Comparative Embryology of Angiosperms Vol.1

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2 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

3 Prof. Dr. Brij M. lohri Dr. Kunda B. Ambegaokar Department of Botany University of Delhi Delhi India Dr. Prem S. Srivastava Department of Botany Hamdard University Hamdard Nagar New Delhi India ISBN DOI / ISBN (ebook) This work is subject to copyright. A ll rights are reserved, whether the whole or part of th e 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 or 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, registered 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 ge neral usc.

4 Dedicated to Professor Karl Schnarf ( ) Professor Panchanan Maheshwari ( )

5 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 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.

6 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

7 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)

8 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, Springer Verlag, 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 ) 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.

9 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

10 Contents Volume 1 1 The Background Diagnostic Characters Developmental Aspects Anther Microsporogenesis Male Gametophyte Ovule Megasporogenesis Female Gametophyte Mature Embryo Sac Pollination Double Fertilization Endosperm Embryo Polyembryony Gametophytic Apomixis Seed Nutrition of the Anther, Ovule and Seed Comparative Studies I Dicotyledons: Archichlamydeae 4 Casuarinales General Features Casuarinaceae Taxonomic Considerations Juglandales General Features Myricaceae Juglandaceae Taxonomic Considerations Balanopales Balanopaceae Leitneriales

11 XIV ' Contents Volume 1 General Features Leitneriaceae.... Didymelaceae.... Taxonomic Considerations Salicales.... Salicaceae.... Taxonomic Considerations.... Fagales.... Betulaceae Fagaceae.... Taxonomic Considerations Urticales.... Rhoipteleaceae Ulmaceae.... Eucommiaceae Moraceae.... Urticaceae.... Taxonomic Considerations Proteales.... Proteaceae Taxonomic Considerations Santalales.... Olacaceae.... Dipentodontaceae..... Opiliaceae Grubbiaceae.... Santalaceae Myzodendraceae.... Loranthaceae: Loranthoideae, Viscoideae.... Taxonomic Considerations Balanophorales Balanophoraceae Taxonomic Considerations Medusandrales Polygonales Polygonaceae ' Taxonomic Considerations Centrospermae

12 Contents Volume 1 XV 16.2 Phytolaccaceae Gyrostemonaceae Achatocarpaceae Nyctaginaceae Molluginaceae Aizoaceae Portulacaceae Basellaceae Caryophyllaceae Dysphaniaceae Chenopodiaceae Amaranthaceae Didiereaceae Taxonomic Considerations Cactales General Features Cactaceae Taxonomic Considerations Magnoliales General Features Magnoliaceae Degeneriaceae Himantandraceae Winteraceae Annonaceae Eupomatiaceae Myristicaceae Canellaceae Schisandraceae Illiciaceae Austrobaileyaceae Trimeniaceae Amborellaceae Monimiaceae Calycanthaceae Gomortegaceae Lauraceae Hernandiaceae Tetracentraceae Trochodendraceae Eupteleaceae Cercidiphyllaceae Taxonomic Considerations Ranuncula1es General Features Ranunculaceae

13 XVI Contents Volume Berberidaceae Sargentodoxaceae Lardizabalaceae Menispermaceae Nymphaeaceae Ceratophyllaceae Taxonomic Considerations Piperales General Features Saururaceae Piperaceae Chloranthaceae Lactoridaceae Taxonomic Considerations Aristolochiales General Features Aristolochiaceae Rafflesiaceae Hydnoraceae Taxonomic Considerations Guttiferales General Features Dilleniaceae Paeoniaceae Crossosomataceae Eucryphiaceae Medusagynaceae Actinidiaceae Ochnaceae Dioncophyllaceae Strasburgeriaceae Dipterocarpaceae Theaceae Caryocaraceae Marcgraviaceae Quiinaceae Guttiferae (Clusiaceae) Ancistroc1adaceae Taxonomic Considerations Sarraceniales General Features Sarraceniaceae Nepenthaceae Droseraceae Taxonomic Considerations Papaverales

14 Contents Volume 1 XVII 24.1 General Features Papaveraceae Capparaceae Cruciferae Tovariaceae Resedaceae Moringaceae Taxonomic Considerations Batales Rosales General Features Platanaceae Hamamelidaceae Myrothamnaceae Crassulaceae Cephalotaceae Saxifragaceae Brunelliaceae Cunoniaceae Davidsoniaceae Pittosporaceae Byblidaceae Roridulaceae Bruniaceae Rosaceae Neuradaceae Chrysobalanaceae Connaraceae Leguminosae (Fabaceae) Krameriaceae Taxonomic Considerations Hydrostachyales General Features Hydrostachyaceae Taxonomic Considerations Podostemales General Features Podostemaceae Taxonomic Considerations Geraniales General Features Limnanthaceae Oxalidaceae Geraniaceae Tropaeolaceae Zygophyllaceae

15 XVIII Contents Volume Linaceae Erythroxylaceae Euphorbiaceae Daphniphyllaceae Taxonomic Considerations Rutales General Features Rutaceae Cneoraceae Simaroubaceae Picrodendraceae Burseraceae Meliaceae Akaniaceae Malpighiaceae Trigoniaceae Vochysiaceae Tremandraceae Polygalaceae Taxonomic Considerations Sapindales General Features Coriariaceae Anacardiaceae Aceraceae Bretschneideraceae Sapindaceae Hippocastanaceae Sabiaceae Melianthaceae Aextoxicaceae Balsaminaceae Taxonomic Considerations lulianiales Celastrales General Features Cyrillaceae Pentaphylacaceae Aquifoliaceae Corynocarpaceae Pandaceae Celastraceae Staphyleaceae Hippocrateaceae Stackhousiaceae Salvadoraceae

16 Contents Volume 1 XIX Buxaceae Icacinaceae Cardiopteridaceae Taxonomic Considerations Rhamnales General Features Rhamnaceae Vitaceae Leeaceae Taxonomic Considerations Malvales General Features Elaeocarpaceae Sarcolaenaceae Tiliaceae Malvaceae Bombacaceae Sterculiaceae Scytopetalaceae Taxonomic Considerations Thymelaeales General Features Geissolomataceae Penaeaceae Dichapetalaceae Thymelaeaceae Elaeagnaceae Taxonomic Considerations Violales General Features Flacourtiaceae Peridiscaceae Violaceae Stachyuraceae Scyphostegiaceae Turneraceae Malesherbiaceae Passifloraceae Achariaceae Cistaceae Bixaceae Sphaerosepalaceae Cochlospermaceae Tamaricaceae Frankeniaceae Elatinaceae

17 xx Contents Volume Caricaceae Loasaceae Datiscaceae Begoniaceae Taxonomic Considerations Cucurbitales General Features Cucurbitaceae Taxonomic Considerations Myrtiflorae General Features Lythraceae Trapaceae Crypteroniaceae Myrtaceae Dialypetalanthaceae Sonneratiaceae Punicaceae Lecythidaceae Melastomaceae Rhizophoraceae Combretaceae Onagraceae Oliniaceae Haloragaceae Theligonaceae (Cynocrambaceae) Hippuridaceae Cynomoriaceae Taxonomic Considerations Umbelliflorae General Features Alangiaceae Nyssaceae Davidiaceae Comaceae Garryaceae Araliaceae Umbelliferae Taxonomic Considerations References see Vol. 2 p Plant Index see Vol. 2 p. 1174

18 Contents Volume 2 II Dicotyledons: Sympetalae Diapensiales.... Diapensiaceae Taxonomic Considerations Ericales.... Clethraceae Pyrolaceae Ericaceae.... Empetraceae Epacridaceae Taxonomic Considerations Primulales Theophrastaceae.... Myrsinaceae.... Primulaceae Taxonomic Considerations Plumbaginales Plumbaginaceae Taxonomic Considerations.... Ebenales.... General Features Sapotaceae Sarcospermataceae Ebenaceae.... Styracaceae Lissocarpaceae Symplocaceae.... Hoplestigmataceae Taxonomic Considerations.... Oleales

19 XXII Contents Volume General Features Oleaceae Taxonomic Considerations Gentianales General Features Loganiaceae Desfontainiaceae Gentianaceae Menyanthaceae Apocynaceae Asclepiadaceae Rubiaceae Taxonomic Considerations Tubiflorae General Features Polemoniaceae Fouquieriaceae Convolvulaceae Hydrophyllaceae Boraginaceae Lennoaceae Verbenaceae Callitrichaceae Labiatae (Lamiaceae) Nolanaceae Solanaceae Duckeodendraceae Buddleiaceae Scrophulariaceae Globulariaceae Bignoniaceae Henriqueziaceae Acanthaceae Pedaliaceae Martyniaceae Gesneriaceae Columelliaceae Orobanchaceae Lentibulariaceae Myoporaceae Phrymaceae Taxonomic Considerations Plantaginales General Features Plantaginaceae Taxonomic Considerations

20 Contents Volume 2 XXIII 50 Dipsacales General Features Caprifoliaceae Adoxaceae Valerianaceae Dipsacaceae Taxonomic Considerations Campanulatae General Features Campanulaceae Sphenocleaceae Pentaphragmataceae Goodeniaceae Brunoniaceae Stylidiaceae Calyceraceae Compositae Taxonomic Considerations III Monocotyledons 52 Helobiae General Features Alismataceae Butomaceae Hydrocharitaceae Scheuchzeriaceae Aponogetonaceae Juncaginaceae Potamogetonaceae Zannichelliaceae Najadaceae Taxonomic Considerations Triuridales General Features Triuridaceae Taxonomic Considerations Liliiflorae General Features Liliaceae Xanthorrhoeaceae Stemonaceae Agavaceae Haemodoraceae Cyanastraceae

21 XXIV Contents Volume Amaryllidaceae Hypoxidaceae Velloziaceae Taccaceae Dioscoreaceae Pontederiaceae Iridaceae Geosiridaceae Burmanniaceae Corsiaceae Philydraceae Taxonomic Considerations Juncales General Features Juncaceae Thurniaceae Taxonomic Considerations Bromeliales General Features Bromeliaceae Taxonomic Considerations Commelinales General Features Commelinaceae Mayacaceae Xyridaceae Rapateaceae Eriocaulaceae Restionaceae Centrolepidaceae Flagellariaceae Taxonomic Considerations Graminales General Features Gramineae Taxonomic Considerations Principes General Features Palmae Taxonomic Considerations Synanthae General Features Cyclanthaceae Taxonomic Considerations Spathiflorae General Features

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