DESCRIBING SPECIES Practical Taxonomic Procedure for Biologists Judith E. Winston COLUMBIA UNIVERSITY PRESS NEW YORK Columbia University Press Publishers Since 1893 New York Chichester, West Sussex Copyright © 1999 Columbia University Press All rights reserved Library of Congress Cataloging-in-Publication Data © Winston, Judith E. Describing species : practical taxonomic procedure for biologists / Judith E. Winston, p. cm. Includes bibliographical references and index. ISBN 0-231-06824-7 (alk. paper)—0-231-06825-5 (pbk.: alk. paper) 1. Biology—Classification. 2. Species. I. Title. QH83.W57 1999 570'.1'2—dc21 99-14019 Casebound editions of Columbia University Press books are printed on permanent and durable acid-free paper. Printed in the United States of America c 10 98765432 p 10 98765432 The Far Side by Gary Larson "I'm one of those species they describe as 'awkward on land." Gary Larson cartoon celebrates species description, an important and still unfinished aspect of taxonomy. THE FAR SIDE © 1988 FARWORKS, INC. Used by permission. All rights reserved. Universal Press Syndicate DESCRIBING SPECIES For my daughter, Eliza, who has grown up (andput up) with this book Contents List of Illustrations xiii List of Tables xvii Preface xix Part One: Introduction 1 CHAPTER 1. INTRODUCTION 3 Describing the Living World 3 Why Is Species Description Necessary? 4 How New Species Are Described 8 Scope and Organization of This Book 12 The Pleasures of Systematics 14 Sources CHAPTER 2. BIOLOGICAL NOMENCLATURE 19 Humans as Taxonomists 19 Biological Nomenclature 21 Folk Taxonomy 23 Binomial Nomenclature 25 Development of Codes of Nomenclature 26 The Current Codes of Nomenclature 50 Future of the Codes 36 Sources 39 Part Two: Recognizing Species 41 CHAPTER 3. SPECIES AND THEIR DISCOVERY 43 Species Concepts 44 viii CONTENTS Processes Affecting Speciation 46 Taxonomic Characters Examples of Ways in Which Biologists Have Discovered New Species 55 Sources 65 CHAPTER 4. ESTABLISHING IDENTITY: THE LITERATURE SEARCH 71 Mistakes and Bad Examples 71 Establishing Identity 73 Where to Find the Taxonomic Literature 75 How to Read the Taxonomic Literature 83 Species Descriptions 83 Taxonomic Literature Searching on the Internet 87 Sources 88 CHAPTER 5. ESTABLISHING IDENTITY: USING MUSEUM COLLECTIONS Collections, Museums, and Herbaria 96 Locating Material 97 Borrowing Material 102 Type Material 103 Visiting Collections: What to Expect and How to Behave 105 Cooperation with Systematists 106 Sources IOJ Part Three: Writing Species Descriptions 113 CHAPTER 6. SPECIES DESCRIPTIONS IN TAXONOMY 1/5 Reasons for Writing Species Descriptions J/J Different Kinds of Taxonomic Publications 116 Form of the Descriptive Paper 125 CHAPTER 7. HEADINGS AND SYNONYMIES 129 Description Headings 129 Synonyms 134 Synonymies 135 New Species 136 Types of Synonymies 137 Terms Used in Synonymies 140 CONTENTS ix Different Kinds of Synonymies 140 References in Headings and Synonymies 144 CHAPTER 8. NAMING SPECIES: ETYMOLOGY 147 Naming Species 147 Brief Review of Latin and Greek 149 Basic Rules of Species Names 153 Descriptive Species Names 156 Geographic Species Names 164 Commemorative Species Names 165 Nonsense Species Names 169 The Etymology Section 169 Sources 171 CHAPTER 9. TYPE AND VOUCHER MATERIAL 173 Rationale for Types and Vouchers 173 Rules of Nomenclature Regarding Types 175 Selection of Types and Vouchers 17s Composition of Type Material 177 Documentation of Type Material 178 Deposition of Types 179 Type Section 182 Sources 185 CHAPTER 10. DIAGNOSIS 189 What Is a Diagnosis? 189 Diagnosis in Zoological Taxonomy 191 Diagnosis in Botanical Taxonomy 191 What Is a Diagnostic Character? 192 The Diagnosis Section: Animals 193 The Diagnosis Section: Plants 197 Additional Uses for Diagnoses 198 CHAPTER 1 1. DESCRIPTION SECTION 201 Descriptive Writing 201 Information Used in the Description Section 203 Writing the Description 210 Telegraphic Style 211 The Description Section: Animals (Examples of Styles for Different Groups) 214 CONTENTS The Description Section: Plants (Examples of Styles for Different Groups) 228 Illustrating Taxonomic Descriptions 231 Sources 238 CHAPTER 12. TAXONOMIC DISCUSSION SECTION 241 Purpose of the Discussion Section 241 Discussion in Descriptions of New Species 242 Evidence to Include 247 Composite Papers 254 The Discussion Section in Other Species Descriptions 256 Taxonomic Ethics 257 CHAPTER 13. THE ECOLOGY SECTION 261 Ecology in Species Descriptions 261 Analysis of Ecological Variation 265 Field Records: Getting the Most from Field Work 265 Ecological Information from Museum Specimens 268 The Ecology Section 269 Sources 273 CHAPTER 14. OCCURRENCE AND DISTRIBUTION 277 Distributional Information in Species Descriptions 277 Parameters of Species Distributions 278 The Distribution Section 281 Distribution Papers 28g Sources 290 CHAPTER 1 5. MATERIAL EXAMINED 293 Practical Value 293 In Original Descriptions 294 In Other Descriptions 295 Material Examined Section 297 Material Examined: Botanical Taxonomy 300 The Material Examined Paper 301 Sources 302 CHAPTER 16. PUBLICATION 303 Criteria of Publication: Zoology 303 Criteria of Publication: Botany 306 CONTENTS xi Preparation of the Manuscript 307 Submission of the Manuscript 311 Final Revision and Publication 312 Journals That Publish Taxonomic Papers 314 Part Four: Beyond Species Description 321 CHAPTER 17. SUBSPECIES 323 Why Are Subspecies Important? 323 Infraspecific Variation 324 Rules of Infraspecific Nomenclature: Zoology 32J Rules of Infraspecific Nomenclature: Botany 329 Deciding When to Name an Infraspecific Taxon 330 Writing Infraspecific Descriptions 332 Sources 334 CHAPTER 18. GENUS-LEVEL DESCRIPTION AND REVISION 337 The Genus Concept 337 When to Describe a New Genus 341 Generic Names 341 Publication of Generic Names 346 Generic Types 346 Examples of Generic-Level Description 349 Problems Caused by Generic Revision 358 Infrageneric Categories and Names 360 Sources 363 CHAPTER 19. KEYS 367 Keys in Taxonomy 367 Key Characters 370 Single-Access (Analytical or Sequential) Keys 371 Multiaccess Keys (Polyclaves) 37s Interactive Identification 378 Key Construction 378 Computerized Key Construction 379 Sources 380 CHAPTER 20. DESCRIPTION OF HIGHER TAXA 383 Family Concepts and Their Use in Taxonomy 384 xii CONTENTS Practical Significance in Biology 385 Describing Families 386 Family-Level Descriptions: Examples 389 Redescriptions of Family-Level Taxa 394 Descriptions of Taxa Above the Family Level 396 Problems with Nomenclature of Higher Taxa 405 Sources 405 CHAPTER 2 1. COMMON PROBLEMS 407 Missing Types 407 Lectotypes 411 Neotypes 416 Necessary Name Changes 422 Replacement Names: Homonymy 424 Conservation of a Name 425 Emendations 427 New Combinations 428 Lack of Information 431 CHAPTER 22. FURTHER STUDIES IN SYSTEMATICS 433 Evolutionary Systematics 434 Phenetics 438 Cladistics 439 Molecular Systematics 446 Biogeography 450 Comparative Biology 451 Sources 451 Literature Cited 455 Subject Index 489 Author Index 503 Taxon Index 513 List of Illustrations Figure 1.1 New equipment and new habitats yield new species Figure 1.2 The Panay cloudrunner, Crateromys heaneyi Figure 1.3 New species from a park, the Virginia Beach bug Figure 1.4 Species description bar graph Figure 2.1 Part of a page from the tenth edition of Linnaeus's Systema Naturae (1758) Figure 3.1 Speciation by cladogenesis Figure 3.2 Patterns of genetic relationships in sexual and asexual populations Figure 3.3 Schematic representation of the decision-making process in species description Figure 4.1 Bad examples. Echinoderm pedicellariae mistakenly described as new species Figure 4.2 Zoological Record Online sample record Figure 4.3 Biological Abstracts on Compact Disc sample record Figure 4.4 Print Version ofBiological Abstracts sample abstract Figure 4.5 Parts of a species description for a new and known species Figure 5.1 Modern compactor shelving holds both old and new specimens Figure 5.2 Computer cataloging and fluid-preserved collection storage Figure 5.3 Fossil collection storage Figure 5.4 Mammal specimens: frozen tissues, skeletons, study skins Figure 5.5 Sample museum loan form xiv LIST OF ILLUSTRATIONS Figure 5.6 Methods of identifying type material in collections Figure 6.1 Rate of description of new taxa Figure 11.1 Regional population variation in fox squirrel pelage color Figure 11.2 Orange and dark purple colonies of the colonial ascidian Distaplia corolla Figure 11.3 Zyzzyzus floridanus, simple line drawing Figure 11.4 Bathypera goreaui, anatomy line drawing Figure 11.5 Fossil insect, twisted-winged parasite, line drawing with stippling Figure 11.6 Comparative plate, characters in species of a new spider genus, Unicorn Figure 11.7 Collospongia auris, composite photographic plate Figure 11.8 Aethopyga linarabonae, photograph of holotype Figure 12.1 Table used to distinguish a new species from others Figure 12.2 Cladogram placing a new species in phylogenetic context Figure 15.1 Locations showing where specimens of two sand lance species were collected Figure 19.1 Portion of a key designed to help identify members of the chief taxonomic subdivisions Figure 19.2 Bracketed key Figure 19.3 Indented key Figure 19.4 Pictorial key Figure 19.5 Branching key Figure 19.6 Matrix key Figure 20.1 Cladogram of families and genera of the superfamily Isochyropsalidoidea Figure 20.2 Strict consensus tree and cladograms from an analysis of the Tropidurus group Figure 21.1 Early "descriptions" of some Mediterranean and Red Sea bryozoans Figure