B.M. Johri . K.B. Ambegaokar . P.S. Srivastava

Comparative Embryology of Angiosperms Vol.1

With 362 Figures in two Volumes

Springer-Verlag Berlin Heidelberg GmbH

Prof. Dr. Brij M. lohri Dr. Kunda B. Ambegaokar Department of Botany University of Delhi Delhi 110007 India

Dr. Prem S. Srivastava Department of Botany Hamdard University Hamdard Nagar New Delhi 110062 India

This work is subject to copyright. A ll rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting. reproduction on microfilm or in any other way, and storage in data banks. Dupli­cation of this publication o r parts thereof is penniued only under the provisions of the German Copyright Law of September9, 1965, in itscuTTent version, and permission for use must always be obtained from Springer-Yerlag. Yiolations are liable for prosecution under the German Copyright Low.

Q Springer-Yerlag Berlin Heidelberg 1992 Originally publish~-d by Springer-Verlag Berlin Hcidelbt-rg in 1992 Soficover reprint orthe hardcover 1st edition 1992

The use of general descriptive names, registe red names, trademarks, etc. in this publication does oot imply, evcn in the absence of a spe<;ific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general usc.

ISBN 978-3-642-76397-7 ISBN 978-3-642-76395-3 (eBook) /978-3-642-76395-3 DOI 10.1007

Professor Karl Schnarf (1879-1947)

Dedicated to

Professor Panchanan Maheshwari (1904-1966)

Preface

The concept for the present book dates back to the early 1960s when Professor P. Maheshwari FRS (deceased May 1966) invited scientists to contribute to a multiauthored book - Comparative Embryology of An­giosperms. Soon thereafter, Professor Gwenda L. Davis (University of New England, Armidale, Australia) joined the University of Delhi as a Visiting Professor and announced that her book, Systematic Embryo­logy of Angiosperms, would soon be published (it appeared in 1966). Therefore, Professor Maheshwari postponed his project for the time being. The only book on embryology available until then (in english) was by Prof. Maheshwari, published in 1950 (McGraw-Hill, New York) al­though G. L. Davis' book (without illustrations) was also useful, but to a limited extent. The rapid increase in knowledge in the years that fol­lowed, with the advent of new techniques such as ultramicroscopy (TEM and SEM), histochemistry, and the study of organic and inorganic ma­terials, provided impetus to the study of embryology, transgressing its traditional confinements to 'compound' microscopy. This resulted in the emergence of new ideas and methodologies, which helped in a better interpretation of data previously restricted to only the description of developmental aspects. As new information on embryology accumulated, the need for a com­prehensive, illustrated book on comparative embryology that could serve researchers as a reference work became imperative. In 1979, when Dr. Klaus Peters of Springer-Verlag visited BMJ with an invitation to prepare a treatise on Embryology of Angiosperms, the book, Experimental Embryology of Vascular Plants (ed. B. M. Johri), was already in its final stage and was published by Springer-Verlag in 1982. The multi authored book Embryology of Angiosperms (ed. B.M. Johri) was published by Springer-Verlag in 1984, and the outline of the book, Comparative Embryology of Angiosperms, was prepared simul­taneously and given to Klaus Peters. Although a very large collection of relevant literature was available to us at the Department of Botany, University of Delhi, it took almost 10 years to survey the exhaustive literature and to complete the manuscript on Comparative Embryolo­gy.

VIII Preface

This book endeavours to consolidate the scattered information on angiosperm embryology, to present a coordinated and coherent inter­pretation of significant data accumulated thus far, and to elucidate the interrelationship of taxa based on comparative embryological find­ings. Comparative Embryology of Angiosperms is a review of the develop­mental processes leading to sexual reproduction in flowering plants, and provides illustrated source material on sexual reproduction, primitive and advanced embryological characters, and phylogenetic relationships. On the basis of embryological data and certain evidence from other areas of study, special emphasis has been placed on the relationship among and within the families and orders of angiosperms. For our work Engler's classification system for angiosperms proposed in Syllabus der Pflanzen­familien, revised by Melchior (1964), has been adopted. This treatise summarizes nearly 5000 publications (up to 1991) on an­giosperm embryology; its resulting ultimate size prompted us to divide it into two volumes. For the convenience of the reader, the preface, acknowledgements and the table of contents are presented in both volumes. For economic reasons, the references for the entire work and the plant index are included only in Volume 2. Each order begins with general features, followed by the embryology of its constituent families, culminating in critical taxonomic considerations on the basis of comparative embryologic data, and the placement of each family in six chosen systems of classification is discussed. A comprehensive bibliography is given at the end of each family and includes the citations referred to in the text. Additional citations (not mentioned in the text) are provided for those who wish to obtain further information. When appropriate, inaccuracies in observation and interpretation are pointed out, alternative interpretations offered, gaps in our knowledge highlighted, and prospects outlined. Comparative Embryology of Angiosperms provides useful information to students and research workers involved in angiosperm embryology, taxonomists, plant breeders, agriculturists and horticulturists, and plant scientists interested in the evolutionary tendencies in flowering plants.

University of Delhi, July 1992 B.M. Johri K. B. Ambegaokar

P. S. Srivastava

Acknowledgements

The preparation of Comparative Embryology of Angiosperms took al­most a decade. We could not have accomplished this undertaking without the academic support and encouragement from our colleagues at the University of Delhi, and plant scientists in India and abroad. Over the last 55 years, B. M. Johri has had the privilege of discussions with embryologists on the reproductive biology of angiosperms, espe­cially at national and international conferences. He is grateful to all of them. We are indebted to fellow embryologists who generously provided rel­evant literature and advance information about ongoing research proj­ects. To name a few: Prof. Martin Bopp (Heidelberg), Dr. F. Bouman and Dr. W. Verkerke (Amsterdam), the late Dr. R. Dahlgren (Copen­hagen), Dr. M.l. Hakki (Berlin), Prof. U. Hamann (Bochum), Dr. R. Wunderlich (Vienna), Prof. M. Yakovlev and Prof. T. Batygina (Leningrad), Dr. H. Tobe (Chiba, Japan), Dr. P.H. Raven (St. Louis, USA), Prof. A. E. Cocucci (Cordoba, Argentina), the late Dr. Thani­koimani (Pondichery, India), Prof. K. K. Lakshmanan (Coimbatore, India), Dr. M.P. Nayar and Dr. B.D. Sharma (Botanical Survey of India, Calcutta), Prof. S. Chanda (Bose Institute, Calcutta) and our colleagues (at the University of Delhi) Prof. H. Y. Mohan Ram, Prof. S.c. Maheshwari, Prof. R.N. Kapil, Prof. N.N. Bhandari and Dr. Ashok Bhatnagar. For advance information regarding the recently published (1989) Dedi­catory Volume in honour of Prof. A. Lebegue, Some Aspects and Orientations in Plant Embryology, we wish to thank the editors, Prof. J. Pare and Prof. M. Bugnicourt (University of Picardie, Amiens, France). For the translation of Russian literature we are obliged to Dr. Balram Sharma (Indian Agric. Res. Inst., New Delhi) and Dr. (Mrs.) Akhilesh Tiwari (University of Delhi); and to Dr. (Mrs.) Sheela Srivastava (Uni­versity of Delhi) for German literature. Dr. M. A. Rau (Mysore) very kindly prepared a number of schematic figures on the development of endosperm in the Sympetalae. Dr. F. Bouman provided scanning elec­tron micrographs of seeds which appear in this volume. Thanks are also extended to Mr. Dev Raj Khosla (Indian Natl. Sci. Acad., New Delhi)

x Acknowledgements

who prepared photocopies of literature and illustrations, so essential for checking the manuscript. Mr. S. B. Ambegaokar (Delhi) was kind enough to compare over 1800 pages of the final manuscript with the earlier draft. Dr. (Mrs.) Kapoor ably assisted us in the collection of literature and indexing the Contents and References. Dr. Kapoor and Dr. Manoj Kumar (University of Delhi) also helped in the preparation of plates. We express our appreciation to Mr. G. A. Shastri and Ms. Sushil Kaur for easy access to the University Library, and to Dr. (Mrs.) Vats ala (Publ. Direc., CSIR, New Delhi), Dr. (Mrs.) Nina Dhiman and Ms. Binny Mathur (Editorial Assistants, Phytomorphology), and our re­search scholars Mr. N. Syed Abbas Ali and Ms. Babeeta Chrungu for their help in checking the periodicals. Mr. Krishan Lal, artist (University of Delhi), very competently designed and prepared several figures, and labelled all the illustrations. Mr. D. Daniel and Mr. S. K. Das (Univer­sity of Delhi) attended to photographic work with professional compe­tence. We are indebted to Dr. Dieter Czeschlik, Life Science Editor, Springer­Verlag, Heidelberg, for his immense patience in waiting to see the com­pletion of this work. Moreover, we would like to thank Mrs. Jean von dem Bussche for copyediting "Sympetalae" (pp. 615-829) in 1987; Dr. G. Forbath, Copy Editor, Ms. Carola Wissmeier, Secretary to Dr. D. Czeschlik, and Ms. Renate Miinzenmayer, Production Editor, Springer-Verlag, for the meticulous handling of this publication. For the high quality of printing by MacMillan India, Bangalore, we have great appreciation. We are obliged to Janab Hakim Abdul Hameed Sahab (Chancellor), Prof. M. Amin (Vice Chancellor) and Prof. Muhammed Iqbal (Dean, Faculty of Science) of the Hamdard University, New Delhi, for their interest and encouragement in this project. Thanks are also due to Ms. Madhumati, Mr. Aniruddha, Ms. Deep­shikha, and Mr. Deepak, research scholars at the Department of Botany, Hamdard University for cross-checking the text and references in the final proofs, and for preparing plant indices. We would like to make special mention of Mr. R. K. Gupta (University of Delhi) for expert secretarial assistance, and typing twice the manu­script of about 2000 pages, and Mrs. Krishna Thakur (University of Delhi) for periodical secretarial assistance. Mr. Sukhbir Singh and Mr. Chandra Pal (University of Delhi) have been very helpful in attending to the day-to-day routine work during the preparation of this book. The Indian National Science Academy, New Delhi, and Springer-Ver­lag, Heidelberg, provided partial financial support for this book. We also wish to record gratitude to Mrs. Raj Johri (wife of B. M. Johri) and Dr. (Mrs.) Sheela Srivastava (wife of P. S. Srivastava) for their support, forbearance and interest during the long periods when Com­parative Embryology of Angiosperms was in preparation. Dr. Lalit M.

Acknowledgements XI

Johri (son) and Mrs. Meera Johri (daughter-in-law) keenly followed the progress of this book and encouraged an expeditious completion. Once again, many thanks to all those who helped us in various ways.

May 28,1992 B.M. Johri K. B. Ambegaokar

P. S. Srivastava

Contents Volume 1

1 The Background ............................ 1 2 Diagnostic Characters . . . . . . . . . . . . . . . . . . . . . . . . . 2 3 Developmental Aspects ....................... 4 3.1 Anther................................... 4 3.2 Microsporogenesis........................... 11 3.3 Male Gametophyte .......................... 15 3.4 Ovule.................................... 19 3.5 Megasporogenesis........................... 29 3.6 Female Gametophyte . . . . . . . . . . . . . . . . . . . . . . . . . 31 3.7 Mature Embryo Sac. . . . . . . . . . . . . . . . . . . . . . . . . . 37 3.8 Pollination................................. 42 3.9 Double Fertilization . . . . . . . . . . . . . . . . . . . . . . . . . . 48 3.10 Endosperm ................................ 53 3.11 Embryo................................... 62 3.12 Polyembryony .............................. 84 3.13 Gametophytic Apomixis. . . . . . . . . . . . . . . . . . . . . . . 88 3.14 Seed ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 3.15 Nutrition of the Anther, Ovule and Seed. . . . . . . . . . . 105 3.16 Comparative Studies ......................... 111

I Dicotyledons: Archichlamydeae

4 Casuarinales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 4.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 4.2 Casuarinaceae.............................. 113 4.3 Taxonomic Considerations ..................... 115 5 Juglandales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 5.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 5.2 Myricaceae................................ 117 5.3 Juglandaceae............................... 117 5.4 Taxonomic Considerations ..................... 119 6 Balanopales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 6.1 Balanopaceae.............................. 120 7 Leitneriales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

XIV

7.1 7.2 7.3 7.4 8 8.1 8.2 8.3 9 9.1 9.2 9.3 9.4 10 10.1 10.2 10.3 10.4 10.5 10.6 10.7 11 11.1 11.2 11.3 12 12.1 12.2 12.3 12.4 12.5 12.6 12.7 12.8 12.9 13 13.1 13.2 13.3 14 15

'15.1 15.2 15.3 16 16.1

Contents Volume 1

General Features Leitneriaceae .............................. . Didymelaceae ............................. . Taxonomic Considerations .............. ...... . Salicales ................................. . General Features ........................... . Salicaceae ................................ . Taxonomic Considerations .................... . Fagales .................................. . General Features ........................... . Betulaceae . . . . . . . . . . . . . . . . . . . . . . . . . ....... . Fagaceae ................................. . Taxonomic Considerations .................. .. . Urticales ................................. . General Features ........................... . Rhoipteleaceae ....................... ..... . Ulmaceae ................................ . Eucommiaceae . . . . . . . . . . . . . . . . . . . . . . . ...... . Moraceae ................................ . Urticaceae ................................ . Taxonomic Considerations ............ ........ . Proteales ................................. . General Features ........................... . Proteaceae . . . . . . . . . . . . . . . . . . . . ............ . Taxonomic Considerations .............. ...... . Santalales ................................ . General Features ........................... . Olacaceae ................................ . Dipentodontaceae ......................... .. Opiliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . Grubbiaceae .............................. . Santalaceae ............. . . . . . . . . . . . . . . . . . . . Myzodendraceae ........................... . Loranthaceae: Loranthoideae, Viscoideae ......... . Taxonomic Considerations ................ .... . Balanophorales .................... ........ . General Features ........................... . Balanophoraceae ....... . . . . . . . . . . . . . . . . . . . . . Taxonomic Considerations ..... ............... . Medusandrales . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . Polygonales . . . . . .......................... . General Features ........................... . Polygonaceae . . . . . . '. . . . . . . . . . . . . . . . . ....... . Taxonomic Considerations ................. ... . Centrospermae .............. .............. . General Features ........................... .

120 120 121 121 122 122 122 123 124 124 124 127 128 129 129 129 130 131 132 146 147 148 149 149 155 155 155 156 158 158 161 162 171 175 192 200 200 201 206 207 207 207 207 210 210 211

Contents Volume 1 XV

16.2 Phytolaccaceae.............................. 211 16.3 Gyrostemonaceae............................ 212 16.4 Achatocarpaceae ............................ 212 16.5 Nyctaginaceae .............................. 212 16.6 Molluginaceae ...... . . . . . . . . . . . . . . . . . . . . . . . . 214 16.7 Aizoaceae .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216 16.8 Portulacaceae .............................. 217 16.9 Basellaceae ................................ 218 16.10 Caryophyllaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219 16.11 Dysphaniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 16.12 Chenopodiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 16.13 Amaranthaceae ............................. 224 16.14 Didiereaceae .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225 16.15 Taxonomic Considerations ..................... 227 17 Cactales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232 17.1 General Features. . . . . . . . . . . . . . . . . . . . . . . . . . . . 232 17.2 Cactaceae ................................. 232 17.3 Taxonomic Considerations ..................... 235 18 Magnoliales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236 18.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236 18.2 Magnoliaceae............................... 237 18.3 Degeneriaceae.............................. 238 18.4 Himantandraceae............................ 240 18.5 Winteraceae ............................... 242 18.6 Annonaceae ............................... 243 18.7 Eupomatiaceae ............................. 246 18.8 Myristicaceae............................... 248 18.9 Canellaceae................................ 252 18.10 Schisandraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252 18.11 Illiciaceae ................................. 254 18.12 Austrobaileyaceae ........................... 255 18.13 Trimeniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255 18.14 Amborellaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260 18.15 Monimiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260 18.16 Calycanthaceae ............................. 263 18.17 Gomortegaceae ............................. 265 18.18 Lauraceae .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 18.19 Hernandiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 18.20 Tetracentraceae ............................. 269 18.21 Trochodendraceae ........................... 269 18.22 Eupteleaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270 18.23 Cercidiphyllaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270 18.24 Taxonomic Considerations ..................... 272 19 Ranuncula1es. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 19.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 19.2 Ranunculaceae.............................. 285

XVI Contents Volume 1

19.3 Berberidaceae .............................. 291 19.4 Sargentodoxaceae ........................... 292 19.5 Lardizabalaceae............................. 292 19.6 Menispermaceae ............................ 294 19.7 Nymphaeaceae.............................. 295 19.8 Ceratophyllaceae ....... . . . . . . . . . . . . . . . . . . . . . 301 19.9 Taxonomic Considerations ..................... 301 20 Piperales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 20.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 20.2 Saururaceae................................ 308 20.3 Piperaceae................................. 309 20.4 Chloranthaceae ............................. 313 20.5 Lactoridaceae .............................. 314 20.6 Taxonomic Considerations ..................... 315 21 Aristolochiales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316 21.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316 21.2 Aristolochiaceae ............................ 316 21.3 Rafflesiaceae............................... 318 21.4 Hydnoraceae............................... 320 21.5 Taxonomic Considerations ..................... 321 22 Guttiferales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 22.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 22.2 Dilleniaceae................................ 324 22.3 Paeoniaceae................................ 325 22.4 Crossosomataceae ........................... 328 22.5 Eucryphiaceae.............................. 329 22.6 Medusagynaceae ............................ 329 22.7 Actinidiaceae............................... 329 22.8 Ochnaceae................................. 330 22.9 Dioncophyllaceae............................ 333 22.10 Strasburgeriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 22.11 Dipterocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 22.12 Theaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336 22.13 Caryocaraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337 22.14 Marcgraviaceae ............................. 337 22.15 Quiinaceae ................................ 339 22.16 Guttiferae (Clusiaceae) . . . . . . . . . . . . . . . . . . . . . . . . 339 22.17 Ancistroc1adaceae ........................... 341 22.18 Taxonomic Considerations ..................... 341 23 Sarraceniales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348 23.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349 23.2 Sarraceniaceae.............................. 349 23.3 Nepenthaceae .............................. 350 23.4 Droseraceae ............................... 351 23.5 Taxonomic Considerations ..................... 353 24 Papaverales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355

Contents Volume 1 XVII

24.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355 24.2 Papaveraceae............................... 355 24.3 Capparaceae ............................... 357 24.4 Cruciferae................................. 360 24.5 Tovariaceae................................ 362 24.6 Resedaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 362 24.7 Moringaceae ............................... 363 24.8 Taxonomic Considerations ..................... 364 25 Batales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366 26 Rosales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366 26.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366 26.2 Platanaceae................................ 367 26.3 Hamamelidaceae ............................ 367 26.4 Myrothamnaceae .... . . . . . . . . . . . . . . . . . . . . . . . . 368 26.5 Crassulaceae ............................... 369 26.6 Cephalotaceae.............................. 372 26.7 Saxifragaceae............................... 372 26.8 Brunelliaceae............................... 374 26.9 Cunoniaceae ............................... 374 26.10 Davidsoniaceae ............................. 375 26.11 Pittosporaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 26.12 Byblidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 26.13 Roridulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 26.14 Bruniaceae ................................ 379 26.15 Rosaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 26.16 Neuradaceae ............................... 382 26.17 Chrysobalanaceae ........................... 382 26.18 Connaraceae ............................... 383 26.19 Leguminosae (Fabaceae) ...................... 383 26.20 Krameriaceae .............................. 392 26.21 Taxonomic Considerations ..................... 393 27 Hydrostachyales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407 27.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407 27.2 Hydrostachyaceae ........................... 407 27.3 Taxonomic Considerations ..................... 408 28 Podostemales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408 28.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408 28.2 Podostemaceae ............................. 409 28.3 Taxonomic Considerations ..................... 415 29 Geraniales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416 29.1 General Features. . . . . . . . . . . . . . . . . . . . . . . . . . . . 416 29.2 Limnanthaceae ............................. 416 29.3 Oxalidaceae................................ 418 29.4 Geraniaceae ............................... 419 29.5 Tropaeolaceae .............................. 420 29.6 Zygophyllaceae ............................. 423

XVIII Contents Volume 1

29.7 Linaceae .......... . . . . . . . . . . . . . . . . . . . . . . .. 424 29.8 Erythroxylaceae............................. 426 29.9 Euphorbiaceae.............................. 427 29.10 Daphniphyllaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430 29.11 Taxonomic Considerations ..................... 431 30 Rutales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438 30.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438 30.2 Rutaceae.................................. 439 30.3 Cneoraceae................................ 441 30.4 Simaroubaceae.............................. 441 30.5 Picrodendraceae............................. 443 30.6 Burseraceae................................ 444 30.7 Meliaceae ................................. 444 30.8 Akaniaceae ................................ 447 30.9 Malpighiaceae .............................. 447 30.10 Trigoniaceae ............................... 450 30.11 Vochysiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451 30.12 Tremandraceae ............................. 455 30.13 Polygalaceae ............................... 455 30.14 Taxonomic Considerations ..................... 459 31 Sapindales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 462 31.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 462 31.2 Coriariaceae ............................... 462 31.3 Anacardiaceae.............................. 463 31.4 Aceraceae................................. 466 31.5 Bretschneideraceae .......................... 467 31.6 Sapindaceae................................ 467 31.7 Hippocastanaceae ........................... 468 31.8 Sabiaceae ................................. 470 31.9 Melianthaceae .............................. 470 31.10 Aextoxicaceae ............. . . . . . . . . . . . . . . . . . 471 31.11 Balsaminaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 472 31.12 Taxonomic Considerations ..................... 475 32 lulianiales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479 33 Celastrales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479 33.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480 33.2 Cyrillaceae ................................ 480 33.3 Pentaphylacaceae............................ 482 33.4 Aquifoliaceae .............................. 482 33.5 Corynocarpaceae ........... . . . . . . . . . . . . . . . . . 483 33.6 Pandaceae................................. 483 33.7 Celastraceae ............................... 483 33.8 Staphyleaceae .............................. 485 33.9 Hippocrateaceae ............................ 486 33.10 Stackhousiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486 33.11 Salvadoraceae .............................. 487

Contents Volume 1 XIX

33.12 Buxaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 488 33.13 Icacinaceae ................................ 489 33.14 Cardiopteridaceae ........................... 491 33.15 Taxonomic Considerations ..................... 491 34 Rhamnales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495 34.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495 34.2 Rhamnaceae ............................... 495 34.3 Vitaceae .................................. 497 34.4 Leeaceae.................................. 498 34.5 Taxonomic Considerations ..................... 499 35 Malvales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 501 35.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501 35.2 Elaeocarpaceae ............................. 502 35.3 Sarcolaenaceae ............................. 502 35.4 Tiliaceae ......................... . . . . . . . . . 503 35.5 Malvaceae................................. 504 35.6 Bombacaceae............................... 509 35.7 Sterculiaceae ...................... . . . . . . . . . 510 35.8 Scytopetalaceae ............................. 512 35.9 Taxonomic Considerations ..................... 512 36 Thymelaeales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515 36.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515 36.2 Geissolomataceae............................ 515 36.3 Penaeaceae ................................ 515 36.4 Dichapetalaceae............................. 516 36.5 Thymelaeaceae ............................. 517 36.6 Elaeagnaceae............................... 518 36.7 Taxonomic Considerations ..................... 519 37 Violales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 521 37.1 General Features. . . . . . . . . . . . . . . . . . . . . . . . . . . . 521 37.2 Flacourtiaceae ............. . . . . . . . . . . . . . . . . . 522 37.3 Peridiscaceae............................... 524 37.4 Violaceae ................................. 524 37.5 Stachyuraceae .............................. 525 37.6 Scyphostegiaceae .. . . . . . . . . . . . . . . . . . . . . . . . . . . 526 37.7 Turneraceae................................ 526 37.8 Malesherbiaceae ............................ 528 37.9 Passifloraceae .............................. 529 37.10 Achariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 530 37.11 Cistaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 530 37.12 Bixaceae .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 531 37.13 Sphaerosepalaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . 532 37.14 Cochlospermaceae .......... . . . . . . . . . . . . . . . . . 532 37.15 Tamaricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 533 37.16 Frankeniaceae ....... . . . . . . . . . . . . . . . . . . . . . . . 538 37.17 Elatinaceae ................................ 540

xx Contents Volume 1

37.18 Caricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 541 37.19 Loasaceae ............................... . . 542 37.20 Datiscaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 544 37.21 Begoniaceae ............................... 546 37.22 Taxonomic Considerations ..................... 549 38 Cucurbitales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 556 38.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 556 38.2 Cucurbitaceae .............................. 556 38.3 Taxonomic Considerations ..................... 563 39 Myrtiflorae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 563 39.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 564 39.2 Lythraceae ................................ 564 39.3 Trapaceae ................................. 565 39.4 Crypteroniaceae............................. 567 39.5 Myrtaceae................................. 569 39.6 Dialypetalanthaceae.......................... 571 39.7 Sonneratiaceae.............................. 571 39.8 Punicaceae ................................ 572 39.9 Lecythidaceae .............................. 572 39.10 Melastomaceae ............................. 573 39.11 Rhizophoraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 574 39.12 Combretaceae .............................. 578 39.13 Onagraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 580 39.14 Oliniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 584 39.15 Haloragaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 585 39.16 Theligonaceae (Cynocrambaceae) ................ 586 39.17 Hippuridaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 587 39.18 Cynomoriaceae ............................. 588 39.19 Taxonomic Considerations ..................... 588 40 Umbelliflorae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 601 40.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 601 40.2 Alangiaceae................................ 602 40.3 Nyssaceae ................................. 602 40.4 Davidiaceae................................ 603 40.5 Comaceae................................. 603 40.6 Garryaceae ................................ 604 40.7 Araliaceae................................. 606 40.8 Umbelliferae .................... . . . . . . . . . . . 607 40.9 Taxonomic Considerations ..................... 610

References see Vol. 2 p. 1013 Plant Index see Vol. 2 p. 1174

Contents Volume 2

II Dicotyledons: Sympetalae

41 41.1 41.2 41.3 42 42.1 42.2 42.3 42.4 42.5 42.6 42.7 43 43.1 43.2 43.3 43.4 43.5 44 44.1 44.2 44.3 45 45.1 45.2 45.3 45.4 45.5 45.6 45.7 45.8 45.9 46

Diapensiales .............................. . General Features ........................... . Diapensiaceae ....................... ...... . Taxonomic Considerations .............. ...... . Ericales .................................. . General Features ........................... . Clethraceae . . . . . . . . . . . . . . . . . . . . . . . . . ...... . Pyrolaceae . . . . . ........................... . Ericaceae ................................ . Empetraceae ........ . . . . . . . . . . . . .......... . Epacridaceae . . . . . . . . . . . . . . . . . . . . . . . ....... . Taxonomic Considerations .............. ...... . Primulales . . . . . . . . . . . . . . . . . . . . ............ . General Features ........................... . Theophrastaceae ........................... . Myrsinaceae .............................. . Primulaceae . . . . . . . . . . . . . . . . . . . . . . . ........ . Taxonomic Considerations .............. ...... . Plumbaginales .............. ............... . General Features ........................... . Plumbaginaceae . . . . . . . . . . . . . . . . . . . . ........ . Taxonomic Considerations .................... . Ebenales ................................. . General Features . . . . . . . . . . . . . . . . . . . . . . . .... . Sapotaceae ....................... ........ . Sarcospermataceae . . . . . . . . . . . . . . . . . . . . . . . ... . Ebenaceae ................................ . Styracaceae .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lissocarpaceae . . . . . . . . . . . . . . ............... . Symplocaceae ............................. . Hoplestigmataceae . . . . . . . . . . . . . . . . . . . . . . . . . . . Taxonomic Considerations .................... . Oleales .................................. .

615 615 615 617 617 617 617 618 620 623 624 626 629 629 630 630 633 634 635 636 636 640 641 641 641 642 643 644 645 645 647 647 650

XXII Contents Volume 2

46.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650 46.2 Oleaceae.................................. 650 46.3 Taxonomic Considerations ..................... 651 47 Gentianales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 652 47.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 652 47.2 Loganiaceae ............................... 653 47.3 Desfontainiaceae ............................ 656 47.4 Gentianaceae............................... 656 47.5 Menyanthaceae ............................. 657 47.6 Apocynaceae............................... 658 47.7 Asclepiadaceae ............................. 660 47.8 Rubiaceae................................. 663 47.9 Taxonomic Considerations ..................... 670 48 Tubiflorae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 675 48.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 675 48.2 Polemoniaceae.............................. 675 48.3 Fouquieriaceae ............................. 676 48.4 Convolvulaceae ............................. 676 48.5 Hydrophyllaceae ............................ 685 48.6 Boraginaceae............................... 689 48.7 Lennoaceae................................ 696 48.8 Verbenaceae ............................... 696 48.9 Callitrichaceae.............................. 702 48.10 Labiatae (Lamiaceae) . . . . . . . . . . . . . . . . . . . . . . . . . 707 48.11 Nolanaceae ................................ 711 48.12 Solanaceae ................................ 712 48.13 Duckeodendraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . 715 48.14 Buddleiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715 48.15 Scrophulariaceae ............................ 716 48.16 Globulariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 732 48.17 Bignoniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 733 48.18 Henriqueziaceae ............................ 737 48.19 Acanthaceae ............................... 737 48.20 Pedaliaceae .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . 746 48.21 Martyniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 747 48.22 Gesneriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 748 48.23 Columelliaceae ............................. 752 48.24 Orobanchaceae ............................. 752 48.25 Lentibulariaceae ............................ 755 48.26 Myoporaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 762 48.27 Phrymaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 762 48.28 Taxonomic Considerations ..................... 763 49 Plantaginales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777 49.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777 49.2 Plantaginaceae.............................. 777 49.3 Taxonomic Considerations ..................... 779

Contents Volume 2 XXIII

50 Dipsacales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 779 50.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 780 50.2 Caprifoliaceae .............................. 780 50.3 Adoxaceae ................................ 781 50.4 Valerianaceae .............................. 782 50.5 Dipsacaceae................................ 782 50.6 Taxonomic Considerations ..................... 789 51 Campanulatae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 796 51.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 797 51.2 Campanulaceae ............................. 797 51.3 Sphenocleaceae ............................. 802 51.4 Pentaphragmataceae.......................... 803 51.5 Goodeniaceae .............................. 803 51.6 Brunoniaceae............................... 805 51. 7 Stylidiaceae................................ 806 51.8 Calyceraceae .... . . . . . . . . . . . . . . . . . . . . . . . . . . . 808 51.9 Compositae................................ 808 51.10 Taxonomic Considerations ..................... 821

III Monocotyledons

52 Helobiae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 831 52.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 831 52.2 Alismataceae............................... 831 52.3 Butomaceae................................ 835 52.4 Hydrocharitaceae............................ 838 52.5 Scheuchzeriaceae............................ 843 52.6 Aponogetonaceae ........................... 844 52.7 Juncaginaceae .............................. 846 52.8 Potamogetonaceae........................... 847 52.9 Zannichelliaceae ............................ 850 52.10 Najadaceae ................................ 853 52.11 Taxonomic Considerations ..................... 856 53 Triuridales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 860 53.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 860 53.2 Triuridaceae................................ 860 53.3 Taxonomic Considerations ..................... 861 54 Liliiflorae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 862 54.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 863 54.2 Liliaceae ........................ . . . . . . . . .. 863 54.3 Xanthorrhoeaceae ........................... 871 54.4 Stemonaceae ...................... . . . . . . . . . 872 54.5 Agavaceae................................. 874 54.6 Haemodoraceae............................. 875 54.7 Cyanastraceae .............................. 877

XXIV Contents Volume 2

54.8 Amaryllidaceae ............................. 877 54.9 Hypoxidaceae .............................. 879 54.10 Velloziaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 882 54.11 Taccaceae ................................. 882 54.12 Dioscoreaceae ........................ . . . . . . 883 54.13 Pontederiaceae ............................. 884 54.14 Iridaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 885 54.15 Geosiridaceae .............................. 886 54.16 Burmanniaceae ............................. 887 54.17 Corsiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 889 54.18 Philydraceae ............................... 889 54.19 Taxonomic Considerations ..................... 890 55 Juncales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 898 55.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 898 55.2 Juncaceae ................................. 898 55.3 Thurniaceae................................ 900 55.4 Taxonomic Considerations ..................... 900 56 Bromeliales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 901 56.1 General Features. . . . . . . . . . . . . . . . . . . . . . . . . . . . 901 56.2 Bromeliaceae............................... 901 56.3 Taxonomic Considerations ..................... 902 57 Commelinales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903 57.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903 57.2 Commelinaceae ....... . . . . . . . . . . . . . . . . . . . . . . 904 57.3 Mayacaceae................................ 909 57.4 Xyridaceae ................................ 911 57.5 Rapateaceae ............................... 914 57.6 Eriocaulaceae .............................. 916 57.7 Restionaceae............................... 920 57.8 Centrolepidaceae............................ 921 57.9 Flagellariaceae.............................. 922 57.10 Taxonomic Considerations ..................... 923 58 Graminales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 928 58.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 928 58.2 Gramineae ................................ 928 58.3 Taxonomic Considerations ..................... 943 59 Principes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 944 59.1 General Features .. . . . . . . . . . . . . . . . . . . . . . . . . . . 944 59.2 Palmae ................................... 944 59.3 Taxonomic Considerations ..................... 950 60 Synanthae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 950 60.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 951 60.2 Cyclanthaceae .............................. 951 60.3 Taxonomic Considerations ..................... 952 61 Spathiflorae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 953 61.1 General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . 953

Contents Volume 2

61.2 61.3 61.4 62 62.1 62.2 62.3 62.4 62.5 63 63.1 63.2 63.3 64 64.1 64.2 64.3 64.4 64.5 64.6 64.7 65 65.1 65.2 65.3 66 67 68

Araceae ................................. . Lemnaceae ............................... . Taxonomic Considerations .................... . Pandanales ......................... ...... . General Features ........................... . Pandanaceae ............................. .. Sparganiaceae ............................. . Typhaceae ................................ . Taxonomic Considerations .................... . Cyperales ................................ . General Features ........................... . Cyperaceae ............................... . Taxonomic Considerations .................... . Scitamineae ............................... . General Features ........................... . Musaceae ................................ . Zingiberaceae ..................... ........ . Cannaceae ................................ . Marantaceae ....................... ....... . Lowiaceae ................................ . Taxonomic Considerations ................. ... . Microspermae .......................... ... . General Features ........................... . Orchidaceae ....................... ....... . Taxonomic Considerations .................. .. . Concluding Remarks .............. .......... . References ............................... . Plant Index .... . . . . . . . . . . . . . . . . . . . . . ...... .

XXV

953 957 964 968 968 969 971 972 973 974 974 974 981 982 982 982 983 989 990 991 991 993 993 993

1004 1005 1013 1174