Introduction to Taxonomy of Angiosperms

B.K. Verma Introduction to TAXONOMY OF ANGIOSPERMS

B.K. VERMA Former Professor and Head Department of Botany University of Allahabad Allahabad

New Delhi-110001 2011 INTRODUCTION TO TAXONOMY OF ANGIOSPERMS B.K. Verma

© 2011 by PHI Learning Private Limited, New Delhi. All rights reserved. No part of this book may be reproduced in any form, by mimeograph or any other means, without permission in writing from the publisher.

ISBN-978-81-203-4114-2

The export rights of this book are vested solely with the publisher.

Published by Asoke K. Ghosh, PHI Learning Private Limited, M-97, Connaught Circus, New Delhi-110001 and Printed by Rajkamal Electric Press, Plot No. 2, Phase IV, HSIDC, Kundli-131028, Sonepat, Haryana. Contents

Preface ix

List of Families xi

Chapter 1 INTRODUCTION 1–8 1.1 AIMS OF TAXONOMY 2 1.2 DEFINITION OF TERMS 2 1.2.1 Identification 2 1.2.2 Nomenclature 3 1.2.3 Classification 3 1.2.4 Documentation 4 1.2.5 Units of Classification and Their Concepts 4 1.3 MAJOR UNITS 4 1.3.1 Division 4 1.3.2 Class 5 1.3.3 Order 5 1.3.4 Family 5 1.4 MINOR UNITS 6 1.4.1 Genus 6 1.4.2 Species 7 1.4.3 Subspecies 8 1.4.4 Variety 8 1.4.5 Forma 8

Chapter 2 HISTORY OF PLANT CLASSIFICATION 9–18 2.1 PRE-DARWINIAN SYSTEMS 10 2.1.1 The Age of Herbals 11 2.1.2 Sexual System 12 2.1.3 Systems Based on Form Relationships 13

iii iv Contents

2.2 POST-DARWINIAN SYSTEMS 14 2.2.1 Systems Based on Phylogeny 14 2.3 TAXONOMY IN INDIA 17 2.3.1 Ancient India 17 2.3.2 16th Century Onwards 18 2.3.3 Post-Linnean Botany 18

Chapter 3 IMPORTANT SYSTEMS OF CLASSIFICATION 20–73 3.1 BENTHAM AND HOOKER’S SYSTEM 20 3.1.1 Class I: 21 3.1.2 Class II: Gynospermae 24 3.1.3 Class III: 25 3.1.4 Merits and Demerits of the System 25 3.2 ENGLER AND PRANTL’S SYSTEM OF CLASSIFICATION 27 3.2.1 Class I: Monocotyledoneae 28 3.2.2 Class II: Dicotyledoneae 29 3.2.3 Merits and Demerits of the System 32 3.3 HUTCHINSON’S SYSTEM OF CLASSIFICATION 33 3.3.1 Outline of Classification 35 3.3.2 Merits and Demerits of the System 42 3.4 BESSEY’S SYSTEM OF CLASSIFICATION 43 3.4.1 Phylum: Anthophyta (Angiosperms) 45 3.5 TAKHTAJAN’S SYSTEM OF CLASSIFICATION 49 3.5.1 Division—Magnoliophyta (Angiospermae) 51 3.5.2 Merits and Demerits of Takhtajan’s System of Classification 58 3.6 CRONQUIST’S SYSTEM OF CLASSIFICATION 59 3.6.1 Division: Magnoliophyta (Angiosperms) 60 3.6.2 Differences between Systems of 1968 and 1981 66 3.6.3 Merits and Demerits of Cronquist’s System of Classification 66 3.7 WETTSTEIN’S SYSTEM OF CLASSIFICATION 68 3.7.1 Class I: Dicotyledons 68 3.7.2 Class II: Monocotyledons 71 3.7.3 Merits and Demerits of Wettstein’s System of Classification 72

Chapter 4 PLANT NOMENCLATURE 74–82 4.1 FIRST CODE OF NOMENCLATURE ‘THE PARIS CODE’, 1867 75 4.2 IMPORTANT AND MAJOR RULES OF NOMENCLATURE 75 4.2.1 Rank of Taxa 75 4.2.2 Typification (Articles 7–10) 76 4.3 PRINCIPLES OF PRIORITY (Articles 11–15) 78 4.4 EFFECTIVE AND VALID PUBLICATIONS (Articles 29–45) 78 4.5 AUTHOR CITATION (Articles 46–50) 79 Contents v

4.6 RETENTION AND CHOICE OF NAMES 80 4.7 REJECTION OF NAMES 80 4.8 SOME IMPORTANT DATES IN BOTANICAL NOMENCLATURE 81

Chapter 5 CURRENT TRENDS IN PLANT TAXONOMY 83–121

5.1 ANATOMY AND TAXONOMY 83 5.1.1 Anatomical Characters 83 5.2 EMBRYOLOGY AND TAXONOMY 91 5.2.1 Use of Embryological Characters Among Major Taxa 93 5.2.2 At Family Level 94 5.2.3 At Generic Level 98 5.3 PALYNOLOGY AND TAXONOMY 99 5.3.1 Taxonomic Characters of Pollen Grains 99 5.3.2 Use of Pollen Character in Plant Taxonomy 102 5.4 CHEMISTRY AND TAXONOMY 104 5.4.1 Directly Visible Characters 105 5.4.2 Plant Products 106 5.4.3 Serum Analysis or Serology 109 5.4.4 Biosynthetic Pathways 109 5.4.5 DNA Hybridization 110 5.5 CYTOLOGY AND TAXONOMY 111 5.5.1 Chromosome Number 111 5.5.2 Chromosome Morphology 113 5.5.3 Chromosome Behaviour at Meiosis and in Crosses 114 5.5.4 Heterochromatin 114 5.5.5 Use of Cytological Data at Family Level 115 5.5.6 Use of Cytological Data at Genus Level and Below 117 5.6 ECOLOGY AND TAXONOMY 117 5.6.1 Phenotypic Plasticity 118 5.6.2 Ecological Factors and Genotype 119 5.6.3 Genotype and Physical Environment 119 5.6.4 Genotype and the Biotic Environment 120 5.6.5 Ecotypic Variations 120

Chapter 6 SELECTED FAMILIES OF ANGIOSPERMS: DICOTYLEDONS 122–338 6.1 POLYPETALAE (Thalmiflorae) 122 6.1.1 Order— 122 6.1.2 Order—Parietales 145 6.1.3 Order—Caryophyllineae 167 6.1.4 Order—Guttiferales 172 6.1.5 Order— 176 vi Contents

6.2 POLYPETALAE (Disciflorae) 186 6.2.1 Order— 186 6.2.2 Order— 194 6.2.3 Order— 197 6.3 POLYPETALAE (Calyciflorae) 201 6.3.1 Order— 201 6.3.2 Order— 217 6.3.3 Order—Passiflorales 221 6.3.4 Order—Umbellales 229 6.4 SUB-CLASS—GAMOPETALAE 234 6.4.1 Order—Rubiales 234 6.4.2 Order— 240 6.4.3 Order—Primulales 252 6.4.4 Order— 255 6.4.5 Order—Polemoniales 265 6.4.6 Order—Personales 282 6.4.7 Order— 298 6.5 MONOCHLAMYDEAE 309 6.5.1 Series—Curvembryae 309 6.5.2 Series—Unisexuales 321

Chapter 7 SELECTED FAMILIES OF ANGIOSPERMS: MONOCOTYLEDONS 339–390

7.1 MONOCOTYLEDONEAE (Ovary Inferior) 339 7.1.1 Series—Microspermae 339 7.1.2 Series—Epigynae 345 7.2 MONOCOTYLEDONEAE (Ovary Superior) 362 7.2.1 Series—Coronarieae 362 7.2.2 Series—Calycinae 367 7.2.3 Series—Nudiflorae 372 7.2.4 Series—Glumaceae 376

Chapter 8 ORIGIN OF ANGIOSPERMS 391–405

8.1 THE ANTIQUITY (TIME) OF ORIGIN OF ANGIOSPERMS 391 8.1.1 Lower Cretaceous Angiosperms 394 8.2 THE CRADLE (PLACE) OF THE ANGIOSPERMS 396 8.2.1 Greenland as the Cradle of Angiosperms 396 8.2.2 Tropical Region as the Site of Origin 396 8.2.3 Pacific Islands as the Cradle of Angiosperms 396 8.2.4 South Africa as the Cradle of Angiosperms 396 8.3 THE ANCESTORS OF ANGIOSPERMS 397 8.3.1 Bennettitalean Theory 397 8.3.2 Caytonialean Theory 398 Contents vii

8.3.3 Pteridosperm Theory 400 8.3.4 Pentoxylales Theory 400 8.3.5 Dirhopalostachyacean Theory 401 8.3.6 Czekanowskialean Theory 401 8.3.7 Gonophyll Theory 402 8.3.8 Coniferales—Amentiferae Theory 402 8.3.9 Ephedrales Theory 402 8.3.10 Gnetales—Angiosperm Theory 403 8.3.11 Durian Theory 403 8.3.12 Gigantopteridalean Theory 404

Chapter 9 FIELD AND HERBARIUM TECHNIQUES 406–413

9.1 PURPOSES OF MODERN HERBARIUM 406 9.2 DIFFERENT TYPES OF HERBARIA 407 9.2.1 National or Major Herbaria 407 9.2.2 Regional Herbaria 407 9.2.3 Local Herbaria 407 9.3 IMPORTANCE OF HERBARIA IN BOTANICAL RESEARCH 407 9.3.1 Anatomy 407 9.3.2 Palynology 408 9.3.3 Physiology, Biochemistry and Chemotaxonomy 408 9.3.4 Biosytematics 408 9.4 HERBARIUM TECHNIQUES 409 9.4.1 Collection 409 9.4.2 Pressing 410 9.4.3 Drying 410 9.4.4 Poisoning 410 9.4.5 Mounting 411 9.4.6 Labelling 411 9.4.7 Storage and Arrangement of Herbarium Sheets in a Herbarium 411 9.4.8 Fumigation 412

Appendix I Key to Families Described in the Book 415–418

Appendix II Pro Forma for the Description of a Flowering Plant 419–422

Appendix III Synopsis of Families and Suprafamilial Taxa of Magnoliophyta 423–436

Glossary 437–446

References 447–469

Index of Botanical Names 471–489

Index 491–492 Preface

I wrote a book under the title Introduction to Taxonomy of Flowering Plants in 1976, wherein thirty-three families and three well known systems of classification were covered. It was reprinted in 1982. Due to several unavoidable circumstances, it was not possible to bring out further editions and that title had to be abondoned. In the present book, I have included fifty families and seven systems of classification and have tried to cover the syllabi of most of the Indian universities. The book has been written basically to cater to the needs of undergraduate students, but there are chapters like plant nomenclature, current trends in plant taxonomy, origin of angiosperms, etc. which may be of great use to postgraduate students and also to candidates studying for various competitive examinations. The families have been arranged according to the Bentham and Hooker’s system of classification. The number of genera and species at world level is based on Willis—Dictionary of Flowering Plants and Ferns, 8th edition, and that of India, on Flora of India—Introductory volume, published by Botanical Survey of India (BSI). In each family, information about endemic taxa is given after its general account, and one or two plants have been described for the benefit of students for their practical work. In most cases, the descriptions are based on those given in Flora of Allahabad District (Misra and Verma 1992) and also those based on personal observations. I am thankful to Professor Nupur Bhowmick, Professor D.K. Chauhan and Dr. S.P. Tiwari for providing literature, specially in connection with family and origin of angiosperms. I am also thankful to Professor G.K. Srivastava, Professor G.L. Tiwari, Professor Anupam Dixit of University of Allahabad, and Dr. P.C. Srivastava of Ewing Christian College (ECC), for their valuable help. Dr. R.C. Srivastava of Botanical Survey of India, provided information about Indian plants. His kind help is gratefully acknowledged. I am indebted to Dr. K.K. Khanna, Dr. Anand Kumar and Dr. Vinay Ranjan of Botanical Survey of India for their manifold help. I am also grateful to Dr. Satya Narain and Dr. H.P. Pandey for their kind help in preparation of the book. Mr. Pramod Kumar and Mr. Om Prakash, research scholars of Duthie Herbarium, also helped me immensely. I express my gratefulness to them. Dr. B.K. Misra, Brahmanand College, Kanpur, was kind enough to go through a few chapters and provide helpful suggestions. I thank him for his helpful advice. I am deeply indebted to my wife, Pratibha, who has been a constant source of encouragement to me, resulting in successful culmination of this task. She also took great ix x Preface pains in preparation of the index. My son Abhishek and daughter-in-law Manjaree helped me in various ways. Their help is gratefully acknowledged. My granddaughter Aditi and grandsons Akshay and Amber deserve special mention; they all created an affectionate atmosphere in the family to take up this work. My daughters Anju and Renu and son-in-laws Amit and Ashutosh always encouraged me for this work. I also express my gratitude to them. I thank Mr. H.C. Joshi, Mr. C.P. Joshi, Mrs. Subhada Bhargava and Mr. Radhey Shyam for their support in connection with library. Finally, I express my deep sense of gratitude to Shri Asoke K. Ghosh, Chairman and Managing Director, PHI Learning, New Delhi, for undertaking the publication of my manuscript. I am also indebted to Ms. Pushpita Ghosh, Managing Editor and Marketing Director, Mr. Darshan Kumar, and Mr. K.K. Chaturvedi and others, for taking pains in scrutinizing the manuscript judiciously, and for rendering able expertise and invaluable advise at all stages of the book. In particular, I am most grateful to Dr. Aerum Khan for carefully and critically editing the manuscript. Suggestions, if any, for improvement of the book will be gratefully appreciated.

B.K. VERMA Introduction To Taxonomy Of Angiosperms

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