorset BH12 5BB, UK, email: gesteban@bournemouth.ac.uk

[Chapter 7]

Bland J. Finlay ,     School of Biological and Chemical Sciences, Queen Mary University of London, The River Laboratory, Wareham, Dorset, BH20 6BB, England, email: b.j.finlay@qmul.ac.uk

[Chapter 7]

Nadine Folino-Rorem ,     Department of Biology, Wheaton College, IL, 501 College Avenue, Wheaton, Illinois 60187 USA, email: nadine.c.folino-rorem@wheaton.edu

[Chapter 9]

Stuart R. Gelder ,     Department of Science & Mathematics, ME, University of Maine at Presque Isle, Presque Isle, Maine 04769 USA, email: stuart.gelder@umpi.edu

[Chapter 22]

Jean-Jacques Geoffroy ,     Département Ecologie et Gestion de la Biodiversité, Muséum National d’Histoire Naturelle, UMR 7204 CESCO CNRS-MNHN-UPMC, 4 avenue du petit Château 91800 Brunoy, France, email: geoffroy@mnhn.fr

[Chapter 26]

Stanislav Gorb ,     Spezielle Zoologie, Universität Kiel, Am Botanischen Garten 1-9, 24098 Kiel, Germany, email: sgorb@zoologie.uni-kiel.de

[Chapter 35]

Frederic R. Govedich ,     Department of Biological Sciences, Southern Utah University, Cedar City, Utah 84720, USA, email: govedich@suu.edu

[Chapter 23]

Daniel L. Graf ,     Department of Biology, University of Wisconsin, Stevens Point, 2100 Main St., Stevens Point, Wisconsin 54481, USA, email: dgraf@uwsp.edu

[Chapter 19]

Roberto Guidetti ,     Department of Life Sciences, University of Modena and Reggio Emilia, via Campi 213/D, 41125, Modena, Italy, email: roberto.guidetti@unimore.it

[Chapter 17]

Ben Hanelt ,     Department of Biology, University of New Mexico, Albuquerque, 163 Castetter Hall, New Mexico 87131, USA, email: bhanelt@unm.edu

[Chapter 15]

Horton H. Hobbs ,     Department of Biology, Wittenberg University, Springfield, Ohio, USA, email: hhobbs@wittenberg.edu

[Chapter 32]

Rick Hochberg ,     Department of Biological Sciences, University of Massachusetts-Lowell, One University Avenue, Lowell, Massachusetts 01854, USA, email: rick_hochberg@uml.edu

[Chapter 12]

Ralph W. Holzenthal ,     Department of Entomology, 219 Hodson Hall, University of Minnesota, 1980 Folwell Ave, St. Paul, Minnesota 55108, USA, email: holze001@umn.edu

[Chapter 38]

David J. Horne ,     School of Geography, Queen Mary University of London, Mile End Road, London E1 4NS, UK, email: d.j.horne@qmul.ac.uk

[Chapter 30]

Vincent J. Kalkman ,     Nederlands Centrum voor Biodiversiteit Naturalis Leiden, Darwinweg 2, 2333 CR Leiden, The Netherlands, email: vincent.kalkman@ncbnaturalis.nl

[Chapter 35]

Tobias Kånneby ,     Department of Zoology, Swedish Museum of Natural History, PO Box 50007, SE-104 05 Stockholm, Sweden, email: tobias.kanneby@nrm.se

[Chapter 12]

Siegfried Kehl ,     Department of Animal Ecology II, Universität Bayreuth, Universitätsstr. 30, 95440 Bayreuth, Germany, email: siegfried.kehl@uni-bayreuth.de

[Chapter 39]

Boris C. Kondratieff ,     C. P. Gillette Museum of Arthropod Diversity, Department of Bioagricultural Sciences and Pest Management, Colorado State University, 012A Laurel Hall, Fort Collins, Colorado 80523, USA, email: boris.kondratieff@colostate.edu

[Chapter 36]

David M. Lodge ,     Department of Biological Sciences, University of Notre Dame, Notre Dame, Indiana 46556, USA, email: lodge.1@nd.edu

[Chapter 32]

David A. Lytle ,     Department of Integrative Biology, Cordley Hall, Oregon State University, Corvallis, Oregon 97331, USA, email: lytleda@oregonstate.edu

[Chapter 37]

Renata Manconi ,     Dipartimento di Scienze della Natura e del Territorio (DIPNET), Università di Sassari, Via Muroni 25, I-07100 Sassari, Italy, email: r.manconi@uniss.it

[Chapter 8]

Koen Martens ,     Freshwater Biology, Royal Belgian Institute of Natural Sciences, Brussels, Belgium; and Department of Biology, University of Ghent, Ghent, Vautierstraat 29, 1000 Brussels, Belgium, email: martens@naturalsciences.be

[Chapter 30]

Patrick J. Martin ,     Biologie des Eaux, Institut royal des Sciences naturelles de Belgique douces 29, rue Vautier, B-1000, Bruxelles, Belgium, email: patrick.martin@sciencesnaturelles.be

[Chapter 21]

William E. Moser ,     Department of Invertebrate Zoology, National Museum of Natural History, Washington, DC, Smithsonian Institution, Museum Support Center, MRC 534, 4210 Silver Hill Road, Suitland, Maryland 20746, USA, email: moserw@si.edu

[Chapter 23]

Diane R. Nelson ,     Department of Biological Sciences, East Tennessee State University, Johnson City, Tennessee 37614, USA, email: janddnelson@yahoo.com

[Chapter 17]

Carolina Noreña ,     Departamento Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), Madrid, España, c/o Jose Gutierrez Abascal 2, 28006 Madrid, España, email: norena@mncn.csic.es

[Chapter 10]

Brian J. O’Neill ,     Department of Biological Sciences, University of Wisconsin – Whitewater, 269 Wyman Mall, Whitewater, Wisconsin 53190 , USA, email: oneillb@uww.edu

[Chapter 33]

George O. Poinar Jr.  ,     Department of Zoology, Oregon State University, Corvallis, Oregon 97331, USA, email: poinarg@science.oregonstate.edu

[Chapter 14]

Heather C. Proctor ,     Department of Biological Sciences, CW405 Biological Sciences Building, University of Alberta, Edmonton, T6G 2E9, Canada, email: hproctor@ualberta.ca

[Chapter 25]

Roberto Pronzanto ,     Dipartimento di Scienze della Terra, dell’Ambiente e della Vita (DI.S.T.A.V.), Università di Genova, Area Scientifico-Disciplinare 05 (Scienze biologiche), Settore BIO/05, Genova, Italy, email: pronzato@dipteris.unige.it

[Chapter 8]

Mark Pyron ,     Department of Biology, Ball State University, Cooper Life Sciences Building, CL 121, Muncie, Indiana 47306, USA, email: mpyron@bsu.edu

[Chapter 18]

Lorena Rebecchi ,     Department of Life Sciences, University of Modena and Reggio Emilia, via Campi 213/D, 41125, Modena, Italy, email: lorena.rebecchi@unimore.it

[Chapter 17]

Vincent H. Resh ,     Department of Environmental Science, Policy, and Management, University of California, 305 Wellman Hall, Berkeley, California 94720, USA, email: resh@berkeley.edu

[Chapter 6]

Anthony Ricciardi ,     Redpath Museum, McGill University, Montreal, Quebec H3A 2K6, Canada, email: tony.ricciardi@mcgill.ca

[Chapter 5]

Blanca Ríos-Touma ,     Centro de Investigación de la Biodiversidad y el Cambio Climático, Museo de Zoología, Universidad Tecnológica Indoamérica, Quito, Ecuador. [Also contact at: Department of Landscape Architecture & Environmental Planning, 300 Wurster Hall, University of California, Berkeley, California 94553, USA.], email: briostouma@gmail.com

[Chapter 38]

D. Christopher Rogers ,     Kansas Biological Survey and Biodiversity Institute, Higuchi Hall, University of Kansas, 2101 Constant Avenue, Lawrence, Kansas 66047-3759, USA, email: branchiopod@gmail.com

[Chapters 1, 3, 24, 27-28]

David M. Rosenberg ,     Emeritus Scientist Freshwater Institute, Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Manitoba, R3T 2N6, Canada, email: drosie@mymts.net

[Chapter 6]

Göran Sahlén ,     Ecology and Environmental Sciences, Halmstad University, Högskolan i Halmstad, Box 823, Kristian IV: s väg 3, 30118 Halmstad, Sweden, email: Goran.Sahlen@hh.se

[Chapter 35]

John B. Sandberg ,     California Department of Fish and Wildlife, CSUC Research Foundation, California State University, Holt Hall, 1205 W 7th St., Chico, California 95929, USA, email: jsandberg@csuchico.edu

[Chapter 36]

Michel Sartori ,     Museum of Zoology, Palais de Rumine, Place Riponne 6, CH-1014 Lausanne, Switzerland, email: michel.sartori@uni-hamburg.de

[Chapter 34]

Andreas Schmidt-Rhaesa ,     Zoological Museum, University Hamburg, Martin Luther-King. Platz 3, 20146 Hamburg, Germany, email: andreas.schmidt-rhaesa@uni-hamburg.de

[Chapter 15]

Isa Schön ,     Freshwater Biology, Royal Belgian Institute of Natural Sciences, Vautierstraat, 29, 1000 Brussels, Belgium, email: Schoen@naturalsciences.be

[Chapter 30]

Alison J. Smith ,     Department of Geology, Kent State University, Kent, Ohio 44242, USA, email: alisonjs@kent.edu

[Chapter 30]

Bruce P. Smith ,     Department of Biology, Ithaca College, Ithaca, New York 14850, USA, email: smithb@ithaca.edu

[Chapter 25]

Hilary A. Smith ,     Department of Biological Sciences, University of Notre Dame, Notre Dame, Indiana 46556, USA, email: hilary.a.smith.964@nd.edu

[Chapter 13]

Ian M. Smith ,     Canadian National Collection of Insects and Arachnids, Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Ave., Ottawa, Ontario K1A 0C6, Canada, email: smithi@agr.gc.ca

[Chapter 25]

Terry W. Snell ,     School of Biology, Environmental Science and Technology, Georgia Institute of Technology, Atlanta, Athens, Georgia 30332, USA, email: terry.snell@biology.gatech.edu

[Chapter 13]

Malin Strand ,     Coordinator, Marine Invertebrates, Swedish Species Information Centre, Swedish University of Agricultural Sciences, Box 7007, SE 75007 Uppsala, Sweden, email: malin.strand@slu.edu

[Chapter 11]

Eduardo Suárez-Morales ,     El Colegio de la Frontera Sur (ECOSUR), Unidad Chetumal, P.O. Box 424. Chetumal, Quintana Roo 77014, Mexico, email: esuarez@ecosur.mx

[Chapter 29]

Frank Suhling ,     Institut für Geoökologie, Technische Universität Braunschweig, Langer Kamp 19c, Raum 303, 38106 Braunschweig, Germany, email: f.suhling@tu-bs.de

[Chapter 35]

Per Sundberg ,     Department of Biological and Environmental Sciences, University of Gothenburg, P.O. Box 463, SE-405 30 Gothenburg, Sweden, email: per.sundberg@zool.gu.se

[Chapter 11]

Robin E. Thomson ,     Department of Entomology, University of Minnesota, Hodson Hall, 1980 Folwell Avenue, St. Paul, Minnesota 55108, USA, email: thom1514@umn.edu

[Chapter 38]

James H. Thorp ,     Kansas Biological Survey and Department of Ecology and Evolutionary Biology, Higuchi Hall, University of Kansas, 2101 Constant Avenue, Lawrence, Kansas 66047, USA, email: thorp@ku.edu

[Chapters 1–4, 24, 27, 33]

Tarmo Timm ,     Institute of Agricultural and Environmental Sciences, Centre for Limnology, Estonian University of Life Sciences, Rannu, Tartumaa 61117, Estonia, email: tarmo.timm@emu.ee

[Chapter 21]

Jan van Tol ,     Nederlands Centrum voor Biodiversiteit Naturalis, Leiden, Darwinweg 2, 2333 CR Leiden, The Netherlands, email: jan.vantol@ncbnaturalis.nl

[Chapter 35]

Piet F.M. Verdonschot ,     Freshwater Ecology, Alterra, Wageningen UR, P.O. Box 47, 6700 AA Wageningen, The Netherlands, email: piet.verdonschot@wur.nl

[Chapter 20]

Robert L. Wallace ,     Department of Biology, Ripon College, Ripon, 300 Seward Street, Ripon, Wisconsin 54791, USA, email: wallacer@ripon.edu

[Chapter 13]

Alan Warren ,     Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK, email: a.warren@nhm.ac.uk

[Chapter 7]

Gary A. Wellborn ,     Department of Biology, University of Oklahoma, 730 Van Vleet Oval, Norman Oklahoma 73019, email: gwellborn@ou.edu

[Chapter 31]

Bronwyn W. Williams ,     Department of Zoology, Southern Illinois University, 1125 Lincoln Drive, MC 6501, Carbondale, Illinois 62901, USA, email: bwwilliams@siu.edu

[Chapter 22]

Jonathan D.S. Witt ,     Department of Biology, University of Waterloo, 200 University Avenue West, Waterloo, Ontario, N2L 3G1Canada, email: jwitt@uwaterloo.ca

[Chapter 31]

Timothy S. Wood ,     Department of Biological Sciences, Wright State University, 3640 Colonel Glen Highway, Dayton, Ohio 45435, USA, email: tim.wood@wright.edu

[Chapter 16]

Donald A. Yee ,     Department of Biological Sciences, University of Southern Mississippi, 118 College Drive #c005018, Hattiesburg, Mississippi 39406-0001, USA, email: donald.yee@usm.edu

[Chapter 39]

Acknowledgments for Volume I

Many people contributed to this volume in addition to the chapter authors and those acknowledged in individual chapters. We greatly appreciate all our colleagues who have contributed information, figures, or reviews to Volume I 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 Vince Resh, who suggested a number of possible chapter authors in addition to writing his own chapter.

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 our association with Sean Coombs and Candice Janco in the U.S. offices of Elsevier, as well as past editors who assisted us in producing T&C I–III.

James H. Thorp

D. Christopher Rogers

About the Editors

Dr James H. Thorp has been a Professor in the Department of Ecology and Evolutionary Biology at the University of Kansas (Lawrence, KS, USA) and a Senior Scientist in the Kansas Biological Survey since 2001. Prior to returning to his alma mater, Prof. Thorp was a Distinguished Professor and Dean at Clarkson University, Department Chair and Professor at the University of Louisville, Associate Professor and Director of the Calder Ecology Center of Fordham University, Visiting Associate Professor at Cornell University, and Research Ecologist at the University of Georgia’s Savannah River Ecology Laboratory. He received his Baccalaureate degree from the University of Kansas and both Masters and PhD degrees from North Carolina State University. Those degrees focused on zoology, ecology, and marine biology, with an emphasis on the ecology of freshwater and marine invertebrates.

Dr Thorp is currently on the editorial board of two journals (River Research and Applications and River Systems) and is a former President of the International Society for River Science (ISRS). He teaches three courses at the University of Kansas (Principles of Ecology, River and Lake Ecology, and Marine Biology) and has both Masters and doctoral graduate students working on various aspects of the ecology of organisms, communities, and ecosystems in rivers, reservoirs, and wetlands.

Prof. Thorp’s research interests and background are highly diverse and span the gamut from organismal biology to the ecology of communities, ecosystems, and macrosystems. He works on both fundamental and applied research topics using descriptive, experimental, and modeling approaches in the field and laboratory. While his research emphasizes aquatic invertebrates, he also studies fish ecology especially as related to food webs. He has published more than one hundred refereed journal articles, books, and chapters, including three single-volume editions of Ecology and Classification of North American Freshwater Invertebrates (edited by J.H. Thorp and A.P. Covich). Prof. Thorp is now embarked on a major project to expand from the previous North American emphasis on freshwater invertebrates to the fourth edition’s global coverage of this subject in perhaps nine volumes of Thorp and Covich’s Freshwater Invertebrates.

D. Christopher Rogers is a research zoologist at the University of Kansas with the Kansas Biological Survey and is affiliated with the Biodiversity Institute. Christopher specializes in freshwater crustaceans (particularly the Branchiopoda and the Decapoda) and the invertebrate fauna of seasonally astatic wetlands on a global scale. He has numerous peer reviewed publications in crustacean taxonomy and invertebrate ecology, as well as published popular and scientific field guides and identification manuals to freshwater invertebrates. Christopher is an Associate Editor for the Journal of Crustacean Biology and a founding member of the Southwest Association of Freshwater Invertebrate Taxonomists. He has been involved in aquatic invertebrate conservation efforts all over the world.

Preface to the Fourth Edition

When Alan P. Covich and I initially conceived the first edition of Ecology and Classification of North American Freshwater Invertebrates in the mid-1980s (subsequently published in 1991 by Academic Press), the possibility of a second edition (2001) was only vaguely considered, much less a third (2010). Between the second and third edition, I tried to convince some European scientists to develop independently a similar approach for their fauna, but a lack of interest at that time doomed this proposed Elsevier book project. Less than a year after the third edition, an acquisition editor at Elsevier (Candace Janco) inquired about a fourth edition for 2015 at an annual meeting of the North American Benthological Society (now Society for Freshwater Sciences). Other project commitments prevented Alan from continuing as a coeditor, and I felt that 2015 was too early for a fourth edition without major changes in the scope of the project or detail of the keys. At that point a kernel of what was possibly a creative but certainly audacious idea began to sprout. This soon grew to a proposal I submitted with D. Christopher Rogers to develop an approximately 9-volume series covering inland water invertebrates of the world. (Christopher had coauthored two chapters in the third edition and has now coauthored A Field Guide to Freshwater Invertebrates of North America with me.) Although the prospectus ultimately submitted to Elsevier included books covering all major zoogeographic regions, the initial contract was limited to three volumes encompassing the world’s major book markets (Europe and the USA).

While I am the sole editor of the book series at this time, Christopher has been a major partner in developing ideas for the fourth edition and is an editor on the other volumes (senior editor on the third). As we made significant progress on the first three volumes, we began contacting some potential coeditors and authors to develop volumes for other zoogeographic regions and negotiations with some of those volumes are now underway.

Based on my feelings and recommendations of Academic Press, I have named our book series Thorp and Covich’s Freshwater Invertebrates in order to: (a) associate present with past editions, unite current volumes in the fourth edition, and link to possible future editions; (b) establish a connection between the ecological and general biology coverage in volume I with the taxonomic keys in the remaining volumes; and (c) give credit to Alan Covich for his work on the first three editions. For the sake of brevity, I refer to the current edition as T&C IV. Whether T&C V will ever appear is certainly problematic, but who knows!

Our concept for T&C IV included producing one book (volume I) with 6 chapters on general environmental issues applicable to many invertebrates, followed by 35 chapters devoted to individual taxa at various levels (order to phylum, or even multiple phyla in the case of the protozoa). Volume I was designed both as an independent book on ecology and general biology of various invertebrate taxa and as a companion volume for users of the keys in the regional taxonomic volumes, thereby reducing the amount of duplicate information needed for the taxonomic volumes.

The perhaps eight taxonomic volumes will contain both keys for identifying invertebrates in specific zoogeographic regions and descriptions of detailed anatomical features needed to employ those keys. The multilevel keys are formatted to enable users to work easily at the level of their project/course need and their scientific experience. For that reason, we separated keys by major taxonomic divisions. For example, a student in a college course might work through one or more of the initial crustacean keys to determine the family to which a freshwater shrimp belongs. In contrast, someone working on an environmental monitoring project might need to identify a crayfish to genus or even species, and thus would use the relevant, detailed keys that require more background experience. We have also designed the keys, where possible, to proceed from a general to a specific character within a couplet. These changes to the key are one of Christopher’s major contributions to T&C IV.

While the vast majority of authors in T&C I–III were from the USA or Canada, we attempted in T&C IV to attract authors from many additional countries in six continents. Although we largely succeeded in this goal, we expect future editions of T&C to continue increasing the proportion of authors from outside North America as our books become better known internationally.

Our goals for T&C IV are to improve the state of taxonomic and ecological knowledge of inland water invertebrates, to help protect our aquatic biodiversity, and to encourage more students to devote their careers to working with these fascinating organisms. These goals are especially important because the verified and probable losses of species in wetlands, ponds, lakes, creeks, and rivers around the globe exceed those in most terrestrial habitats.

James H. Thorp

Editor-in-Chief, T&C IV

Preface to Volume I

Readers familiar with previous three editions of the Thorp and Covich (T&C) volumes will notice a dramatic change in T&C IV, as we have expanded from a North American concentration to worldwide coverage. This volume consists of 6 general chapters on inland water habitats and invertebrates, and 35 chapters on the ecology and general biology of specific groups. The greatest difference between this volume of T&C IV and the chapters in T&C III is that all taxonomic keys have been shifted to separate volumes. This has allowed an increase in the ecological coverage in volume I and more detailed keys in subsequent volumes. This enabled us to expand from the North American-centric coverage in previous editions to the global ecological perspective characterizing T&C IV, which is one of the reasons we increased the international representation of authors. In the process we added some groups not occurring in the Nearctic (e.g., aquatic millipedes) or whose numbers were not large in the Nearctic (e.g., freshwater crabs and shrimp, which dominate tropical freshwater habitats). Readers of the earlier editions will note that we have expanded coverage of the ecology and general biology of insects from a single chapter in T&C III to 9 chapters in volume I. However, the taxonomic keys in Volume II for the Nearctic Region remain limited to family level to avoid duplication with the excellent existing text on North American aquatic insects by Merritt, Cummins, and Berg (2008). The literature-cited sections in volume I represent a compromise between T&C II and III, in that we have returned to inclusion of author names and dates in the text (rather than just numbered citations) and have expanded the number of allowed citations at the end of each chapter, as in T&C II; however, the average number of references per chapter in this printed volume is often less than in the first two volumes in order to save space. T&C IV continues our policy from the previous edition of being in color for volume I; indeed, we have considerably augmented the number of color figures. We believe this helps students, in particular, appreciate better that bright colorful invertebrates are not entirely limited to marine and terrestrial habitats.

This edition is strongly focused on species found in fresh through saline inland waters, with a nonexclusive emphasis on surface waters. Again, most estuarine and parasitic species are not covered in this book, but we do discuss species whose life cycles include a free-living stage (e.g., Nematomorpha) and species that live in hard freshwaters through to brackish waters even though they may be normally associated with estuarine or marine habitats in some parts of their life cycles (e.g., some shrimp and crabs).

It is our hope that scientists and students from around the world will enjoy this volume, but it is also important that we interest the younger generation (especially the kindergarten through middle school group of children) in freshwater invertebrates. One way that this could happen is to present them with interesting books to read on these organisms and freshwaters in general. Few have been written, but one of the more interesting ones includes biology, ecology, and even rudimentary taxonomy! That book is entitled The Secret Life of Streams and was authored by Lynell Marie Garfield, with contributions from Daniel Devine, Jason Barnes, and Sandra L. Silva (see www.clearmountainstream.com). This charming children’s book was published in 2013 by Lucy Bat Books (www.luckybatbooks.com) and has an ISBN number of 1-9390-5133-9). We strongly recommend you consider getting this book for your younger children.

Editors, Volume I

James H. Thorp,  and D. Christopher Rogers

Section I

Introduction

Outline

Chapter 1. Introduction to Invertebrates of Inland Waters

Chapter 2. Overview of Inland Water Habitats

Chapter 3. Collecting, Preserving, and Culturing Invertebrates

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, KS, USA

D. Christopher Rogers     Kansas Biological Survey and Biodiversity Institute, University of Kansas, Lawrence, KS, USA

Walter W. Dimmick     Lawrence, KS, USA

Abstract

This introductory chapter to Volume I of the fourth edition of Thorp and Covich’s Freshwater Invertebrates discusses alternative definitions of species (biological, phylogenetic, evolutionary, and cohesion), the importance of understanding phylogenies when studying the ecology of individual species and communities, the construction of phylogenetic trees, and the use of systematics and taxonomies in classifying organisms. This chapter concludes with a key to major groups of inland water invertebrates (phyla and classes in some cases) of the world. This key is meant to provide assistance to readers trying to decide which group they are examining and to point to the correct chapter for more information.

Keywords

Global ecology; Keys to phyla and classes; Phylogenetic analyses; Phylogenetic trees; Species concepts; Systematics; Taxonomy

Chapter Outline

Introduction 3

Species and Phylogenies 4

Species and Concepts of Species 4

Biological Species Concept 5

Phylogenetic Species Concept 6

Evolutionary Species Concept 6

Cohesion Species Concept 6

The Role of Phylogenies in Studies of Ecology and Behavior 7

Phylogenetic Trees 8

Using Systematics and Taxonomy to Classify Organisms 9

Taxonomic Keys to Invertebrates of Inland Waters 10

Key to Freshwater Invertebrates 11

Mollusca Classes 11

Annelid Groups 12

Arthropod Subphyla 12

Tetraconata: Crustacean Classes and Malacostracan Orders 12

Tetraconata: Hexapoda Classes 13

Tetraconata: Hexapoda: Insecta Orders 13

References 21

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 pill bug curled in a ball within their palm, ants marching steadily onward with heavy loads, anemones waving their tentacles in a tide pool, or a crayfish scurrying between refuges under stream rocks. It is true that many inland water invertebrates may initially seem drab and uninspiring next to their more colorful and often larger marine relatives. Yet, once the curious observer penetrates beyond superficial appearances and thoroughly examines their diverse structural, physiological, behavioral, and general ecological adaptations, few fail to be impressed by these fascinating freshwater creatures. This childhood fascination led progressively, if not inevitably, to collegiate studies of invertebrates and then to a life-long career in science for many authors contributing to this and other volumes in our fourth edition.

Aquatic scientists focusing on invertebrates conduct research in a wide diversity of fields. Some are fascinated by the diversity of invertebrates and the relationships among species over evolutionary history, or they may be involved in conducting environmental assessments in which knowledge of invertebrate identification is typically essential. Others seek to mitigate human diseases and parasitism by understanding and controlling those where freshwater invertebrates play a role. Many more study the population biology or culturing of invertebrates involved in food webs, which contribute directly (freshwater crayfish) or indirectly (such as zooplankton prey of lake fish) to human diets. Many more investigate the ecology of invertebrates either from an environmental monitoring perspective or just to satisfy their scientific curiosity.

In this chapter, we discuss invertebrate taxonomy and classification. We also include a taxonomic key to help the reader begin the process of identifying inland water invertebrates at higher taxonomic levels. This dichotomous key will lead to the appropriate chapter in volume I for information on the ecology and general biology of various groups and to more detailed identification keys in the other volumes of this series. Additional information on freshwater invertebrates can be found in various textbooks on invertebrate zoology and aquatic ecology. For amateur invertebrate zoologists and students needing initial guidance to species in the field and laboratory, we recommend the use of more introductory texts, such the Field Guide to Freshwater Invertebrates of North America (Thorp and Rogers, 2011).

Species and Phylogenies

The identification and definition of natural biological units is one of the major guiding principles for biologists that endeavor to study, discover, and document the world’s biodiversity. Unless you properly identify the organism you are studying, you will be hindered in drawing any justifiable conclusions. Genera and species are examples of natural biological units as they result from evolution. The formal name of a natural biological unit is a taxon. In contrast, a population is an example of a qualitative unit that is often not a natural unit but rather a convenient partition of geographic variation.

The principal intellectual tools used to define natural taxa have emerged from the scientific disciplines of taxonomy, systematic biology, and population genetics. Species concepts and phylogenetic analyses are central to systematic biology. Population genetics is a method of direct study of connectivity between populations. Sometimes, analyses of molecular variation enable the discovery of cryptic species, which are difficult to distinguish by morphological characters alone. This has resulted in the recognition of multiple species that were formerly thought to be a single wide-ranging species.

There are several different species concepts, and no one species concept is necessarily better than another. There are various groups of organisms wherein one concept works better than another, depending on the evolution of that group. Similarly, most species can be defined using two or more species concepts. These species concepts are not competing concepts, but rather different tools to be used sometimes in concert, sometimes individually, to describe and define biodiversity. In fact, the appropriateness of a particular species concept can vary with the evolutionary patterns of that group, and most taxa can be defined using two or more species concepts. Although this situation may seem inconvenient for a new student or a researcher who is primarily interested in ecological or behavioral studies, an informed viewpoint about species concepts and phylogenetics can only help a serious scholar of invertebrates. Cataloging biodiversity is paramount in our modern world, given the alarming amount of local extirpation and global extinction of species (particularly because of anthropogenic factors).

Although it may appear that taxonomy changes capriciously, the real issue is that taxonomic concepts are hypotheses to be tested. A species may move from one genus to another or move from species to subspecies and back as new data becomes available. The goal is to have taxonomic and systematic classification and hierarchy converge with biological reality.

Species and Concepts of Species

Evolution is a phenomenon of lineages, and the termini of these lineages are called species. Species are the result of historical natural processes (Darwin, 1859). Anagenesis is the modification of lineages through mutation, gene flow, natural selection, and random genetic drift. Cladogenesis is the creation of a new lineage by splitting a preexisting lineage or the result of hybridization between two preexisting lineages. Consequently, the first logical step to understanding current and past biodiversity is the identification of species and other natural units that result from evolution (Mayden and Wood, 1995).

Scientists often need to identify biodiversity units below the genus level, but ultimately, unit definitions must be based on appropriate species concepts. Species concepts play a critical role in the interpretation of variation because species provide an upper boundary for population comparisons. Whether a particular population or group of populations is distinctive enough to be recognized as a species continues to be a key issue in many taxonomic and conservation disputes.

Taxonomists identify and describe species, and classify them according to a hierarchy. This is accomplished by following the scientific method, as is any branch of science. A species is a name for a discrete group of organisms (see species concepts below). The definition of that name is, in essence, a theory, constructed by the taxonomist based on available morphological, molecular, behavioral, and/or ecological data. Although these theories are meant to be tested, as are all scientific theories, it should be remembered that species are unstable through time unlike most scientific subjects. They evolved from other species, they are still under selective pressures, and they may evolve away from the original described theory or may become extinct or both.

The scientific literature about the nature of species and the best method of identifying species is enormous and was previously contentious. Species concepts almost always fall into one of two categories: operational species concepts or ontological species concepts. Operational species concepts provide specific criteria to determine whether or not two different populations belong to the same species. The biological species concept of Mayr (1942) is a classic example of an operational species concept. The operation used by Mayr is the discovery of barriers to sexual reproduction that are assumed to define species boundaries. Ontological species concepts provide a theoretical species definition, but do not specify a method for their identification. Wiley’s (1981) evolutionary species concept is an example of an ontological species concept. Species are defined as lineages expected to persist through time, but no method (operation) is prescribed for the discovery of these lineages. In short, ontological concepts can be understood as theoretical species definitions. Operational species concepts provide only a method of how to discover species and are, by definition, limited by their particular methodology.

Below are brief definitions of the biological, phylogenetic, cohesion, and evolutionary species concepts along with comments on their strengths and limitations. These influential concepts provide a good basis for understanding many issues regarding species concepts. For a more thorough discussion and evaluation of species concepts, we recommend an article by Mayden and Wood (1995). Suffice to say that not all species concepts will work with all groups, some groups can be defined by only one concept, some groups have different species depending on which concept is used, and still other groups have species that are defined the same way regardless of which concepts is used. This is a reflection of the diversity of life and the diversity of ways that species can evolve. 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. Because biological diversity is enormous, it seems highly unlikely that any single operational concept could be devised to encompass all species.

Biological Species Concept

Species are groups of actually or potentially interbreeding populations that are reproductively isolated from other such groups (Mayr, 1942).

This 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 organismal 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 (O’Brien and Mayr, 1991).

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 (Dobzhansky, 1935, 1970). 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 gene flow barriers that define the boundaries of the reproductive community and gene pool thereby preserving the integrity of the species’ genetic system. In practice, however, isolating mechanisms are rarely studied and species are usually diagnosed by differences in morphological features.

A fundamental drawback to the biological species is the concept is that it is exclusively defined in terms of sexual reproduction. Asexual and cyclically parthenogenetic 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. 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 (Templeton, 1989) from the viewpoint of population genetics. 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. Freely hybridizing species are known from plants, insects, and vertebrates (Templeton, 1989).

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 isolation of a subpopulation from the main 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. However, isolation mechanisms can be geographical, temporal, or behavioral. In each case, it is a barrier to reproduction and gene flow. Hence, the evolutionary forces responsible for allopatric speciation may not be influenced by the isolating mechanisms that are a fundamental aspect of the biological species concept.

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 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).

Phylogenetic Species Concept

A species is an irreducible cluster of organisms diagnostically distinct from other such clusters and containing a parental pattern of ancestry and descent (Cracraft, 1989).

Phylogenetic methods focus primarily on evolutionary relationships and ancestry. Several species concepts have been proposed that are based on the methods and philosophy of phylogenetic systematics (Cracraft, 1989; Nixon and Wheeler, 1990; Nelson and Platnick, 1991). The definition presented above was proposed by Cracraft (1989), 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, molecular (e.g., allozymes, mtDNA haplotypes, nucleic acid sequences, and amino acid sequences), behavioral, and ecological characters are used to diagnose different species. Individual characters or groups of characters that are determined to be primitive or shared as a common character between groups are called plesiomorphies. Characters that are determined to be recently derived and unique to a lineage are called apomorphies. 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, in part, 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.

Even though phylogenetic species concepts have several positive features, several drawbacks are inherent to these concepts (Mayden and Wood, 1995). One serious disadvantage is that all species are considered to be evolutionary units that are equivalent in comparative biological studies (Cracraft, 1983). However, not all species or any taxonomic level are equal, in that each has different ecological roles, functions, and evolutionary histories. Another disadvantage is that these concepts are prone to either underestimate or seriously overestimate the number of species present. Finally, it is frequently difficult to tell if a character is apomorphic or plesiomorphic, or even if that character evolved more than once. In these situations, a misunderstanding of the characters may result in obscured relationships or present false relationships.

Evolutionary Species Concept

A species is a single lineage of ancestor–descendant populations that maintains its identity from other such lineages and has its own evolutionary tendencies and historical fate (Wiley, 1981).

The evolutionary species concept was refined and popularized in Wiley’s book (1981) on the principles of phylogenetic systematics. The evolutionary and phylogenetic concepts are related but different. The evolutionary species concept is ontological and simply states that individual species exist as lineages of ancestor–descendant populations which maintain their own characteristic identity with its peculiar evolutionary tendencies and historical fate. It does not specify a method for identifying species, in contrast to the phylogenetic species concept, which is operational rather than ontological.

The evolutionary concept is advantageous because it is applicable to living, extinct, sexual, and asexual groups. It also emphasizes that species can be maintained 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 species descriptions 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 founded on phenotypic continuity or discontinuity and not on comparisons of gene flow among populations.

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. That is, it is limited to diagnosable species—an action predicated on human abilities.

Cohesion Species Concept

A species is the most inclusive group of organisms having the potential for demographic and genetic exchangeability (Templeton, 1989). (Genetic exchangeability is the exchange of genes between individuals through sexual reproduction.)

The cohesion species concept was proposed and described in detail by Templeton (1989). His concept emphasizes the evolutionary processes that hold evolutionary lineages (species) together through time. Because biological species are defined by gene flow, the evolutionary processes (e.g., isolating mechanisms) that curtail or promote gene flow define the boundaries of sexually reproducing species. Templeton pointed out, however, that gene flow is only one of the microevolutionary forces acting on populations and that random genetic drift, demography, and natural selection also have a role in defining evolutionary lineages because they can act on both sexual and asexual populations.

Because 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. Thus, 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 no longer exhibit complete demographic exchangeability, their cohesion to the overall population has consequently been diminished. Templeton’s (1989) 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 greatly illuminated the mechanisms of cohesion, but his cohesion concept is really contained within the framework of the evolutionary 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 (Mayden and Wood, 1995). Nevertheless, the cohesion concept articulated by Templeton (1989) complements the evolutionary species concept by identifying the mechanisms that are responsible for the speciation process. Templeton’s cohesion mechanisms deserve careful consideration because they play a critical role in the process of speciation and generation of biodiversity. Students are encouraged to consider Templeton’s 1989 article because it is especially relevant to the diverse reproductive strategies of aquatic species of invertebrates.

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. Phylogenies play a critical role because they provide an intellectual framework for comparative studies of morphology, behavior, and ecology at various levels. 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 (Brooks and McLennan, 1991). Although some of the reasons that a phylogenetic basis for comparative studies is needed are summarized below, the landmark book by Brooks and McLennan (1991) should be consulted for comparative studies of species.

Phylogenetic analyses are perhaps the best way to discover biologically meaningful groups of species that are statistically comparable. (Many ecological questions can, of course, be studied at community and greater organizational levels using genera or even higher taxonomic levels.) Although statistical techniques routinely assume that samples are independently drawn from a model distribution of samples, species are not independent and 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 connections. 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, both the pattern and timing of speciation and character evolution must be considered because all species are a mosaic of characters that have evolved over time. 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!

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. No repeatable methods were widely used to formulate phylogenetic hypotheses. The development of numerical taxonomy in 1957 in separate articles by Charles Michener and Peter Sneath and later extensive elaboration by Robert Sokal and Peter Sneath 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 fundamental problem with this approach is that the choice of characters and the method of weighing character values were based entirely upon the type of data selected for analyses. Although the method of phenogram development may have been objective, the data were still subjective.

The idea of basing phylogenetic hypotheses on overall similarity was soon challenged by the emerging discipline of cladistics. Over a decade of often rancorous debate ensued between the disciples of phenetics and cladistics following the 1966 English translation of Hennig’s Phylogenetic Systematics. 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 because of convergent or parallel evolution. This type of similarity is not only devoid of phylogenetic information, it is also likely to be positively misleading because a character or an attribute of two or more species may be similar because of 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. In contrast, 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 a 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