The Gymnosperms

THE GYMNOSPERMS

Chhaya Biswas D B.M. Johri

Springer-Verlag Berlin Heidelberg GmbH

Cover photograph: Taxus baccata. Courtesy Prof. B.D. Sharma, JNV University, Jodhpur, India

Dr. Cbhaya Biswas Formerly Principal, Gargi College University of Delhi South Campus New Delhi 110 049, India

Prof. (Retired) B.M. Johri Central Reference Library University of Delhi, Delhi 110 007, India

Copyright © I 997 Springer-Verlag Berlin Heidelberg

Originally published by Springer-Verlag Berlin Heidelberg New York in 1997

All rights reserved. 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 permission of the publisher.

Exclusive distribution in North America (including Canada and Mexico), Europe and Japan by Springer-Verlag Berlin Heidelberg New York.

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ISBN 978-3-662-13166-4 ISBN 978-3-662-13164-0 (eBook) DOI 10.1007/978-3-662-13164-0

To Our Teacher

Professor Panchanan Maheshwari (1904-1966)

Preface

The Gymnosperms is a well-illustrated comprehensive account of living and fossil plants of this group. Chapters 1 and 2 give a general account, and describe similarities and dissimilarities with pteridophytes and angiosperms. Chapter 3 deals with classification. The next 18 chapters (4-21) deal sequentially with fossil and living taxa. Phylogenetic relationships are considered for each order. Chapter 22 discusses the in vitro experimental studies on the growth, development and differentiation of vegetative and reproductive organs and tissues. Chapter 23 summarizes the economic importance of gymnosperms. Chapter 24 gives the conciuding remarks.

Thus, there is a complete coverage of significant findings concerning morphology, anatomy, reproduction, development of embryo and seed, cytology, and -evolutionary trends and phylogeny. Ultrastructural and histochemical details are given wherever considered necessary.

There is a comprehensive list of literature citations, and a plant index. This book is essentially meant for the postgraduate students in India and abroad. Undergraduate students can also use it profitably. The entire course should be taught in 25-30 lectures/hours and about 75 hours of field and laboratory work.

CHHAYA BISWAS

BRu MoHAN JoHRI

Acknowledgements

For the preparation of this book we are grateful to: Dr. B.S. Venkatachala, former Director, Birbal Sahni Institute of

Palaeobotany, Lucknow, for allowing Dr. Chhaya Biswas to work in the Library of the Institute, and to Dr. H.K. Maheshwari, Dr. Sheila Chandra, Dr. A.K. Srivastava, Dr. Usha Bajpai, Dr. Sukh Devi, Dr. S.C. Srivastava and Dr. Jayashree Bannerjee for helping her to update the information, fruitful discussions and suggestions concerning fossil members of the gymnosperms, to the Librarian of the Institute for providing the literature.

(The Late) Professor P.N. Mehra, Department of Botany, Punjab University, Chandigarh, for scrutinizing the outlines of this book and giving valuable advice, for providing his monograph on the Indian Conifers, Gnetophytes and Phylogeny of Gymnosperms; Professor B.D. Sharma, Department of Botany, JNV University, Jodhpur, for a critical review of several chapters of the manuscript; to Professor S.P. Bhatnagar, and Dr. S.S. Bhojwani, Department of Botany, University of Delhi, for providing relevant literature and fruitful discussions; Dr. P.S. Srivastava, Department of Botany, Hamdard (deemed) University, New Delhi, for important literature and valuable suggestions; eur colleagues in Gargi College, University of Delhi, Dr. Bharati Bhattacharyya for useful suggestions, Dr. Asha Juneja and Dr. Lalita Sehgal for making available several publications from their personal collections, and Dr. Gita Mathur for scrutinizing some of the chapters; our students Arindam Bhattacharyya and Somdutta Sinha Roy for technical assistance in the preparation of the manuscript.

Professor Upendra Baxi (former Vice-Chancellor), Professor A.L. Nagar (former Pro-Vice-Chancellor), Professor A.P. Srivastava (former Librarian) and Mr. Sher Singh (Deputy Librarian), University of Delhi, for providing an office in the Library where this work was completed.

Mr. Kishen Lal and Mr. S.K. Das for preparing the photographs, Mr. R.K. Gupta for typing the manuscript, and Professor N.N. Bhandari (then Head of the Department of Botany, University of Delhi) for allowing us to use the departmental facilities during the preparation of this book.

Our family members, especially Mr. Alok Biswas and Mrs. Raj Johri, for their encouragement and interest in this work.

We wish to express our gratitude to all the persons and institutions mentioned above.

CHHAYA BISWAS

B.M. JOHRI

Contents

Preface vii Acknowledgements ix

1. Introduction 1-11

Antiquity and Fossil History 1 Geographical Distribution 4

Cycadales and Ginkgoales 4 Pinaceae 6 Taxodiaceae 6 Cupressaceae 7 Podocarpaceae 7 Araucariaceae 7 Cephalotaxaceae 7 Taxales 8 Ephedrales 8 Welwitschiales 8 Gnetales 8

Characteristic Features 8 Gymnosperms and Pteridophytes 9 Gymnosperms and Angiosperms 10

2. Seed Development 12-19

Microsporangium 12 Microsporogenesis 12 Male Gametophyte 13 Ovule 14 Megasporogenesis 14

Megaspore Wall 15 Female Gametophyte 16

Archegonia 16 Pollination and Fertilization 17

Pollination 17 Fertilization 18

Embryo 19 Seed 19

Seed-Coat 19 Temporal Considerations 19

3. Classification 20-23

xii Contents

4. Progymnospermopsida

Phylogenetic Considerations of Archaeopteris 25

5. Gymnospermopsida

Gymnospermopsida-Cycadophytes (Chapters 6-8)

6. Pteridospermales

Lyginopteridaceae 29 Lyginopteris oldhamia (Calymmatotheca hoeninghausi) 29

Habit 29 Root 30 Stem 30 Leaf 30 Sporangia 33 Seed 33

Phylogenetic Considerations 34

7. Cycadeoidales

Cycadeoidaceae 38 Cycadeoidea 38

Stem 38 Leaves 38

Reproduction 39 Phylogenetic Considerations 42

8. Cycadales

Fossil Cycads 44 Nilssonia 46 Baenia 47 Androstrobus manis 47

Living Cycads 47 Root 47 Stem 48 Leaf 49

Reproduction 50 Spermatozoid 51

Ovule 52 Pollen Tube 52 Embryo 54 Chromosome Number 55 Cycadaceae 56 Cycas 56

24-26

27

28-35

36-43

44-81

Morphology 56 Anatomy 59

Root 59 Stem 59 Leaf 61

Reproduction 63 Male Cone 63

Microsporangium 66 Microsporogenesis 66 Male Gametophyte 67

Megasporophyll 68 Ovule 68 Megasporogenesis 71 Female Gametophyte 71 Archegonium 71

Pollination 73 Fertilization 77 Embryogeny 77

Differentiation of Embryo 79 Seed 79 Chromosome Number 80 Temporal Consideration 80 Phylogenetic Considerations 81

Contents xiii

Gymnospermopsida-Gymnosperms of Uncertain Relationship (Chapters 9-12)

9. Caytoniales

Caytoniaceae-Caytonia 82 Leaf 82 Pollen-bearing Organs 82 Ovule-bearing Organs 84

Phylogenetic Considerations 85

10. Glossopteridales

Glossopteridaceae-Glossopteris 86 Root 86 Leaf 88 Male Fructifications 88 Ovulate Structures 89

Phylogenetic Considerations 92

11. Pentoxylales

Pentoxylon 94 Habit 94

82-85

86-93

94-97

xiv Contents

Stem 94 Leaf 95 Microsporangiate Organs 96 Ovulate Cone 96

Phylogenetic Considerations 97

12. Ginkgoales

Fossil Taxa 98 Ginkgoaceae 100 Ginkgo 100 Morphology 100 Anatomy 104

Root 104 Stem 104 Leaf 105

Reproduction 105 Male Cone 105

Microsporangium 105 Microsporogenesis 107 Male Gametophyte 107

Ovule 107 Megasporogenesis 108 Female Gametophyte 109

Pollination 117 Post-Pollination Development of Male Gametophyte 117

Fertilization 121 Embryogeny 122

Differentiation of Embryo 122 Seed 124

Germination 125 Chromosome Number 125 Temporal Considerations 125 Phylogenetic Considerations 125

Gymnospermopsida-Coniferophytes (Chapters 13-16)

98-126

13. Cordaitales 127-136

Cordaitaceae-Cordaites 127 Habit 127 Root 127 Stem 128 Leaf 131

Fructifications 131 Cordaianthus concinus 133 Cordaianthus pseudofluitans 133

Phylogenetic Considerations 134

Contents xv

14. Voltziales 137-143

Morphology 137 Anatomy 138 Reproductive Organs 139 Lebachiaceae 139 Voltziaceae 139

Ullmania 142 Glyptolepis 142 Voltziopsis 142

Phylogenetic Considerations 142

15. Coniferales 144-146

15.1 Pinaceae 146-188

Pinus 146 Morphology 146 An~tomy 150

Root 150 Stem: Shoot Apex 150 Compression Wood 156 Dwarf Shoot 156 Leaf 156

Reproduction 157 Male Cone 157

Microsporangium 158 Pre-meiotic Changes 158 Meiotic Changes 160 Post-meiotic Changes 160 Microsporogenesis 163 Male Gametophyte 166

Female Cone 166 Ovule 166 Megasporogenesis 168 Female Gametophyte (First period of growth) 168

Pollination 168 Post-pollination Development of Male Gametophyte 170 Female Gametophyte (Second period of growth) 171

Fertilization 175 Embryogeny 178

Proembryo 178 Differentiation of Embryo 178

Seed 181 Germination 184

Chromosome Number 185 Temporal Considerations 186

15.2 Taxodiaceae 188-202

Cryptomeria japonica 189

xvi Contents

Morphology 189 Anatomy 189

Stem 189 Leaf 191

Reproduction 191 Male Cone 191

Microsporangium 192 Microsporogenesis 192 Male Gametophyte 192

Female Cone 194 Ovule 194 Megasporogenesis 197 Female Gametophyte 197

Fertilization 199 Embryogeny 199

Diferentiation of Embryo 201 Seed 201

Germination 201 Chromosome Number 201 Temporal Considerations 202

15.3 Cupressaceae

Biota orienta/is 203 Morphology 203 Reproduction 205 Male Cone 205

Microsporangium 205 Microsporogenesis 205 Male Gametophyte 206

Female Cone 207 Ovule 207 Megasporogenesis 207 Female Gametophyte 207 Archegonial Complex 209

Pollination 212 Post-pollination Development of Male Gametophyte 212

Fertilization 213 Embryogeny 215

Differentiation of Embryo 217 Seed 218

Germination 218 Chromosome Number 219 Temporal Considerations 221

15.4 Podocarpaceae

Podocarpus 222 Morphology 222

202-221

221-251

Anatomy 222 Root 222 Stem. Shoot Apex 222 Leaf 224

Reproduction 227 Male Cone 227

Microsporangium 227 Microsporogenesis 231 Male Gametophyte 233

Female Cone 234 Ovule 234 Megasporogenesis 236 Female Gametophyte 236

Pollination 237 Post-Pollination Development of Male Gametophyte 239

Fertilization 242 Embryogeny 242

Differentiation of Embryo 245 Seed 248

Germination 250 Chromosome Number 250 Temporal Considerations 251

15.5 Araucariaceae

Araucaria 252 Morphology 252 Anatomy 254

Stem 254 Leaf 254

Reproduction 257 Male Cone 257

Microsporogenesis 257 Male Gametophyte 257

Female Cone 259 Megasporogenesis 259 Female Gametophyte 260

Pollination 262 Embryogeny 262 Chromosome Number 265

15.6 Cephalotaxaceae

Cephalotaxus 265 Morphology 267 Anatomy 268

Leaf 268 Reproduction 270 Male Cone 270

Contents xvii

251-265

265-290

01iii Contents

Microsporangium 270 Microsporogenesis 271 Male Gametophyte 272

Female Cone 273 Ovule 274 Megasporogenesis 278 Female Gametophyte 278

Pollination 282 Post-pollination Development of Male Gametophyte 282

Fertilization 283 Embryogeny 284 Seed 287

Germination 289 Chromosome Number 289 Temporal Considerations 289

15.7 Phylogenetic Considerations of Coniferales

16. Taxales

Taxaceae 292 Taxus 292 Morphology 292 Anatomy 293

Stem 293 Leaf 294

Reproduction 295 Male Cone 295

Microsporangium 295 Microsporogenesis 295

Ovule 296 Megasporogenesis 298

Pollination 302 Post-pollination Development of Male Gametophyte 302

Fertilization 303 Embryogeny 304 Seed 308

Germination 308 Chromosome Number 308 Temporal Considerations 309 Phylogenetic Considerations 309

17. Gnetopsida

18. Ephedrales

Ephedraceae 312 Ephedra 312

290-291

292-310

311

312-342

Morphology 312 Anatomy 313

Root 313 Stem 314

Reproduction 321 Male Strobilus 321

Microsporangium 321 Microsporogenesis 325 Male Gametophyte 327

Female Strobilus 327 Ovule 327 Female Gametophyte 330

Pollination 335 Fertilization 336 Embryogeny 337 Seed 338 Chromosome Number 341 Temporal Considerations 341

19. Welwitschiales

Welwitschiaceae 343 Welwitschia 343 Morphology 343 Anatomy 346

Root 346 Stem 347 Leaf 349

Reproduction 352 Cone Bracts 352

Male Strobilus 354 Microsporangium 355 Male Gametophyte 355

Female Strobilus 356 Megasporogenesis 356 Female Gametophyte 356

Pollination 359 Post-pollination Development of Male Gametophyte 360

Fertilization 360 Embryogeny 360 Seed 364

Germination 364 Chromosome Number 365

20. Gnetales

Gnetaceae 366 Gnetum 366

Contents xix

343-365

366-404

xx Contents

Morphology 366 Anatomy 366

Root 366 Stem 367 Leaf 375

Reproduction 378 Male Strobilus 378

Microsporangium 378 Microsporogenesis 381 Male Gametophyte 383

Female Strobilus 384 Ovule 384 Megasporogenesis 389 Female Gametophyte 389

Pollination 391 Fertilization 393 Embryogeny 396 Seed 400

Germination 402 Chromosome Number 402 Temporal Consideration 403

21. Phylogenetic Considerations: Ephedra, Welwitschia and Gnetum

22. In Vitro Experimental Studies

Vegetative Tissues and Organs 408 Microspore and Male Gametophyte 411 Female Gametophyte 413 Embryo 423 Conifer Biotechnolgy 433 Somatic Embryogenesis 433 Genetic Transformation 436 Protoplast Culture 436 Micropropagation 437

23. Economic Importance

General Aspects 440 Woods 441

AJaucariaceae 442 Cupressaceae 442 Pinaceae 443 Podocarpaceae 444 Taxodiaceae 445 Taxaceae 445 Ginkgoaceae 445

405-407

408-439

440-456

Paper and Board 445 Resins 446 Hard Resins 44 7

Copal 447 Kauri Copal 447 Manila Copal 447 Amber 448 Sandarc 448

Oleoresins 449 Turpentine 449 Canada Balsam 450 Venice Turpentine 450

Tannins 451 Essential Oils 451 Fatty Oils 451 Food Supplements 452 Pharmaceuticals 454

24. Concluding Remarks

Seasonal Changes in Secondary Growth 457

Population and Wood Requirement 457

Pollution 458 Symbiosis 458 Reproductive Biology Pollen Biology 459 Life History 459

References Plant Index

458

Contents xxi

457-459

460-489 491-494

The Gymnosperms