The Genera of Araceae

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ARACEAE S J M ayo

J Bogner

P C Boyce

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ARACEAE S J M ayo

J Bogner

WITH CONTRIBUTIONS FROM

J.C. French and R. Hegnauer ILLUSTRATIONS BY

E. Catherine

P C Boyce

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© Copyright The Trustees, Royal Botanic Gardens, Kew First published 1997 ISBN 1 900347 22 9

Cover design by John Stone Book design by Jeff Eden Page make-up by Media Resources, Information Services Department, Royal Botanic Gardens, Kew

Printed in The European Union by Continental Printing, Belgium.

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C O N T E N T S - TO R E T U R N TO T H I S PA G E P R E S S C

Dedication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .vii Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viii Preface

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Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi

A

GENERAL PART

1. History

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2. Vegetative Morphology

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3. Vegetative Anatomy (by J.C. French)

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4. Inflorescence and Floral Morphology

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5. Inflorescence and Floral Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 6. Fruits and Seeds

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7. Seedling Morphology 8. Embryology 9. Cytology

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10. Palynology

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11. Phytochemistry and Chemotaxonomy (by R. Hegnauer) 12. Ecology and Life Forms 13. Pollination Biology 14. Dispersal

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15. Geography 16. Uses

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17. Cultivation

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18. Conservation

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19. Fossil Record

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20. Phylogenetic relationships within the Monocotyledons

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21. Phylogenetic relationships within Araceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 22. Previous classifications

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CONTENTS

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CONTENTS

B

TAXONOMIC PART

23. Synopsis of the Classification of Araceae

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24. Family Description of Araceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 25. Key to the Genera of Araceae and Acoraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 26. Descriptions of the Tribes and Genera of Araceae

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27. Description of Acoraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 28. References and selected taxonomic literature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291 29. Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. Appendix: Table 9. Fungal Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 10. Schott’s (1860) classification of Aroideae . . . . . . . . Table 11. Engler’s (1876b) classification of Araceae . . . . . . . Table 12. Engler’s (1920b) classification of Araceae . . . . . . . Table 13. Grayum’s (1990) classification of Araceae . . . . . . . Table 14. Bogner & Nicolson’s (1991) classification of Araceae Table 15. Generic country lists . . . . . . . . . . . . . . . . . . . . . . . Table 16. Colour plates: photo credits and vouchers . . . . . . . 31. Index to scientific names . . . . . . . . . . . . . . . . . . . . . . . . . . . 32. Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. Descriptions of new taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . Colour Plates

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D E D I C AT I O N

We dedicate this book to the memory of Heinrich Wilhelm Schott and Heinrich Adolf Gustav Engler, the two great founding fathers of Araceae systematics, and also to Nicholas Edward Brown, whose studies of the family, while less widely known, were of the highest standard.

DEDICATION

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F O R E WO R D

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By Dan H. Nicolson

It has been almost 150 years since H.W. Schott published his monumental Genera Aroidearum (The Genera of Araceae). A new treatment of the family has been a desideratum for a century and urgently needed during the last fifty years. What has been particularly needed is an illustrated work that can be used by anyone to recognize unknowns and help learn the terms necessary for accurate understanding of the taxa. It is often taken for granted that the family is more or less completely known and information is readily available. The first problem is that the keystone works by Schott (1794-1865) and Engler (1844-1930) are expensive, if available for purchase, and not to be found in public libraries. If you are fortunate enough to find them, you discover the second problem. They are in Latin! In spite of such problems there has been a revival of interest in the family. An astonishing number of workers have been studying plants in the field and cultivation and applying new and sophisticated techniques. New tools, such as the scanning electron microscope (which revolutionized comparative study of pollen) and techniques, such as molecular systematics and cladistics, have revolutionized thinking about relationships. At last it has been possible to stimulate three people, scarcely more than entering middle life and with all the necessary modern training, to collaborate and synthesize what is known about the genera of Araceae, including at least one crisp, fresh and new full page drawing of each genus. Comparison of this work with its predecessor shows the distance we have travelled. Do not think that the last word has been spoken. New species, even genera, are still turning up and some of them are testing the hypotheses that we wish were facts. Smithsonian Institution, Washington, D.C. March 1994

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P R E FA C E

The Araceae, or aroids, are plants which are very familiar to everyone but paradoxically little known. Monstera deliciosa, Epipremnum pinnatum ‘Aureum’ (syn. Epipremnum aureum), Philodendron scandens, Dieffenbachia maculata and Aglaonema commutatum may be counted as among the world’s most popular house plants. But most rarely flower in their domestic environment, and the fact that they are all aroids is appreciated by relatively few people. Once their inflorescences appear, however, it is obvious they belong together, with their characteristic cowl-like spathe and central, fleshy spike, known as the spadix. The most familiar aroid inflorescences are those of Anthurium andraeanum and Zantedeschia aethiopica, which are used the world over as cut flowers. The family is extraordinarily diverse in appearance, with the foliage being probably its most widely appreciated feature. The perforated leaf of Monstera deliciosa (the subject of our cover) is one of the most instantly recognizable plant images the world over, while the velvet, pendent leaves of Anthurium waroqueanum are unforgettably elegant. The beauty of many other genera, such as Philodendron, Alocasia and Arisaema is also very largely due to their superb foliage. In sharp contrast, the inflorescences of certain aroids are quite repulsive; that of Helicodiceros muscivorus, for example, resembles nothing so much as the rear end of a decomposing mammal corpse, while so bizarre is the foul-smelling Amorphophallus konjac that some years ago it was chosen as the basis for a film dramatization of the “triffids” of science fiction. Towering over all such lesser monstrosities is Amorphophallus titanum, one of the vegetable wonders of the world, which attracts large crowds at botanic gardens when from time to time the great tuber yields a huge phallic inflorescence smelling of rotten fish. Why is it then, that aroids are not better known as a plant group? We think the reason is a dearth of accessible literature. Apart from Deni Bown’s excellent “Aroids. Plants of the Arum Family”, published in 1988 for the non-specialist reader, there have been no general and comprehensive treatments of the Araceae, either technical or lay, since the publication of Engler’s taxonomic monograph (in Latin and German) for his “Das Pflanzenreich” series in the first two decades of this century. We have tried to contribute towards filling this gap with a general taxonomic treatment of the family which extends to the level of genus. While it is unashamedly technical, and may seem somewhat forbidding to the general reader, we hope that the book’s publication will encourage more people to study and enjoy these wonderful plants, which are among the

most beautiful and dramatic that the vegetable kingdom has to offer. The idea for this book germinated in 1980 when the first international workshop on the systematics of Araceae was held at the Marie Selby Botanical Garden, Sarasota, Florida, organized by Dr Michael Madison. Madison and Simon Mayo began a manuscript but the project never came to fruition. In 1987, with the encouragement of Professor Grenville Lucas, then Keeper of the Kew Herbarium, Mayo and Josef Bogner of the Munich Botanic Garden resolved to tackle the task anew and Peter Boyce joined us soon afterwards. Eleanor Catherine, the artist, completed the team at a later stage. The format of this book is modelled on that of the Genera Palmarum by Dr Natalie Uhl and Dr John Dransfield. The delimitation of all the genera has been critically re-examined in the light of modern studies. Since A. Engler’s last monograph in “Das Pflanzenreich” various new genera have been described and old ones reduced to synonymy. F. Gagnepain described Pycnospatha from Thailand, H. Jumelle Arophyton and Carlephyton from Madagascar and S. Buchet Colletogyne, also from Madagascar. M. Hotta contributed four new genera, Heteroaridarum, Pedicellarum and Phymatarum from Sarawak, and Furtadoa from Sumatra and the Malay Peninsula. D.H. Nicolson and colleagues contributed Bognera and Filarum from tropical America and Hottarum from Borneo. G.S. Bunting described the extraordinary aquatic Jasarum, and Lasimorpha, a synonym of Cyrtosperma according to Engler, has been reestablished by A. Hay. Hay also recently described Anaphyllopsis from tropical America and Lazarum from Australia. Despite these changes, the total number of genera treated here (105 without Acorus) is much the same as that presented by Engler (108 without Acorus). Cladistic analysis of morphological and molecular data in recent years has, however, meant that the classification has changed considerably since Engler’s time. We have deliberately laid the primary emphasis on the preparation of completely revised descriptions of the genera, together with analytical illustrations for each. The plates are all original drawings by Eleanor Catherine, based on a combination of herbarium, spirit (from the Kew spirit collection) and living specimens, supplemented when necessary by photographs. The chapters of the General Part are intended to be summaries, in some cases quite brief, of various aspects of Araceae which are of taxonomic and general interest. In two cases, the treatments are much more detailed, namely chapter 3 on vegetative anatomy, and chapter 11 on phytochemistry and chemotaxonomy, which we

P R E FA C E

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were very fortunate to receive from Professor J.C. French and Professor R. Hegnauer, respectively. Their contributions are the first detailed modern reviews in English of these subjects for the Araceae . We have omitted any general treatment of the molecular systematics of the family, which is being studied, in particular, by Professor French and his colleagues at Rutgers University, New Jersey. They have generously allowed us access to their most important phylogenetic conclusions, thus greatly improving our discussion of the family’s phylogeny (chapter 21). We have included material on Acorus, including a generic treatment, despite the fact that we accept that this genus does not belong to the Araceae. However, it was felt that it would be convenient to the non-specialist reader, who might expect to find something on

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the genus in a general treatment of aroids. In contrast, we have not included any systematic account of the Lemnaceae, which according to the molecular work of French and colleagues, seem clearly to be embedded within the Araceae (see chapter 20). Our reasons are again pragmatic. The taxonomy of the Lemnaceae has been comprehensively revised in recent years by Professor E. Landolt and French’s results became available only in the final stages of preparation of this book. Throughout the text we have employed the terms “aroid” and “araceous” as synonymous adjectives referring to any member of the family, and the noun “aroid” likewise. The reader should therefore not interpret “aroid” as referring only to members of the subfamily Aroideae.

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A C K N OW L E D G E M E N T S

A book of this kind cannot be prepared without the help of many people besides the authors. Our greatest debt is to Eleanor Catherine, whose professionalism, patience and superb facility in line illustration has produced such a unique and wonderful set of plates. It is likely that her work will be the most valuable contribution that this book can make to advancing knowledge of the Araceae and it is no exaggeration to say that all aroid-lovers are in her debt. The growth of activity in aroid systematics in recent decades has been notable for the friendly spirit of collaboration that has always prevailed and because of this we have been able to call on the help of many other specialist colleagues. To all these we are extremely grateful. Dr Dan H. Nicolson undertook the task of reviewing our manuscript and contributing a foreword. The generic descriptions have been revised by the following specialists who have been very generous in allowing us to use unpublished manuscripts and in providing new data and insights on taxa and character fields concerning which their knowledge is much more profound than our own:- Mr Julius Boos, Dr Thomas B. Croat, Professor James C. French, Dr Michael H. Grayum, Dr A. Hay, Professor R. Hegnauer, Mr Wilbert Hetterscheid, Professor Niels Jacobsen, Dr Z. Kvaček, Dr Gitte Petersen, Professor Robin Scribailo, Dr Mikhail Serebryanyi, Dr Elke Seubert, Dr M. Sivadasan, Professor Thomas Stützel, Dr Sue Thompson, Professor Hans-Jürgen Tillich and Dr Guanghua Zhu. Professor Tom Ray contributed new data on shoot organization. These contributions have greatly enriched this book, but its shortcomings are necessarily our responsibility alone. We would also like to thank the following staff members of the Royal Botanic Gardens, Kew, who, among many others, provided essential support:Professor Ghillean T. Prance, Dr Charles Stirton, Professor Grenville Lucas, Dr Michael Lock, Dr John Dransfield, Dr Phillip Cribb, Miss Mary Gregory, Mr

Milan Svanderlik, Mr Jeff Eden, Mrs Christine Beard, Dr Geoff Kite, Mrs Margaret Newman, Mr John Stone, Mr John Woodhams, Mr Michael Marsh, Mr John Hale, Mr John Norris, Mr Phillip Brewster. It is also a pleasure to thank Cássia Mônica Sakuragui for preparing the original maps. We are very grateful to the directors and staff of the following herbaria for lending us material:- A, AAU, B, BK, BKF, BM, BO, BRUN, C, DNA, E, FI, G, HRB, K, KEP, L, LE, M, MO, NY, P, SAN, SAR, SEL, SGN, SI, U, US, YUKU, YUNU. J. Bogner wishes to thank the Directors of the following institutes for financial support: the Smithsonian Institution (Washington DC, USA) for a field study in Sarawak, the Centre National de la Recherche Scientifique (CNRS, Paris, France) for the publication of the Araceae for “Flore de Madagascar et des Comores” and the Royal Botanic Gardens, Kew (UK) for visits to RBG Kew in connection with the preparation of this book. The living plants used in the preparation of this book were cultivated at the Botanischer Garten Munich and the Living Collections Department of the Royal Botanic Gardens, Kew and we are very grateful to the Curators and staff of these institutes for their helpful collaboration. Other colleagues should be mentioned who have prepared the ground in some important way; Deni Bown for her splendid pioneering book “Aroids: Plants of the Arum family”, Dr George Bunting, who was responsible for starting the modern renaissance in aroid systematics, Dr Michael Madison, Mrs Betty Waterbury and Mrs Libby Besse who with their friends and colleagues, started the International Aroid Society and its excellent journal Aroideana, and finally Professor P.B. Tomlinson who provided the impetus for continuing the international workshops on Araceae by organizing the second (Harvard Forest 1984) and leading the third (Berlin 1987) meetings.

ACKNOWLEDGEMENTS

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GENERAL

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H I S TO RY

The word “arum” is derived directly some of the finest botanical artists of from ancient Greek “aron” and indithe day (Riedl & Riedl-Dorn 1988). vidual species of the Araceae have The pencil drawings are of herbeen recorded by many botanists barium specimens and material and historians since those ancient preserved in alcohol from times. Theophrastus (ca. herbaria all over Europe and 371–285 B.C.) recorded Arum the water colour plates were in his treatise (Prime 1960), painted from living plants and Hernandez (1790) grown at Schönbrunn, the described a number of tropiimperial gardens near Vienna. cal aroids and their uses by The colour plates represent Aztec people. European each plant in astonishing Araceae were described in detail and thoroughness and detail by botanists such as are among the most impresFuchs (1542) and Ray (1686) sive analyses ever achieved in while R. Dodoens (1574) the medium of botanical illusarranged all Araceae known to tration. Only relatively few him into a single group. were published (Schott J. P. de Tournefort (1700) 1853–1857, 1857, 1858, Peyritsch created a “class” without a name 1879). Today the Icones are part which grouped three European of the collections of the Vienna genera (Arum, Dracunculus and Natural History Museum. About 80 Arisarum), characterized by the posplates (including subfamily Lasioideae) session of a “monopetalous” flower. are lost, while Schott’s herbarium of This concept of the aroid infloresAraceae, consisting of 1379 specicence as a flower with a single petal mens, was destroyed at the end of Figure 1. H.W. Schott (1794–1865). also influenced Linnaeus (1753, the Second World War (Schott 1984, Photograph c. 1860. 1754) who classified the known Riedl & Riedl-Dorn 1988). species according to his artificial Schott’s most important works sexual system. It was not until later that the infloreswere the account of Araceae in the Meletemata botancence was recognized as a spike (spadix) of tiny ica (Schott 1832), the Genera Aroidearum (Schott 1858) flowers surrounded by an often colourful bract and the Prodromus systematis Aroidearum (Schott (spathe). A.L. de Jussieu (1789) established the Araceae 1860). Some of his new genera were first published in as a natural family but recognized only a few small or the cultural periodical Wiener Zeitschrift für Kunst, rather broadly conceived genera, probably because of Literatur, Theater und Mode as part of a series of articles the paucity of good material of non-European taxa. entitled Für Liebhaber der Botanik (Schott 1829a–e, All the climbing species were grouped under the name 1830). He also published many papers on Araceae in the Pothos and most of the terrestrial species were placed Oesterreiches Botanisches Wochenblatt, which later in the genera Arum and Dracontium. became the Oesterreichische Botanische Zeitschrift (from Modern systematic studies of the Araceae began 1858, the 8th volume, onwards). with the work of the Austrian botanist and gardener Schott described many new genera and species Heinrich Wilhelm Schott. He was the first monographer and created the first major natural classification of the of the family and the first botanist to make careful whole family. Though his taxon concepts were narcomparative studies of aroid inflorescences, flowers row, many of his genera and species have withstood and fruits. Using these observations he was able to the test of time. He created the basis of Araceae taxput the family on a sound taxonomic footing. His work onomy, not only for Engler, who soon followed him is documented in his writings (Riedl 1965a, b) and by in studying the family comprehensively, but also for an outstanding archive of scientific illustrations (Schott succeeding generations. A notable aspect of Schott’s 1984). This consists of a collection of 4400 superb work was that he used a combination of herbarium coloured and black and white plates of Araceae, the material, living plants and field work in the study of a Icones Aroidearum, which were prepared at his direclargely tropical plant group at a time when such a tion and personal expense and employed the talents of wide-ranging approach was most unusual.

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Riedl (1965a, b, pers. comm.) Other significant works on the has given details of Schott’s life family published during Schott’s and work. He was born on 7th lifetime include Kunth’s treatment January 1794 in Brünn in for his Enumeratio Plantarum Moravia (now Czech Republic). (Kunth 1841), which was the first At the age of seven he moved to post-Linnean treatment at species Vienna, where his father had level, and Blume’s Rumphia become head gardener at the (Blume 1836–1837), which was botanical garden of the important especially for Asian University. There he soon came genera and included very fine into contact with eminent coloured plates. botanists. N. von Jacquin stimuThe second great monographer lated and directed the early of Araceae was Adolf Engler. interest of the boy to study Right from the start he established plants and once, when severely himself as an authority on the ill as a youth, he received a visit family by a series of prodigious from Alexander von Humboldt works. His first major publication which made a lifelong impreson the family outlined a new syssion upon him. tem on phylogenetic lines (Engler The young Schott attended 1876b) which was substantially lectures in botany and other different from Schott’s classificadisciplines (agriculture, chemtion, especially at the higher istry) at the University, but his ranks. The following year he prefirst employment was as an sented a pioneering comparative assistant gardener under the study of araceous shoot organidirection of his father. In 1815 zation, based on original he became gardener for the colobservations (Engler 1877). His Figure 2. H.A. Engler (1844–1930). lection of Austrian flora at the treatment of Araceae for Martius’s Photograph taken when Engler was Belvedere Palace. Then, at von Flora Brasiliensis followed nineteen years old. Jacquin’s recommendation he (Engler 1878), and immediately was chosen to become a parafterwards a comprehensive ticipant in the famous Austrian-Bavarian monograph at species level for the entire scientific expedition to Brazil which family appeared in De Candolle’s included the botanist Mikan, the Monographiae Phanerogamarum zoologist Natterer, the mineralogist (Engler 1879). His account for and botanist Pohl, the botanist Beccari’s Malesia (Engler 1883a) C.F.P. von Martius and zoologist was followed by the family treatJ. von Spix. ment for Die natürlichen Between 1817 and 1821 he Pflanzenfamilien (Engler 1887– worked in Brazil as a member 1889). At about this time he of this group of naturalists and also published two papers in made acquaintance with a the Botanische Jahrbücher rich tropical flora (Schreiber which further developed his 1822). His main duty was to phylogenetic interpretations establish a garden in Rio de of morphological and anatomJaneiro to prepare plants for ical trends in the Araceae the long journey to Europe. (Engler 1883b, 1884). When he returned to Vienna With the initiation of the in 1821 he took up gardening Das Pflanzenreich series in again, eventually rising to 1900, Engler embarked on his become the Director of the second monograph of the entire botanical and zoological gardens family, completed in 1920 (Engler at the imperial palace of Schön1905, 1911, 1912, 1915, 1920a, brunn, a position he held until his 1920b, Engler & Krause 1908, 1920, death on 5th March 1865. Schott was Krause 1908, 1913) with the help of his made a Doctor “honoris causa” and assistant Kurt Krause (1883–1963). Engler a member of the Kaiserliche and Krause doubled Schott’s total Akademie der Wissenschaften for of about 900 species to around Figure 3. H.A. Engler in his later years his work in botany and horticulture. 1800. An important feature of this

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treatment was the large number of institute. He was by now an extremely illustrations, prepared by J. Pohl. able organizer and under his direcThese are mostly original but tion a new botanical garden and some were copied from Schott’s botanical museum were planned illustrations. The large living and built at Dahlem, then on collection of Araceae at the the outskirts of Berlin. Started Botanischer Garten Berlinin 1897, the new garden was Dahlem was an important inaugurated in 1910. In 1902 resource for Engler’s studies. he journeyed to the Cape and He carried out little field Transvaal in South Africa, East work but he was an accomAfrica and Egypt. In 1905 he plished synthesiser of data visited South and East Africa from the notes and collecagain and also Zimbabwe, tions of others. Botanists from Zambia and the Zambezi all over the world sent him region, after which he went material and information. to India and Sri Lanka, where Diels’s (1931) biography he paid particular attention to gives a detailed account of economic plants and Araceae. Engler’s life and work, from His journey then took him to which we have selected the Bogor, Singapore, Malaysia, main facts. Engler was born on Burma, the Himalaya and 25th March 1844 in Sagan, Lower Calcutta. In 1913 he went on a Silesia (today Poland), the son of world tour that took him to a tradesman. In 1848 his mother Southwest Africa, China, Japan, took him to the provincial capital of Hawaii, California and New Breslau, where the young Engler England. He was awarded honFigure 4. N.E. Brown (1849–1934). grew up. From his early years he orary doctorates of the Universities Photograph c. 1900. took a great interest in natural hisof Cambridge, Cape Town, tory and during his University years Uppsala and Geneva and the gold studied with the famous palaeobmedal of the Linnean Society. He otanist and teacher H.R. Goeppert, later becoming a retired officially as Director at Berlin on 31st March teacher himself for a short time in Breslau (today 1921 but continued his scientific work until his death Wroclaw). In 1869 he became acquainted with A.W. on 10th October 1930. Eichler who had succeeded C. von Martius as editor of K. Krause continued to work on Araceae at Berlin the “Flora Brasiliensis” and this contact was to be of until 1942, publishing new species collected by vargreat significance for his future career. In April 1871 he ious field botanists and collectors, especially those was employed as a scientific assistant at the Botanische working in South America, but he did not take up his Sammlungen (herbarium and living collections) in studies again after the Second World War ended in Munich. Here, under the guidance of Nägeli’s direc1945 and died in obscurity in 1963. A manuscript torship, he matured scientifically for eight years until he that he completed in 1942 for the second edition of was 34 years old. The botanic garden and herbarium, Die natürlichen Pflanzenfamilien was lost in 1943 together with the library (Bayerische Staatsbibliothek) when the Berlin Botanical Museum was largely provided excellent working conditions. He worked up destroyed by war action. his professorial qualification (habilitation) in 1872 and Nicholas Edward Brown (1849–1934), working at later taught at the University. Engler’s period at Munich the Herbarium of the Royal Botanic Gardens, Kew, was clearly the most creative of his life as far as the made significant contributions to Araceae systematics. Araceae were concerned, since it was there that he His largest and most important publication on made most of his important innovations and insights Araceae was the family treatment for Flora of Tropical into the family’s taxonomy. Africa (Brown 1901), but in addition to this he pubIn 1879 Engler became a professor ordinarius at lished various other flora treatments, many new the University of Kiel, where he also became Director species and various new genera. Many of his new of the botanical garden and institute. However, after the taxa were based on living material introduced into the death of Goeppert in 1884 he was appointed to sucgardens at Kew. Brown’s work is particularly notable ceed his former professor at the University of Breslau for his meticulous descriptions based on accurate which included the directorship of the botanic garden. observation and it is evident that his grasp of Araceae There he began work on Die natürlichen Pflansystematics was profound. Brown was an almost exact zenfamilien together with K. Prantl. After five years, he contemporary of Engler. He began work at the Kew moved to Berlin in October 1889 to become the Herbarium as an assistant in 1873 and retired in 1914 Director of Germany’s largest botanical garden and as an Assistant Keeper.

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Joseph Dalton Hooker (1883), in Bentham & Hooker’s Genera Plantarum, published a classification of Araceae based largely on Schott’s (1860) work which was later revised and modified by John Hutchinson (1934, 1959, 1973), another Kew botanist. Hutchinson made an impact on Araceae taxonomy because his treatment included a key to all genera in English. However, aroid specialists have found his system to be artificial and it was never widely used. T. Nakai (1943) published a new classification in which he recognized the Pistiaceae, Cryptocorynaceae and Acoraceae as separate families from the Araceae. He also made important contributions to the systematics of the genus Arisaema. After 1945 a period of relative inactivity followed, until the 1950s when George S. Bunting, Monroe Birdsey, Dan H. Nicolson and Josef Bogner began working on the family. M. Hotta began to publish on Araceae in the 1960s (Hotta 1965, 1966a–b, 1967) and later he presented a classification for Eastern Asian and Malesian genera and an influential study of vegetative and floral morphology (Hotta 1970, 1971). Since then he has made many important investigations of aroids in the Malay Archipelago region (Hotta 1976, 1981, 1982, 1984, 1985, 1986a–b). Other important studies carried out during this period are those by G. Thanikaimoni (pollen morphology, 1969) and C. Marchant (1970, 1971a–b, 1972, 1973, cytology). Floristic and revisionary studies in the 1950s and 1960s that should be mentioned are those of H.C.D. de Wit (Cryptocoryne, Lagenandra), H. Riedl (Flora Iranica, Eminium), Bunting (Spathiphyllum) and Nicolson (Aglaonema). In the 1970s and 1980s, the pace of work on the systematics of Araceae increased. In 1978 the International Aroid Society was formed in Florida, USA and the journal Aroideana founded, originally under the energetic editorship of M.T. Madison. Aroideana generated a significant expansion of scientific and horticultural interest in the family and continues to play an important role. A series of international workshops on Araceae systematics was established, beginning at Sarasota (1980), and continuing at Harvard Forest (1984), Berlin (1987), Moscow (1992), Tokyo (1993), and Kunming (1995), with further meetings planned for Sydney (1998) and St Louis (1999).

These events have done a great deal to foster international collaboration. Bogner (1979a) updated the Engler classification, adding newly described genera and taking account of new synonymy. Nicolson (1983) published an English translation of Engler’s classification, including the accepted genera described since 1920. Bogner & Nicolson (1991) published a revised synoptic key to all the genera, which incorporates a number of important changes from Engler’s concepts, particularly in the subfamilies Pothoideae and Lasioideae. M.H. Grayum (1984, 1990) presented a new phylogenetic classification which is especially notable for recommending the removal of Acorus from Araceae and the large scale reorganization of Engler’s subfamilies. Grayum’s work, in particular, has been a stimulus to other workers to initiate new studies of taxonomically significant character fields, largely due to his comprehensive review of the technical literature. Recent comparative character surveys of particular significance are the studies of J.C. French (anatomy, also with P.B. Tomlinson), T. Ray and P. Blanc (shoot morphology), G. Petersen (cytology), M. Grayum (palynology), H.-J. Tillich (seedling morphology), D. Barabé, L. Chrétien, S. Forget and M. Labrecque (floral anatomy), W.N. Carvell (floral anatomy), E. Seubert (seed anatomy) and J.C. French, M. Chung & Y. Hur (cpDNA molecules). There are many centres of research on Araceae systematics now active throughout the world, including those in Brazil (I.M. Andrade, C. Barros, E. Gonçalves, A. Lins, M. Nadruz Coelho, M.L. Soares, F. Ramalho, C. Sakuragui, J. Waechter). Calicut (M. Sivadasan), Copenhagen (N. Jacobsen, G. Petersen), Kew (S.J. Mayo, P.C. Boyce, G. Kite), Kunming (Li Heng), Kyoto and Kagoshima (M. Hotta), Leiden (W. Hetterscheid), Moscow (M. Serebryanyi), Munich (J. Bogner), Rutgers University, New Jersey (J.C. French, M. Chung, Y. Hur), St. Louis (T.B. Croat, M.H. Grayum, G. Zhu), Sydney (A. Hay), Tokyo (J. Murata), Washington DC. (D.H. Nicolson), and until recently, at Yaoundé (C. Ntépé-Nyame†). With this increase in taxonomic activity, new revisions and floristic studies are now in progress which should eventually lead to a much more complete knowledge of the family.

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Root Roots in Araceae are always adventitious and dimorphic roots are often found in climbing hemiepiphytes, e.g. Monstera deliciosa, Philodendron bipinnatifidum. In some genera, e.g. Arisarum, Arum, Biarum, Cryptocoryne, specialized contractile roots occur which prevent the stem from rising too near to the soil surface; more details of root structure are given in Chapter 3.

Stem

Pinellia, and hypogeal stolons in Colocasia, Cryptocoryne, some Spathiphyllum spp. and Lasimorpha. Shoot organization shows a range of interesting variation within the family and can be taxonomically useful (Engler 1877, Blanc 1977a,b, 1978, 1980, Ray 1986, 1987a–c, 1988, 1990). In virtually all genera the mature stem is a sympodium composed of sympodial units (articles) each of which has a more-or-less determinate structure, beginning with a prophyll and ending with an inflorescence or aborted inflorescence. Foliage leaves and cataphylls (reduced sheath-like leaves) occur in a sometimes very regular sequence within each unit. Continued growth of the stem takes place in most genera by the development of a “continuation shoot” in the axil of the leaf (foliage leaf or cataphyll) situated at the second node below the spathe insertion. In the subfamily Orontioideae it arises at the first node below the spathe. The production of more than one inflorescence to form a floral sympodium commonly takes place by the development of short units consisting of a prophyll, a spathe and a spadix. The first unit arises in the axil of the

The stem varies from an aerial elongated axis with extended internodes, as in the many climbing hemiepiphytes, to a hypogeal rhizome or subglobose tuber. Climbing genera with long internodes are commonest in the more primitive tribes, i.e. those with bisexual flowers. Geophytes are found throughout the family but are especially common in the most advanced subfamily, the Aroideae. Abbreviated aerial stems, resulting in rosulate plant forms, are also commonly found, as in many epiphytic species of Philodendron and Anthurium. Some, generally larger, species have an Continuation shoot arborescent habit, in which the main Spathe axis is a fleshy (Alocasia, Xanthosoma) or fibrous (Philodendron) stem, or a Spadix pseudostem of petiole sheaths Prophyll (Arisaema, Typhonodorum). Shoot Peduncle types apparently specialized for vegeSympodial leaf tative reproduction occur in various forms. Flagelliform shoots, or “flagellae”, equivalent to aerial stolons, have been observed in Amydrium, Cercestis, Pedicellarum, Philodendron, Pothos, Rhaphidophora, Rhodospatha, and Syngonium among others. They consist of branches (usually in the form of continuation shoots) in which the internodes become very much longer and more slender than in the flowering zone of the stem, and the leaves often Monopodial leaf become reduced in size, sometimes to small, scale-like cataphylls. Flagelliform shoots grow rapidly, and thus encounter new host trees on which flowering stems later develop. Bulbils, which appear to be dispersed by birds, occur in Remusatia, while tubercles occur in Amorphophallus bulbifer, Dracontioides, Dracontium, and Figure 5. Schematic diagram of a common form of shoot architecture in Araceae.

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leaf immediately below the spathe and succeeding ones in the axils of the prophylls. These floral sympodia have a range of structural variation which may be quite complex; extreme forms are found in Homalomena (Ray 1988). Anomalous shoot organization of an apparently unique type occurs in Gymnostachys, while in most tribe Potheae and Heteropsis the flowering axes occur as lateral short shoots on monopodial main vegetative axes of apparently indeterminate growth.

Leaf In virtually all genera the leaf is clearly differentiated into an expanded blade, petiole and petiole sheath; exceptions are Gymnostachys and some Biarum species. The sheath normally clasps the subtended internode, at least basally, and has an annular insertion (except many Potheae and most Heteropsis). The foliage leaves that occur nearest the end of sympodial shoot articles (sympodial leaves) often have short or very reduced sheaths, particularly when the article apex aborts and fails to develop an inflorescence, e.g. in Philodendron. The terminology adopted here to describe leaf shape requires some explanation as it differs from traditional practice. For descriptive purposes, the leaf is divided into the anterior division, corresponding to that part of the blade surrounding the midrib, and two posterior divisions, which, when present, are those portions of the leaf blade which extend basally on each side of the petiole insertion. In many genera, e.g. those comprising the Monsteroideae, there are no posterior divisions and the blade is composed entirely of the anterior division. In other taxa, such as the Lasioideae, many species have deeply sagittate leaves with very strongly developed posterior divisions, sometimes exceeding the anterior division in length. In strongly sagittate leaves, each posterior division usually has a well developed basal rib, which performs the same mechanical support role as the midrib does for the anterior division. In cordate and cordate-sagittate leaves the basal ribs may be short or even absent, with the individual primary lateral veins arising independently at the base of the midrib. On the other hand, pedately divided leaves, as seen for example in Philodendron goeldii or Sauromatum venosum, have a central, undivided anterior division while the posterior divisions are represented by the lateral series of pedate segments on either side. Here the basal ribs are represented by the “arms” on which the segments of the posterior divisions are inserted and which arch back from the midrib insertion at the apex of the petiole. Leaf blade size and shape is exceedingly diverse. Size may range from diminutive (e.g. Ambrosina) to gigantic (e.g. Alocasia, Amorphophallus, Anchomanes, Cyrtosperma, Xanthosoma ). Shape varies from linear (Biarum, Jasarum) to dracontioid (tribe Thomsonieae, Anchomanes, Dracontium, Pseudohydrosme, Pycnospatha, ), through elliptic, ovate, cordate, sagittate, hastate, trifid or trisect, pedatifid, pinnatifid, pedatisect,

pinnatisect and radiatisect. Sometimes the posterior divisions of pedatifid and pedatisect leaves are twisted spirally so that the leaf segments resemble a spiral staircase (Eminium, Helicodiceros). Bipinnatifid, tripinnatifid and partially quadripinnatifid leaves also occur. Dracontioid leaves may be viewed as elaborated forms of sagittate, hastate, trisect or pedatisect leaves in which the anterior and posterior divisions of the leaf blade have become highly dissected during ontogeny by differential growth of the margin, necrosis of parts of the blade, or a combination of both processes. The mature leaf can be compared to an umbrella which has been blown inside out by a gale — the spokes correspond to a much-divided system of major leaf ribs and the canopy is rent into numerous “tattered” lamina lobes which adhere to the ribs and are more regularly shaped at the tips of the major leaf veins. Gonatopus has a somewhat similar leaf form while Zamioculcas and certain species of Anaphyllum and Anaphyllopsis have completely pinnatisect leaves. The first foliage leaf of a seedling is always entire (sagittate–hastate) in the genera Anaphyllopsis, Anaphyllum, Anchomanes, Dracontium, and Pycnospatha, but divided (dracontioid type) in the genera Gonatopus and Amorphophallus. Heteroblasty is a striking and sometimes taxonomically useful feature (e.g. Madison 1977a) of a number of climbing genera (Cercestis, Monstera, Philodendron, Pothos, Rhaphidophora, Rhodospatha, Syngonium). It occurs both in ontogeny from seedling to the mature plant and in association with the development of flagelliform shoots. A very striking form of heteroblasty is shown in certain genera (e.g. Monstera, Pothos) where the juvenile leaves have very short petioles and their blades are held flat against the host tree in a regular, overlapping sequence giving the appearance of roof shingles or tiles; these are consequently known as shingle plants. Monstera dubia is a well known example which also has beautifully variegated leaf blades in this growth phase. Perforated (fenestrate) leaves are another peculiarity of Araceae in genera such as Dracontioides, Monstera, Rhaphidophora, and in juvenile leaves of Anchomanes and certain species of Cercestis. A number of genera have species with peltate leaves (tribe Colocasieae, Anthurium, Caladium, Homalomena). The midrib is almost always present, being absent only in Gymnostachys and Pistia. The major veins which compose the midrib and basal ribs and branch laterally from them are termed primary lateral veins. Secondary, tertiary and higher orders of lateral veins are recognized by their relative thickness and/or their hierarchical level of branching. The primary lateral veins may be arcuate-parallel (e.g. Ambrosina), pedate (e.g. Sauromatum) or radiate (e.g. many Arisaema spp.) but most commonly are pinnately arranged. Even in pedatifid (-sect) and radiatisect leaves, the primary lateral veins of each segment are generally pinnate. Except in deeply divided leaves, the primary lateral veins always run throughout the leaf blade, ultimately joining together at the leaf apex (Ertl 1932).

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The primary lateral veins generally run to the margin first, where they form a marginal vein which then runs to the leaf apex. In some species, most primary lateral veins curve arcuately within the margin to fuse together at the apex, and in these cases only the lowermost primaries run into the margin to form a marginal vein. In other genera either one or several of the primary lateral veins form a submarginal collective vein (brochidodromous pattern) which lies parallel to the marginal veins. The finer venation may be reticulated (e.g. Anthurium) or may run essentially parallel to the pinnately arranged primary lateral veins (e.g. Philodendron). This is often referred to as “parallel” or “striate” venation in aroid literature. In this book we refer to the latter type as parallel-pinnate, to distin-

guish it from true, grass-type parallel venation, which in Araceae occurs only in Gymnostachys. A third type of fine venation (“colocasioid venation”) has been recognized by previous authors for tribes Colocasieae and Caladieae. In this pattern the finer veins branch almost at right angles from the primary lateral veins and then arch strongly towards the leaf margin, often fusing on the way to form a more-or-less sinuose interprimary collective vein and then finally joining into a submarginal collective vein. Intermediates occur between most recognized types. Venation patterns and their development require new investigation, particularly because of their great potential for facilitating identification of species and genera. Ertl’s study (1932) is the only large scale comparative survey yet made of this important character field.

ANTERIOR DIVISION

Midrib Primary lateral vein

Petiole insertion

Submarginal collective vein

POSTERIOR DIVISION

POSTERIOR DIVISION

ANTERIOR DIVISION

B Basal ribs Petiole ANTERIOR DIVISION

POSTERIOR DIVISION

A

Petiole

POSTERIOR DIVISION

Basal ribs Petiole

C

Figure 6. Leaf division types: A, no posterior division development; B, moderate posterior division development; C, extreme posterior division development.

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V E G E TAT I V E A N ATO M Y By James C. French

The vegetative anatomy of the Araceae is among the most diverse of any family of monocotyledons. Stem vasculature is the most diverse of any monocotyledon group and virtually every known type of secretory structure occurs, including resin canals, laticifers, extrafloral nectaries, mucilage cavities and intravaginal squamules.

Root The first root to emerge from the seed is a short-lived extension of the radicle. Subsequent roots arise from the stem and may form lateral roots. The stem is the normal site of root origin in Araceae, a pattern referred to by Troll (1949) as “secondary homorhizy” and typical of monocotyledons.

Anchor and feeder roots Aerial roots of epiphytic and climbing Araceae are often specialized into anchor roots (Haftwurzeln) that serve to attach the plant to the substrate which provides support, and feeder roots (Nährwurzeln) which extend to the soil and supply water and dissolved nutrients (Went 1895). A number of anatomical, morphological and physiological differences between anchor and feeder roots have been described (Schimper 1888, Lierau 1888, Wettstein 1904, Gaulhofer 1907, Linsbauer 1907, Porsch 1911, Solereder & Meyer 1928, Funke 1929, Goebel & Sandt 1930, Birdsey 1955, Guttenberg 1968, Madison 1977a). Both anchor and feeder roots typically arise close to the node, although the former may arise along the internode as well in some species of e.g. Anthurium, Epipremnum, Monstera, Rhaphidophora and Scindapsus (Madison 1977a, Croat & Baker 1978, Hotta 1971). According to Troll (1941), the association between anchor and feeder roots can be highly specific, as in the monophyllous sympodium of Philodendron. In P. scandens a feeder root develops near the insertion of the foliage leaf, while anchor roots develop nearby at the insertion of the subadjacent prophyll. In Araceae with root dimorphism anchor roots are more numerous than feeder roots at the same node. The morphology and physiology of anchor and feeder roots are markedly different. The former are typically relatively narrow, agravitropic and appear to be negatively phototropic, which tends to bring them into contact with the substrate. Numerous root hairs are typically produced on the side adjacent to the substrate, but may develop all over the root (Troll 1941). Anchor root

growth is relatively limited and often dependent upon contact with a substrate. With no contact, growth ceases precociously, whereas prolonged contact stimulates elongation. Anchor roots never attain the enormous lengths of feeder roots. In Monstera (Madison 1977a) the stems not in contact with a substrate produce only a tuft of dried anchor roots, while those in contact produce anchor roots 20–30cm in length. The latter may surround tree trunks, for example. Root hairs serve to attach the root to a substrate, often forming a pseudoparenchymatous layer (Went 1895). Some anchor roots may lack root hairs and appear to be cemented by dried mucilage, as in Syngonium. Feeder roots are thicker than the associated anchor roots and are capable of considerable elongation. In some species of Monstera they may extend 30m to the forest floor (Madison 1977a). Feeder roots are considered positively gravitropic, although their tips do not hang vertically in some cases (Linsbauer 1907). Feeder roots may hang freely in the air for their entire length or they may adhere to a tree trunk, e.g. Syngonium (Troll 1941) even though no root hairs are produced. Mucilage may play a role in their adhesion as it is often more abundant than on anchor roots (Went 1895). Feeder roots do not generally branch before reaching the soil unless injured. However, Philodendron melanochrysum and some Syngonium species normally produce aerially branched feeder roots (Troll 1941). The lateral roots of decapitated feeder roots also lack root hairs and show positive gravitropism. Once the typically unbranched feeder roots of most species enter the soil they branch profusely. In Monstera (Madison 1977a), dimorphic roots do not appear until plants have reached about a metre above the ground and the stem is 7–10 mm thick. After a prolonged period without contact, as in a pendent shoot, only tufts of non-growing anchor roots are produced. Hinchee (1981) studied root development in Monstera before and after entry into the soil. Unfortunately Hinchee used the term Haftwurzeln erroneously, applying it to feeder roots prior to their entry into the soil, and reserved the term Nährwurzeln for their subterranean branches. This misapplication of terminology invalidates comparisons that are made with previous studies of true anchor and feeder roots (Guttenberg 1968) since Hinchee did not study Haftwurzeln. Some climbing Araceae, such as Scindapsus pothoides (= S. hederaceus) do not show a clear distinction between anchor and feeder roots (Went 1895). In these instances anchor roots develop all over the stem, but are larger and longer at the nodes and may reach to the soil. Internodal roots are also found in

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Anthurium sect. Polyphyllium (Croat & Baker 1978), as in A. clidemioides. Some feeder roots of Araceae are capable of contraction (Wettstein 1904, Rimbach 1922). Anchor roots, on the contrary, do not appear to contract. Rimbach found that contraction was absent in feeder roots of Monstera deliciosa but present in Philodendron bipinnatifidum where he observed that a feeder root contracted by about 20% over 5 months.

Root growth The average length of the elongation zone of Philodendron species has been observed as 20–50 mm, with the maximum found in P. selloum (= P. bipinnatifidum), amounting to over 90 mm. Hinchee (1981) reported that the elongation zone in Monstera aerial (probably feeder) roots may be as long as 105 mm. Actual rates of growth for Philodendron aerial roots were less than 10 mm/day for most species. The large roots of P. selloum (= P. bipinnatifidum) grew exceptionally fast, from 7–21.5 mm/day. Daily growth rates in Monstera deliciosa ranged from 3–40 mm/day for aerial roots. According to Hinchee (1981) aerial roots of Monstera exhibit an exponential increase in growth rate during aerial growth. Once roots enter the soil their rate of growth declines and maintains a relatively constant value.

Adventitious roots Adventitious roots arise from the inner stem cortex at some distance from the shoot apex, and they often have a broad attachment to the vascular cylinder of the stem, e.g. in some Pothoideae and Monsteroideae (French & Tomlinson 1981a, b). The adventitious roots of Pothos, Pothoidium and Heteropsis have a close spatial relationship to leaf traces. In Pothos and Pothoidium each lateral leaf trace diverges from the central cylinder to the leaf through the region of the root trace attachment. In Heteropsis only the mid-vein leaf trace diverges through the region of root attachment (French & Tomlinson 1981a). The origin of lateral roots in Araceae is typical of many monocotyledons and has been studied in Anthurium, Colocasia, Monstera and Zantedeschia. Pistia is atypical in several respects. Firstly, lateral roots arise very close (110–350 µm) to the root cap and apex junction (Charlton 1983). The lateral root primordia of Pistia tend to arise regularly along the length of the xylem strands in the central cylinder, but no evidence was found for regular spacing between the rows of root primordia (Charlton 1983). Secondly the roots of Pistia are unusual in the Araceae in their development of a persistent pocket or “Tasche”, which is a kind of “substitute” root cap. This structure also occurs in some other aquatic monocotyledons, e.g. Lemna, Spirodela, Eichhornia (Guttenberg 1968).

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Nest roots Another type of aerial root has been described in Anthurium ellipticum and is characteristic of many other species of Anthurium sect. Pachyneurium, the “bird’s nest” or “litter basket” anthuriums. This is the socalled nest root (Schimper 1888, Bruhn 1910). These exhibit negative gravitropism (Troll 1941) and branch profusely forming an “impenetrable” nest. The development of these roots was studied by Bruhn (1910) who found that their formation was typically inhibited when the roots were surrounded by moist moss or earth. Bruhn concluded that they resulted from damage to the root apex. However, more recent field studies suggest that nest roots are mainly for feeding. In these epiphytes the rosulate leaves form a “basket” into which leaves, twigs and other detritus gather. The nest roots seem to be especially adapted to exploit this food resource, growing directly up into the mass of detritus and ramifying within it (Croat 1991).

Contractile roots These roots occur in numerous Araceae which possess a tuberous habit, for example Typhonium (Banerji 1947), Arum (Rimbach 1897), other genera of subfamily Aroideae (Rimbach 1898), subfamily Orontioideae (Lysichiton, Orontium, Symplocarpus), some genera of subfamily Lasioideae (Hotta 1971), and some other aquatic species, as in Cryptocoryne, but not Calla (Dudley 1937). Detailed studies of root contraction have been made in two species of Arum. Rimbach (1897) followed the life history of Arum maculatum in the field, presumably in Germany, from seed germination to adult plant and studied in particular the phenology of shoot and root growth. Lamant & Heller (1967) studied the mechanism of contraction in Arum italicum. They observed that contraction was related to: 1) radial expansion of cortical parenchyma, which results in longitudinal tension, and 2) release of such tension by the collapse of proximal outer cortical tissues and shortening of the stele. Similar mechanisms occur in other contractile roots and were discussed in more detail by Ruzin (1979) and Jernstedt (1984). Galil (1978) found active lateral contractile roots in shallow, horizontally attenuated rhizomes of Arisarum, which pull the rhizome laterally rather than downwards. A second type of rhizome, which grows vertically at deeper levels, lacks contractile roots and is non-mobile.

Root epidermis The epidermis of most aroid roots is composed of a single layer of cells which nearly always includes both ground cells and root hairs. More than one epidermal layer has been reported for a number of genera,

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including Aglaonema, Anthurium, Gonatopus and Homalomena (Lierau 1888, Solereder & Meyer 1928), but these results were largely based on studies of mature roots only, which do not provide conclusive evidence. Root hairs were reported to be absent from subfamily Calloideae (sensu Krause 1908) but the entire family is clearly not “without root hairs” as described by Cronquist (1981). In fact two types of root hair development have been described (Leavitt 1904). In Type I, any protodermal cell may form a root hair, as in Aglaonema, Arisaema, Caladium, Dieffenbachia and Zantedeschia. In Type II, root hairs arise only from specialized trichoblasts, as in Anthurium and Monstera. Root hairs are usually not formed on feeder roots of hemiepiphytes until they penetrate the soil (Solereder & Meyer 1928, Lierau 1888, Guttenberg 1968, Hinchee 1981). However, anchor roots form abundant root hairs, in some cases from every cell (Guttenberg 1968). In many species the root hair tips of anchor roots are modified by branching or are flattened and become cemented to the substrate and to each other, possibly by exudates from the root (Richter 1901, Lierau 1888, Madison 1977a). Mucilage has frequently been observed on roots of climbing Araceae (Went 1895, Guttenberg 1968). Unusual scales have been described on the roots of Pothos and Monstera resulting in a striped appearance (Richter 1901). One of the most distinctive anatomical features of the roots of some Anthurium species is the presence of a true multiple epidermis which resembles the velamen of Orchidaceae and some other monocotyledons (Guttenberg 1968) in having a thick water-absorbing layer that is white when dry. Limited developmental studies have shown that the protoderm of these species exhibits tangential cell divisions resulting in multiple layers of epidermis (Guttenberg 1968). In some Anthurium species the velamen has long been recognized (Leitgeb 1865, Tieghem 1867, Lierau 1888, Schimper 1888). The multiple epidermis of Anthurium gracile exhibits a white velamen when dry (Croat 1984). Engler (1920b) noted that the multiple epidermis of some Anthurium species, particularly in section Pachyneurium, contains dead cells with fibrous thickenings and granular structures on the innermost tangential wall layer. Densely cytoplasmic “passage cells” have also been observed, which are similar to cells found in some Orchidaceae (Deshpande 1956). Fibrous thickenings have been observed in A. affine, A. crassinervium, A. maximum, A. willdenowii, among others, according to Lierau (1888), Leitgeb (1865), Schimper (1888) and Solereder & Meyer (1928). Some Anthurium species exhibit a smooth-walled multiple epidermis (A. huegelii (= A. hookeri), A. cucullatum (= A. andicola); Schimper 1888, Leitgeb 1865).

Root cortex In the roots of many Araceae an exodermis is present beneath the epidermis, which becomes the outer protective layer if the latter is lost (French 1987b). The exodermis or “Interkutis” is suberized and the outer walls may become thickened as in some Anthurium species. Patterns of cell shape are variable, with some roots having exclusively elongated exodermal cells, as in Calla. Other roots have alternating long and short cells, as in Monstera deliciosa (Sinnott & Bloch 1946, Hinchee 1981) or Anthurium (Leitgeb 1865). Although an exodermis is present in most Araceae examined so far (Olivier 1881, Schimper 1888, Solereder & Meyer 1928, Hinchee 1981, French 1987b), it is by no means universal. An exodermis is reportedly absent from Pistia (Guttenberg 1968). Beneath the exodermis a specialized multilayered sclerotic hypodermis develops in the roots of seven genera: Anubias, Cercestis, Culcasia, Furtadoa, Homalomena, Montrichardia and Philodendron (Lierau 1888, Solereder & Meyer 1928, French 1987a). The organization of the ground tissue of the cortex of Araceae varies considerably among different genera. In species with aerial roots the outer cortex contains chloroplasts. Collenchyma has also been reported in this region in feeder roots of both Monstera (Hinchee 1981) and Philodendron (Porsch 1911) as well as other root climbers. The ground tissue of the cortex is usually composed of relatively thin-walled, unlignified parenchyma cells. In many aquatic or semi-aquatic genera and some others, large schizogenous intercellular spaces, or lacunae, are present. These occur in Amorphophallus, Anchomanes, Calla, Colocasia, Dracunculus, Philodendron and Syngonium (Lierau 1888). The feeder roots of Syngonium develop large intercellular spaces, while the anchor roots do not (Went 1895). Trichosclereids have been reported in intercellular spaces only in Monstera deliciosa, Rhaphidophora decursiva and Scindapsus pteropodus (=Rhaphidophora pteropoda) (Lierau 1888, Solereder & Meyer 1928) despite their common occurrence in other organs. Thick-walled cells, some with pits, are mentioned by several authors (Lierau 1888) as occurring in the cortex of some Monsteroideae, but have not been described in detail. A variety of secretory tissues occurs in the cortex. Resin canals are present in the cortex of Philodendron, Homalomena and Furtadoa (Trécul 1865, 1866, Lierau 1888, Porsch 1911, Engler 1912) and in Culcasia and Cercestis (French 1987b). More details of organization are given in the section on resin canals. Laticifers also occur in the root cortex of a limited number of species, including Syngonium (Weiss 1866). Numerous crystalcontaining and tanniniferous idioblasts also occur in the root cortex (Solereder & Meyer 1928). The inner cortex typically contains narrower cells arranged in concentric rings adjacent to relatively smaller intercellular spaces in Amorphophallus, Colocasia, Dieffenbachia, Homalomena, Philodendron, Scindapsus,

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Syngonium and other genera (Solereder & Meyer 1928). The inner cortical region of many Monsteroideae and some Pothoideae contains a sheath of brachysclereids, fibres or sclerotic parenchyma, one to several layers thick (Tieghem 1867, Lierau 1888, Went 1895, Solereder & Meyer 1928, Sinnott & Bloch 1946, Hinchee 1981). This sheath may be in direct contact with the endodermis as in Spathiphyllum kochii (Lierau 1888) or separated from it by one or more cell layers. The latter condition occurs in Epipremnum pinnatum (Tieghem 1867), several species of Rhaphidophora (Solereder & Meyer 1928), Pothos celatocaulis (= R. korthalsii) (Lierau 1888) and P. scandens (Went 1985). An endodermis with a Casparian strip is probably present in roots of all Araceae (Solereder & Meyer 1928). Particularly in older aerial roots, the walls of the endodermis become thickened and suberized, thus obscuring the Casparian strips. In most species suberization begins first in the endodermis adjacent to the phloem and only later progresses to the region adjacent to the xylem. This pattern is typical of monocotyledons.

Vascular cylinder of roots The vascular cylinder of roots of Araceae is typically cylindrical and exhibits an alternating, radial arrangement of xylem and phloem. However, lobing of the cylinder occurs in some species of Philodendron and in Cercestis. Some species of Epipremnum and Philodendron have converging V-shaped xylem strands with narrower phloem strands in their angles and between them. Anomalous organization of vascular tissues occurs in some Monstereae (Monstera, Rhaphidophora, Scindapsus), Anthurium and Philodendron. In these genera the vascular region of the central cylinder consists of interspersed strands of phloem and xylem. The latter are generally one vessel in width while the phloem strands comprise several wide sieve elements (Hinchee 1981). Ground tissue of the central cylinder often becomes sclerotic in older roots. A pericycle has been reported (Hinchee 1981) and a pith region may be present or absent. Meyer (1925) followed the course of the inner vascular tissue in roots of Anthurium and Monstera and found that phloem and xylem strands remained separate in Anthurium wagenerianum. In Monstera deliciosa connections were observed between vascular strands, both in the periphery and the centre of the root.

Root apex Relatively few studies of apical organization have been made in Araceae. Guttenberg (1968) described an open pattern of apical organization. Hinchee (1981) demonstrated the presence of a quiescent centre in both the feeder root and its subterranean branches. More attention has been given to the structure of

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THE GENERA OF ARACEAE

the root caps of dimorphic aerial roots. Earlier workers (Haberlandt 1914) reported the presence of normal “statolith” starch grains in the columellas of gravitropic feeder roots of Monstera deliciosa (and other species), which has been confirmed by Hinchee (1981). In agravitropic clasping roots it is generally agreed that the size of the columella is smaller than in feeder roots and fewer “statocytes” are present (Haberlandt 1914). Several reports contend that columella starch grains are “sluggish or motionless” in clasping roots (Gaulhofer 1907, Haberlandt 1914), however Linsbauer (1907) found mobile grains. Development of the mature cortex was studied in Monstera deliciosa by Bloch (1946), Sinnott & Bloch (1946) and Hinchee (1981). The exodermis has alternating short and long cells resulting from unequal divisions that occur 2.5–3.5mm from the apex. Beneath the exodermis certain cells in longitudinal files undergo unequal divisions resulting in a smaller basal cell which then develops into a trichosclereid (Bloch 1946). The trichosclereid initials develop adjacent to intercellular spaces into which the cell arms elongate intrusively. Bloch (1946) found trichosclereid initials only at the distal ends of files, but Hinchee (1981) reports finding small, densely cytoplasmic, putative trichosclereid initials elsewhere. Asymmetric cell divisions were also associated with the formation of raphide cells by Kovacs & Rakovan (1975), but were not observed by Hinchee (1981). Sinnott & Bloch (1946) emphasized the role of unequal cell division, internal environment and cell position in the differentiation of cells of the exodermis and trichosclereids. In contrast, the development of the inner layer of brachysclereids adjacent to the endodermis is not dependent on unequal cell division. Similar short sclereids develop at the surface in response to wounding of the root, by redifferentiation of their walled cortical cells, which Sinnott & Bloch (1946) interpreted as an effect of a changed environment.

Periderm formation A cork cambium is generally formed in the cell layer below the exodermis in aerial roots of Monstera (Richter 1901, Hinchee 1981), Anthurium and Rhaphidophora (Olivier 1881), and replaces the exodermis as the outermost protective layer. In Monstera deliciosa the periderm begins to develop about 60 mm from the root apex in feeder roots and 10–15 mm from the apex of subterranean lateral roots (Hinchee 1981). Following cork formation, epidermal cells collapse in Monstera and Philodendron (Engler 1912). Cork may consist of alternating layers of thin-walled and sclerenchyma cells in Monstera (Hinchee 1981). In Philodendron the phellogen arises under layers of the sclerotic hypodermis and forms a wide layer of periderm in some species (Engler 1912). Lenticels also occur in the roots of Anthurium and other species (Weisse 1897). The lenticels of Monstera contain filling tissue as well as closing layers.

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Stem Stem anatomy and morphology are frequently correlated in Araceae, particularly with respect to the overall organization of the vascular system (Engler 1920b, French & Tomlinson 1981a–d, 1983, 1984). Conducting tissue tends to be highly condensed in tuberous species and in rhizomes with short internodes, compared with scandent plants. Climbing genera are largely concentrated in subfamilies Pothoideae and Monster oideae, but climbing hemiepiphytes occur in subfamily Aroideae as well, including Cercestis and Culcasia (tribe Culcasieae), Philodendron (tribe Philodendreae) and Syngonium (tribe Caladieae). Grayum (1984) pointed out that it is sometimes difficult to distinguish between rhizomes and tubers in the Araceae. Tubers are common in subfamily Aroideae (tribes Areae, Arisaemateae, Arisareae, Arophyteae, Caladieae, Colocasieae, Nephthytideae, Spathicarpeae, Thomsonieae, Zamioculcadeae, Zantedeschieae, Zomicarpeae). Hotta (1971) drew a distinction between two types of tuber in Araceae, the Amorphophallus type which is also found in Zamioculcas and Gonatopus, and the Arisaema type, which also occurs in Remusatia. The principal difference between them lies in the way new tubers form and the ease with which they are separated. In Amorphophallus, successive tubers develop from each other without separating, while in Arisaema new tubers are separated from the older tissue by cork layers. Stem epidermis The stem epidermis exhibits many of the same features as that of the leaf with regard to occurrence of trichomes, which are discussed in the section on leaves. Older stems of many Araceae develop a periderm in the outer cortex or epidermis which ultimately leads to the death of the epidermis (French & Tomlinson 1981a–d, 1983).

Stem cortex The cortex is delimited from the central cylinder by its less dense clustering of vascular bundles, and sometimes by the presence of an endodermis. The principal sources of histological variation in the cortex are: 1) width, 2) vascular organization, and 3) various specializations of the ground tissue. In many Monsteroideae, some Pothoideae and other genera like Culcasia, Cercestis and Philodendron, an especially distinct inner cortical boundary is present because the sclerotic sheaths of the peripheral vascular bundles of the central cylinder are fused together. In many Monsteroideae the inner layers of the cortex also become sclerotic. Some species, like

Calla palustris, have no endodermis and widely separated central cylinder bundles. Root traces are typically inserted along the periphery of the central cylinder and serve as useful markers. The vascular bundles in the cortex generally belong to two different systems: 1) leaf traces that traverse the cortex but may remain there for a longitudinally short distance, never exceeding one complete internode, and 2) cortical vascular bundles that constitute a separate system, persisting in the cortex over many internodes. In all species in which leaf traces are followed basipetally (using cinematographic techniques), they appear to enter the cortex from the leaf at the node and traverse it before entering the central cylinder, where they ultimately attach to axial bundles. There is considerable variation in the relative distance over which leaf traces remain in the internode before entering the central cylinder. In most species the major leaf traces enter the central cylinder close to the point of leaf insertion after traversing the cortex at an acute angle, as in Anthurium polyschistum (French & Tomlinson 1981a). The minor leaf traces may remain in the inter-node for a variable distance, in some cases extending to the base of the subjacent internode. Fibre bundles enter the cortex from the leaf in Homalomena and Pothos (and Acoraceae) and end blindly in the inner cortex (Tieghem 1867, Falkenberg 1876, French & Tomlinson 1981a,d) in a manner similar to palms and some other monocotyledons (Zimmermann & Tomlinson 1967, Zimmerman, Tomlinson & LeClaire 1974). A separate cortical vascular system is present in a few genera and tribes, including Anthurium, Caladium, Chlorospatha, Dieffenbachia, Heteropsis, Monstereae, Pothoidium, Pothos, Philodendron, Syngonium and Xanthosoma (French & Tomlinson 1980, 1981a–d, 1983, 1984, Grayum 1984). In addition to the persistent cortical bundles there may also be a series of minor and intermediate leaf traces. It is not uncommon for minor leaf traces to fuse with the cortical system and remain separate from the central cylinder. Cortical vascular bundles show anatomical diversity amongst the genera examined. However, they can usually be recognized in single sections by the relatively small amount of vascular tissue as compared to that of leaf traces. The latter tend to be wider and if they are persistent they are located nearer to the central cylinder. In many Caladieae with a cortical vascular system, the cortical bundles are narrow and form a highly anastomosing system, as in Chlorospatha longipoda and Xanthosoma tarapotense. Laticifers are typically present and are associated with the phloem in Caladium, Dieffenbachia, Philodendron and Xanthosoma. The ground tissue of the cortex generally consists of unlignified parenchyma with small intercellular spaces, except in aquatic genera such as Calla and Montrichardia, where a more extensive system of schizogenous intercellular spaces develops.

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Trichosclereids typically develop in the intercellular spaces of subfamily Monsteroideae, Pothos and Pothoidium (French & Tomlinson 1981a). Chloroplasts are present in the ground tissue beneath the epidermis. A continuous ring or separate strands of collenchyma may develop in the peripheral cortex of only certain genera, including Syngonium (Birdsey 1955), Dieffenbachia (Tieghem 1867), Aglaonema, Asterostigma, Homalomena, Philodendron, Schismatoglottis, Spathantheum and Zantedeschia (Engler 1920b). A wide variety of secretory tissues is present, including resin and mucilage canals and cavities, laticifers and a variety of idioblasts. Many of these structures also occur in other parts of the plant and are discussed later. The inner boundary of the stem cortex is sometimes demarcated by an endodermis with casparian strips, and has been described from Rhaphidophora celatocaulis (= R. korthalsii) (Solereder & Meyer 1928), Scindapsus pictus and Anthurium (Solereder & Meyer 1928) as well as various species of Amydrium, Monstera, Orontium, Rhaphidophora, Spathiphyllum, Symplocarpus, tribe Schismatoglottideae, tribe Peltandreae and Pistia (French & Tomlinson 1981a–d, 1983). There is some positive correlation between the aquatic and rhizomatous habits and the occurrence of an endodermis, but there are also aquatic genera that lack an endodermis, including Calla, Cryptocoryne and Lagenandra. Stem endodermis has been sought in the majority of Araceae genera and shown to be absent (French & Tomlinson 1980, 1981a–d, 1983, 1984). When an endodermis is present it may have several patterns of distribution, including unusual types (French & Tomlinson 1981a–d, 1983). It may encircle the entire central cylinder, interrupted only where leaf traces depart, as in Orontium and Symplocarpus. In Scindapsus and Rhaphidophora the endodermis may develop only on the ventral side of the central cylinder, in association with a vascular plexus of root traces to which the roots attach. An unusual pattern is found in tribes Schismatoglottideae and Peltandreae. The endodermis with casparian strips has two different patterns in these tribes: 1) it surrounds only the individual peripheral axial bundles of the central cylinder (i.e. no endodermis is present around interior bundles or around the entire central cylinder), or 2) it surrounds all of the axial bundles of the central cylinder but does not surround the entire central cylinder or individual leaf traces. The first pattern occurs in some species of Schismatoglottis, Hottarum and Typhonodorum. The second is present in some species of Aridarum, Bucephalandra, Peltandra, Phymatarum, Piptospatha and Schismatoglottis. The occurrence of an endodermis around individual vascular bundles is unusual in angiosperms (cf. Gunnera) but does occur in some ferns and in Equisetum (Esau 1965).

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Organization of stem central cylinder The central cylinder contains ground tissue and vascular bundles that are generally dispersed throughout. In many species, the bundles are somewhat more closely spaced towards the periphery and have a smaller diameter than those closer to the centre of the axis. The ground tissue is rarely lignified except in Pothos, Pothoidium, Heteropsis and in Philodendron subgen. Pteromischum, where usually the peripheral and sometimes the entire ground tissue is lignified. A wide range of secretory elements and idioblastic cells occurs both in the ground tissue and in association with the vascular tissue. The general principles of vascular organization in Araceae follow those of palms and other monocotyledons (Zimmermann & Tomlinson 1967). According to these concepts there are two kinds of vascular bundles in the central cylinder: 1) leaf traces that ultimately enter a leaf when followed distally, and 2) axial bundles that remain in the central cylinder. The leaf traces have protoxylem, which easily distinguishes them from axial bundles, which have only metaxylem. Leaf traces of some genera such as Cercestis and Syngonium can be distinguished from axial bundles by the presence of associated laticifers (French & Tomlinson 1981c, 1983). Leaf traces have a collateral organization throughout the family. Four general types of axial bundles can be recognized: simple collateral bundles, compound bundles, intermediate forms between compound and amphivasal bundles, and amphivasal bundles, including frequently branched forms that seldom have a complete cylinder of xylem. There is a structural continuum between all four categories (Tieghem 1867, French & Tomlinson 1981a–d, 1983, 1984). Compound bundles in Araceae were first observed by Tieghem. He correctly interpreted the presence of these in some species of Philodendron and in Dieffenbachia (French & Tomlinson 1981d, 1984), but did not investigate Montrichardia, Rhodospatha and Stenospermation, which also have compound bundles. Tieghem thought that all Araceae with unisexual flowers had compound bundles. However, in virtually all cases, e.g. Syngonium (French & Tomlinson 1983), these are not true compound bundles but various types of amphivasal bundles that branch and anastomose among themselves. The essential difference between the two types hinges on whether the distinct vascular components can be recognized, as in compound bundles, or whether the xylem elements tend to form a cylinder surrounding a core of phloem, as in amphivasal bundles. In most genera the distinction is clear. However, in Philodendron a wide array of bundle organization is present from simple collateral to compound and amphivasal (French & Tomlinson 1984), and numerous intermediate types are also present. Axial bundles in Dieffenbachia show a similar variation in organization and cannot readily be categorized.

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Simple collateral axial bundles occur infrequently in Araceae and tend to be present in genera with bisexual flowers (Tieghem 1867), although there are numerous exceptions. From the systematic and morphological viewpoints, the distribution of collateral bundles is interesting. They occur in all genera with bisexual perigoniate flowers except Gymnostachys, tribe Spathiphylleae, and subfamilies Orontioideae and Lasioideae. Most genera with collateral bundles have elongated internodes and many are climbers. Most genera with bisexual perigoniate flowers but lacking collateral bundles, such as Spathiphyllum, Holochlamys, subfamily Orontioideae, most Lasioideae and Gymnostachys, have relatively congested internodes, suggesting a strong correlation between this type of vascular bundle and habit. Except for Philodendron and Syngonium, Araceae with unisexual flowers are rarely scandent and amphivasal bundles predominate. Exceptions include collateral bundles in the climbing or prostrate plants of the genera Culcasia and Cercestis. The presence of collateral bundles tends to be correlated with prominent sclerenchyma sheaths, which are well developed in many species. Considerable sclerenchyma occurs in the rigid stems of Pothos, Pothoidium, Heteropsis and Philodendron subgen. Pteromischum. The habit of these genera is also distinctive in the Araceae because the climbing stems give rise to lateral branches which arch outward and hang down forming shrubby or tangled masses of branches. It is possible that increased stem rigidity in these genera is functionally correlated with their habit (Grayum 1984). The second major type of vascular bundle in Araceae is the compound bundle, recognized by Tieghem (1867), which typically consists of two to five or more simple collateral bundles clustered together within a common bundle sheath. Two major characters are used to identify the compound bundle. First, component vascular bundles tend to remain clustered as they follow a common course through the stem, although the distance over which the components remain together is variable. Second, individual components consist of discrete collateral bundles with their phloem strands directed towards the centre of the cluster. In some Araceae there are compound bundles with discrete components that remain together over relatively long distances, as in Rhodospatha (except R. venosa), Montrichardia, Philodendron (some sections only), Stenospermation and Dieffenbachia (some species). In other Araceae there are complexes of vascular tissue that migrate together but have less discrete components, in that the phloem tends to form a solid core rather than separate strands. Examples include Zamioculcas, adult stems of Cercestis, many species of Philodendron and some species of Dieffenbachia. These examples lead to the conclusion that it is not possible to precisely circumscribe compound bundles from a structural viewpoint, since there is a continuous intergradation with amphivasal bundles.

Typical amphivasal bundles with a core of phloem that is irregular or circular in transverse section are also present in Acorus and Araceae, as in many Colocasieae and Caladieae. The phloem is surrounded by xylem that contains tracheary elements having variable patterns of organization; they may be relatively narrow and contiguous (Cryptocoryne) or exist as individual elements, or clusters that have variable spacing (Philodendron).

Course of vascular bundles The three-dimensional organization of stem vasculature of Acorus and about sixty genera of Araceae has been analyzed using cinematic techniques (French & Tomlinson 1980, 1981a–d, 1983, 1984). These methods are necessary to understand the complex organization of stem vasculature in monocotyledons, which contains numerous vascular bundles in a dispersed arrangement. A wide range of types of organization was revealed and these can be grouped conveniently into four patterns. Pattern 1 is similar to that of some palms such as Rhapis (Zimmermann & Tomlinson 1967), and involves a continuing axial bundle apparently following a sigmoid curve through the axis, giving rise to branches which become leaf traces as they bend towards the periphery. Developmental studies in palms have shown that the leaf trace actually branches to form the continuing axial bundle. The Rhapis-type pattern occurs in Acorus, which has amphivasal bundles (Mangin 1880). Mangin’s remarkable study is probably the earliest accurate account of the “palm” pattern in monocotyledons and has unfortunately been long neglected. In Acorus, axial bundles tend to become part of a branched network of peripheral vascular bundles after the departure of leaf traces. This pattern has also been observed in palms such as Chamaedorea. The palm type occurs in Anthurium polyschistum and the rhizome of Spathiphyllum cannifolium, which both have simple collateral bundles (French & Tomlinson 1981a). Culcasia saxatilis and Philodendron hederaceum also have this basic organization. Pattern 2 is generally similar to the palm type because at regular intervals a continuing axial bundle branches from the leaf trace. The principal difference is the occurrence of unpredictable branching and anastomosis of axial bundles between departures of leaf traces. This pattern occurs widely in Araceae. It is present in subfamilies Calloideae and Orontioideae, most Lasioideae, and most Aroideae (French & Tomlinson 1980, 1981a–d, 1983, 1984). Highly condensed pattern 2 vascular systems are typical of species with tuberous stems and rhizomes, or erect stems with condensed nodes, which are common in subfamilies Lasioideae and Aroideae. Individual axial bundles in pattern 2 typically have some variation of the amphivasal pattern of organization.

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Pattern 3 exhibits a relatively distinct organization and is restricted to a few genera of Araceae. It has not been described in any other monocotyledon. The basic feature of pattern 3 involves the formation of a series of narrow “bridges”, or lateral branches from both leaf traces and axial bundles basally, which then aggregate to form a new axial bundle that distally departs as a leaf trace into a leaf. The pattern is referred to as the “basal aggregation” type because axial bundles appear to arise by the aggregation of a series of bridges or branches. Basal aggregation occurs in Anadendrum, Pothoidium, Pothos and certain Monstereae, including Amydrium, Epipremnum, Monstera, Rhaphidophora, and Scindapsus. Pattern 4 includes a relatively small number of genera with distinct compound vascular bundles, including Dieffenbachia, Montrichardia, Rhodospatha (all but R. venosa), Stenospermation, some Philodendron spp., and genera such as Cercestis and Zamioculcas, with intermediate conditions. In the compound bundles of Montrichardia, Philodendron, Rhodospatha and Stenospermation, the individual components are relatively distinct and can remain independent over several millimetres. A predictable pattern of changes in course and number of components in a bundle was not detected, in contrast to the compound bundles of Cyclanthaceae (French, Clancy & Tomlinson 1983) and Pandanaceae (Zimmermann, Tomlinson & LeClaire 1974). Components may fuse together laterally, separate from the compound bundle or may split apart in various patterns. New bundles may be added from surrounding compound or simple bundles. In these genera there is an apparently random series of interchanges between components both within one bundle and between adjacent bundles. This unpredictable feature also occurs in Dieffenbachia, in which sclerenchyma is minimal and bundle components are not well separated in some species. In Dieffenbachia and in some Philodendron species there are closely spaced bifurcations of bundles and loose association of components, which intergrade with amphivasal bundles. A similar situation exists in Cercestis (adult stem) and in Zamioculcas.

occur in climbing Cyclanthaceae (French, Clancy & Tomlinson 1983). Three major patterns of bud trace insertion are recognizable in the Araceae. In the first pattern two clusters of bud traces follow an arching course from the bud into the central cylinder and through the subperipheral or peripheral area. The second pattern also involves the formation of distinct clusters, but these migrate around the outside of the central cylinder in the inner cortex. They subsequently enter the central cylinder with leaf traces and often form a cluster around an individual trace. The third pattern is largely inconspicuous and nondescript. It involves the fusion of a few traces directly with the surface bundles of the central cylinder opposite the bud. This pattern can intergrade with the first type and is typically associated with the rhizomatous or tuberous habit. The formation of extensive bud trace systems (patterns 1 and 2) in climbing species of Araceae appears to be an adaptation which ensures adequate vascular connections with lateral branches that have no other immediate source of nutrients.

Internode development This has been studied in a small number of climbing species, using measurements of cell length, mitotic index and marking studies of elongating internodes (Fisher & French 1976, 1978, French & Fisher 1977b, French 1977). The stems of these Araceae contain socalled uninterrupted meristems, in which an acropetal wave of cell maturation proceeds through successive internodes, involving an end to cell division, an abrupt increase in cell length, and ultimately cessation of growth. Marking experiments show that elongation continues for a longer time and at a slightly higher rate in the upper region of the internodes of Anthurium, Epipremnum and Philodendron, leading to differential growth of the upper region (French & Fisher 1977b, French 1977).

Leaf

Epidermis Bud traces Both the organization of bud traces and their pattern of insertion into the main axis vary considerably in Araceae. There are systematic correlations as well as associations with the basic biology of stem growth. Unfortunately this subject has not received much study in monocotyledons so there is minimal information for broad comparisons. While most Araceae have one bud per node, some, like Xanthosoma and Dracontium, develop multiple buds and others like Remusatia produce bulbils (Engler 1920b, Möbius 1935). Some similar types of bud trace insertion also

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THE GENERA OF ARACEAE

Trichomes and larger emergences such as prickles are highly unusual in leaves or stems of Araceae (Solereder & Meyer 1928, Grayum 1984) and detailed anatomical descriptions are usually lacking. Unicellular hairs occur in some species of Xanthosoma (e.g. X. pubescens) and short 2-celled hairs are reported for Anubias barteri (Solereder & Meyer 1928). Multicellular hairs occur in abundance on leaves and stems of some neotropical Homalomena species (section Curmeria), a few species of Schismatoglottis and in Pistia. Trichomes are also reported for leaves of Bognera (Madison 1980), Callopsis, and some species

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of Stylochaeton (Bogner 1984c, Mayo 1985a) but detailed anatomical details are not available. Large, scale-like or filamentous emergences are present on leaves of some species of Philodendron section Philodendron (syn. sect. Polyspermium) (Krause 1913, Engler 1920b). Scale-like emergences are found on stems of Syngonium podophyllum var. peliocladum (Birdsey 1955, Croat 1982). The petioles, leaf blades and stems of some genera are armed with sharp prickles, as in Anchomanes, Cyrtosperma, Lasia, Lasimorpha, Podolasia, Pseudohydrosme and occasionally Nephthytis (Grayum 1984). Some neotropical species of Homalomena have numerous recurved prickles on their petioles. Prickle-like emergences also occur on the stem of Montrichardia (Engler 1911, 1912, 1920b). The cuticle of Araceae varies considerably in thickness and morphology (Solereder & Meyer 1928), but no systematic study has been made. Cuticular ridges occur in a number of genera including Culcasia, Dieffenbachia, Peltandra, Pothos, Spathiphyllum and others (Dalitzsch 1886, Webber 1960, Pant & Kidwai 1966). One or both leaf surfaces may have ridges. Waxy deposits on the epidermis have been reported from Caladium, Colocasia, Lagenandra, Remusatia (Gonatanthus) (Solereder & Meyer 1928), Syngonium (Birdsey 1955), Carlephyton glaucophyllum and Xanthosoma violaceum (Bogner pers. comm.). The types of wax crystalloids in Araceae have a greater similarity to those of dicotyledons than monocotyledons (Behnke & Barthlott 1983). Hydrophilic globules occur on the upper epidermis of Orontium and lower epidermis of Peltandra. Franke (1967) and MaierMaercker (1981) have studied transpiration through the cell walls and cuticle of Zantedeschia. The organization of stomatal guard and subsidiary cells in the leaves of Araceae has been partially surveyed in mature leaves (Dalitzsch 1886, Solereder & Meyer 1928, Webber 1960, Pant & Kidwai 1966, Bunting 1968, Grear 1973, Grau 1983). Not surprisingly, there is great variability in the number of subsidiary cells between taxa and among individuals. The range is from zero (anomocytic) as in Orontium, to two (paracytic), or four to eight (tetracytic) in Rhaphidophora (a more complete list is given by Grayum 1984). The development of subsidiary cells in Araceae appears to follow the perigenous pattern, that is, to have guard and subsidiary cells from the same parent cell, which is typical of monocotyledons. Very few aroid stomata have been examined developmentally, however, which prevents the application of Tomlinson’s (1974) developmental system of classification. Araceae lack the “oblique” cell divisions found in other monocotyledons such as palms (Tomlinson 1974). In the leaves of most aroids the stomata are either present only on the abaxial surface (hypostomatic) or are largely concentrated there (see Grayum 1984 for a complete list). Shaw (1993) has studied the distribution of stomata on the leaves of Monstera deliciosa. In Pistia the leaves are largely epistomatic, i.e. stom-

ata are present on the adaxial surface only, while Orontium is entirely epistomatic. The guard cells are typically level with the other cells of the epidermis, but they may also be sunken, as in Orontium, Ariopsis and Pistia. Ground epidermal cells tend to have a polygonal outline, with 5–8 straight sides and the surface usually lacks papillae or trichomes. Undulate cell walls occur in one or both epidermal layers and over all or part of the anticlinal walls in a few genera. Exceptionally wide ground epidermal cells reaching 94 µm in width occur in Zamioculcas, whereas in other Araceae they range from 36–71 µm (Pant & Kidwai 1966). Epidermal cells of some genera are papillate, giving the leaves a velvety sheen (Dalitzsch 1886, Engler 1920b, Solereder & Meyer 1928, Birdsey 1955). The papillate epidermis of the spathe of Arum *is highly unusual in having intercellular spaces between the epidermal cells. This feature is part of a syndrome of adaptive characters linked to the pollination mechanism and is correlated with the absence of stomata in the spathe (Knoll 1923).

Leaf mesophyll The mesophyll of Araceae is predominantly bifacial, with a thicker spongy layer below the palisade (Dalitzsch 1886). Isobilateral leaves are rare and include some species of Anthurium and Montrichardia in which the spongy layer is virtually absent and the palisade is nearly uniform. In Typhonodorum two palisade layers are separated by a region of aerenchyma. Acorus calamus (Acoraceae) exhibits a band of 3–5 layers of chlorophyllous isodiametric cells beneath the epidermis, and a central region of large air cavities, which significantly vary in size in different populations (Kaplan 1970, Röst 1979b). The palisade parenchyma cells are generally short, relatively wide and may have arms, as for example in Gonatopus (Solereder & Meyer 1928). Long narrow palisade cells are of more limited occurrence and have been observed in some species of Anthurium (Dalitzsch 1886, Solereder & Meyer 1928). The spongy mesophyll consists of flattened cells with arms that are arranged in tiers, as in Anthurium and Anubias. The spongy mesophyll has a distinctive chambered structure in leaves of many species of Homalomena, Philodendron, Piptospatha, Schismatoglottis and Typhonium (Engler 1920b, Solereder & Meyer 1928, Bunting 1968). In Typhonodorum stellate parenchyma occurs in the partitions of such chambers. In some Araceae, a hypodermis is present, consisting of one to several layers of parenchyma cells lacking chloroplasts, as in Anthurium and other genera (Dalitzsch 1886, Solereder & Meyer 1928). In Philodendron Engler (1912) reported a cell layer without chloroplasts containing a red pigment lying adjacent to the lower epidermis. Williams (1994) made a more recent study of some Philodendron species.

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Leaf vasculature Early studies of leaf venation by Engler led him to conclude that it was not a highly significant character for the classification of Araceae (Engler 1920b). Nevertheless, he used venation as a major character in the definition of his subfamilies, especially his Philodendroideae and Aroideae (Engler 1920b), with parallel-pinnate and reticulate venation respectively. Ertl (1932), based on a survey of over 40 genera, developed an explanation for the apparently variable nature of venation in Araceae based on differences in spacing between the primary lateral veins. No detailed study has been made since, and although Hotta (1971) essentially endorsed Ertl’s conclusions, this hypothesis must be regarded as tentative. Ertl concluded that overall leaf size and shape, that is, broad versus linear, as well as degree of dissection, have an important relationship to venation pattern. Although these morphological and developmental correlations are important, systematic position is also significant. Some groups, such as tribes Colocasieae and Caladieae, have a characteristic venation pattern that is independent of leaf shape. In these tribes there is a tendency for quaternary veins to arch over tertiary veins (Ertl 1932, Birdsey 1955). Ertl identified two extremes of venation pattern, Groups I and II, and an intermediate Group III. The continuous intergradation between the extremes is used as an argument for a single basic plan for araceous vasculature. Group I has basically parallel or parallel-pinnate (striate) venation and contains Gymnostachys, Calla, some Monsteroideae such as Stenospermation and Spathiphyllum, and the tribes of Aroideae grouped by Engler (1920b) into subfamily Philodendroideae (Aglaonemateae, Anubiadeae, Dieffenbachieae, Homalomeneae, Peltandreae, Philodendreae, Schismatoglottideae and Zantedeschieae), together with tribes Cryptocoryneae and Ambrosineae. Group II has a reticulate pattern, superficially like that of dicotyledons, and includes Anthurium, subfamily Lasioideae, and most other tribes of subfamily Aroideae including Pistieae. Group III contains Pothos, some Monsteroideae such as Monstera, and tribes Caladieae and Colocasieae. Ertl’s work was a pioneering study and there is a need for a new study of leaf venation using modern terminology and a broad taxonomic approach. Araceae petioles contain a variety of arrangements of vascular bundles but no detailed systematic comparison has been made. Transverse sections of petioles are shown in works by Engler (1905, 1911, 1912, 1915, 1920a, b), Ertl (1932), Troll (1939) and Birdsey (1955). The mechanical support tissue of the petiole may be in the form of a continuous peripheral ring as in Aster ostigma, Homalomena, Montrichardia, Philodendron, Spathantheum, or separate peripheral strands of collenchyma as in Anchomanes, Dieffenbachia, Dracontium,

18

THE GENERA OF ARACEAE

Philodendron, Schismatoglottis, Zantedeschia and most Caladieae and Colocasieae (Dalitzsch 1886, Birdsey 1955). In some genera such as Amorphophallus and Pseudodracontium, the vascular bundles have collenchymatous bundle sheaths (Dalitzsch 1886). In others, the support tissue is primarily bundle sheath sclerenchyma and may be fused together to form a peripheral layer (Anthurium, Rhaphidophora, Spathiphyllum). Fibre bundles may be present, with or without phloem, as in Anthurium. The vascular bundles may be rather distant from one another and have large sclerenchyma sheaths, as in Cyrtosperma and Lasia. In Orontium and Calla mechanical tissue is absent from the petiole (Solereder & Meyer 1928).

Leaf tubercles and regeneration Tubercles regularly develop at the juncture of leaflet and petiole in Pinellia ternata (Hansen 1881, Linsbauer 1934, Troll 1939), at the apical end of petiole in Typhonium bulbiferum (Sriboonma et al. 1994) and at the first and second order divisions of the leaf of Amorphophallus bulbifer (Troll 1939). Tubercles in Pinellia may also form spontaneously along the petiole or can be induced in the basal part by cutting into segments (Linsbauer 1934). Tubercles may develop in Typhonium violifolium at the leaf apex, the petiole apex and at the apex of the sheath (Sriboonma et al. 1994). Regeneration of tubers, leaves and roots from leaf segments is well known in Zamioculcas zamiifolia and Gonatopus boivinii (Engler 1881, Schubert 1913, Cutter 1962). Isolated entire leaflets of Zamioculcas and Gonatopus spontaneously develop a basal swelling, followed by the formation of roots and up to 3 buds, over a 6–9 week period for Zamioculcas. Leaf regeneration in Gonatopus is more rapid. The results of experimental manipulation of isolated leaflets grown in culture show that any part of the compound leaf is capable of regeneration, and that new shoots arise in association with the cut ends of the largest veins, preferentially at the proximal end. When midvein tissue is present, regeneration at lateral veins is absent, even though there is no vascular connection with the midvein. When only lateral veins are present, regeneration proceeds at their proximal ends (Cutter 1962). Plantlets may also regenerate from leaf blades in Schismatoglottis (Bogner, pers. obs.).

Geniculi (pulvini) A pulvinus (swelling) or geniculum (joint) occurs at the distal end of the petiole in nearly all Pothoideae, Monsteroideae and certain Aroideae (Anubias, Bognera, tribes Culcasieae, Zamioculcadeae, rarely in Philodendron and Homalomena). Details are given in the generic and tribal descriptions in this volume. The

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mechanical tissue in the geniculum consists of collenchyma in a peripheral ring or collenchymatous bundle sheaths, whereas elsewhere in the petiole it is formed by sclerenchyma (Dufour 1886, Dalitzsch 1886). The petiole of Gonatopus boivinii has a geniculum in the middle which can reorient the blade. Some reorientation is also possible from a second geniculum near the base of the leaf (Troll 1939).

Ligules Ligules were regarded by Engler as a modified leaf sheath with a free extension at its tip. They occur in Calla, various Schismatoglottideae, Dieffenbachia and some species of Philodendron. A membranous “stipule” has been reported in Pistia (Engler 1920a).

Leaf structure of Acorus (Acoraceae) Much attention has been devoted to the leaf of Acorus, especially to its early development. Kaplan (1970) presented several conclusions based on a rigorous comparative developmental study of Acorus. He concluded that its unifacial leaves are distinct from those of the Araceae because of the intense adaxial meristematic activity that begins during apical growth. Adaxial growth leads to a “bulge” that arches over the shoot apex. The leaf of Acorus is thus a foliar structure and not a flattened petiole. Earlier workers mistakenly interpreted young leaves as having two successive apices, one that led to a hyponastic arching primordium and a second that overtopped the original, a pattern referred to as sympodial growth. Kaplan (1970) observed that continued adaxial meristematic activity and suppression of the marginal meristems leads to the formation of the radially flattened leaf. A narrow secondary midrib does, however, develop. Morphologically and developmentally the leaf of Acorus is similar to the phyllode of Acacia, which also develops an active adaxial meristem. The phyllode theory of Arber, which attempted to interpret leaves of Acorus as flattened petioles, is thus based on an erroneous interpretation. It is interesting that leaves of different populations of Acorus calamus studied by Kaplan (1970) differ in their degree of apical growth, which is correlated with radial expansion. The Iowa population (probably a diploid) exhibited more prolonged apical growth and less radial growth than the Wisconsin population (probably a triploid). One would predict that the tetraploid should exhibit even shorter apical growth and greater radial expansion. Vascular differentiation in the leaf of Acorus begins first in the centre of the axis and later is bidirectional, as is the activity of a plate meristem at the leaf margins. Subsequent growth in length is accomplished predominantly through the activity of a basal intercalary meristem.

The leaf anatomy of Acorus has received much attention (Dalitzsch 1886, Solereder & Meyer 1928, Ertl 1932, Kaplan 1970, 1973a). Röst (1979b) was particularly interested in the biosystematics of Acorus calamus and A. gramineus and he discriminated between the two using a variety of leaf characters, including the organization of chlorenchyma, aerenchyma and sclerenchyma. Three morphologically distinct varieties of A. calamus are recognized (Röst 1979b). Each can be distinguished by ploidy level and on the basis of one leaf characteristic, namely the number of air canals in a prescribed area of a transverse section (0.62mm2) of the lamina above the sheath (Röst 1979b). The diploid variety has relatively few canals (less than 17), the tetraploid has numerous canals (more than 77), and the triploid variety is intermediate (26–51 canals). Using Röst’s criteria it appears that Kaplan’s (1970) Iowa population is the native American diploid (Acorus calamus var. americanus) and the Wisconsin population is the triploid (A. calamus var. calamus). No tetraploids were studied by Kaplan (1970). The triploid is sterile, mainly Eurasian, and probably was introduced to North America by European settlers.

Special Topics

Trichosclereids These are filamentous, branched sclerenchyma cells (sclereids) up to 7 mm long which develop in the intercellular spaces of vegetative and floral tissues of all genera of subfamily Monsteroideae except Anadendrum and Heteropsis, and two genera of subfamily Pothoideae, Pothos (Tieghem 1867, Engler 1920, Solereder & Meyer 1928, Nicolson 1959) and Pothoidium (French & Tomlinson 1981a). Trichosclereids rarely develop in roots. They played a significant role in Engler’s classification of the Araceae, which emphasized vegetative anatomical features. The term trichosclereid was first applied by Bloch (1946) and Sinnott & Bloch (1946) in their classic study of idioblast development in Monstera deliciosa. A variety of other terms had been used previously including “raphide”, which was in common use in the early 19th century to refer to anything needle-shaped (Nicolson 1959, 1960b). The term was confusing because raphides of calcium oxalate, which are crystals, also occur in the same tissue. Schott correctly illustrated trichosclereids in various monsteroid genera, but referred to them as “raphides” (Schott 1832, Schott 1858). Hasskarl (1842) carried this confusion one step further by naming a monsteroid genus Rhaphidophora on the basis of correctly illustrated trichosclereids. Schleiden (1839) correctly identified “raphides” (= trichosclereids) as “Bastzellen”, and more detailed accounts by other anatomists followed (Sueur 1866, Tieghem 1867, Wiesner 1875). Tieghem (1867) made

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a number of observations of trichosclereids as part of a general survey of the anatomy of the Araceae, and was the first to notice that they occurred primarily in monsteroid taxa, an observation used by Engler in defining subfamily Monsteroideae taxonomically (Engler 1920b). However, Engler emphasized the presence of trichosclereids to such an extent that he placed several genera which lack these cells but otherwise display clear monsteroid affinities in his subfamily Pothoideae. These genera, which included Anadendrum, Epipremnopsis (= Amydrium) and Heteropsis, are today widely recognized as belonging to subfamily Monsteroideae (Nicolson 1984a, Bogner & Nicolson 1991, this volume), and trichosclereids are now known to occur in Amydrium. As a result of various studies, a rather general understanding of the morphology, occurrence and development of trichosclereids has emerged (Nicolson 1960b). In the Araceae, trichosclereids have several distinctive features although there is considerable quantitative variation among species. They begin development as small, nearly isodiametric cells which form 1–4 hair-like outgrowths that elongate into the surrounding tissue, forming an irregular network within the intercellular spaces. Their branches grow apically, are often of irregular shape and may themselves branch. Trichosclereids that have two branches have a peglike central body which identifies the location of the original cell and is the site of attachment in the tissue. In contrast, other types of sclereid, such as libriform fibres, may be long and filamentous but do not have branches arising from a cell body. Trichosclereids may also have four outgrowths resulting in an H-shape. The number, diameter and length of the branches are the primary sources of variation. In Spathiphyllum trichosclereids appear to be longer and narrower than those from Rhaphidophora and other Monstereae (Tieghem 1867, Nicolson 1960b). It has long been recognized that in tribe Spathiphylleae there are more numerous trichosclereids in each intercellular space than in tribe Monstereae. Estimates of over 50 (Solereder & Meyer 1928) and between 15–20 (Nicolson 1960b) have been made. This contrasts greatly with the 1–4 trichosclereids in each intercellular space found in tribe Monstereae (Nicolson 1959). In addition, some trichosclereids intergrade with astrosclereids in foliage leaves, e.g. in Monstera. The distribution of trichosclereids in the plant body follows a general pattern throughout subfamily Monsteroideae (sensu Engler 1920b). They are usually present in the stem, petiole, leaf blade and inflorescence, but have been reported in the roots in only three species (Solereder & Meyer 1928). It is not uncommon for trichosclereids to be absent from one part of the plant body (Nicolson 1959, 1960b). In some Epipremnum species and Monstera punctulata, trichosclereids are absent from the leaf blade (Nicolson 1959, Madison 1977a).

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THE GENERA OF ARACEAE

Laticifers Early studies (Hanstein 1864, Trécul 1865, 1866) demonstrated the presence of articulated non-anastomosing laticifers in most subfamilies, including Calloideae, Lasioideae, Orontioideae and Aroideae, and articulated, anastomosing laticifers in tribes Caladieae and Colocasieae (except Ariopsis). Both clear and milky latex was described by early workers, who detected abundant tannins in the latex. The clear, tannin-rich latex of some Araceae has been compared with the exudate produced by articulated cells in the phloem of some Leguminosae (Trécul 1865, 1866, Esau 1974), which may also have perforated end walls. All early workers considered Araceae to have laticifers that produce latex. However, Solereder & Meyer (1928) took a more neutral position, refraining from the use of the term laticifers. They used the terms “secretory files” or “secretory tubes” to describe the non-anastomosing and anastomosing laticifers, respectively. Engler (1920b) emphasized the presence of anastomosing laticifers in defining his subfamily Colocasioideae (equivalent to our tribes Caladieae and Colocasieae combined). However, Weiss (1866) found that non-anastomosing laticifers are present in roots of Syngonium; he made no further observations on other genera of Engler’s subfamily Colocasioideae. Engler (1920b) also defined his subfamilies Pothoideae and Monsteroideae in part because of their lack of laticifers. Individual “secretory” cells, not arranged in files, are present in these taxa, but their chemical and structural similarity to laticifers remains unstudied. Laticifers in Araceae typically occur in association with vascular bundles, lying on the periphery of the phloem in the leaf and stem. In stems, laticifers occur only in leaf traces or cortical bundles, not in axial bundles (French & Tomlinson 1981c–d, 1983). In roots, laticifers also occur in association with phloem, but may be present in the xylem, pericycle or ground tissue (Weiss 1866, Lierau 1888). They are visible in sections because of dark-staining contents and a relatively wide lumen compared to adjacent phloem cells. Laticifers may occur in the root cortex in Philodendron and various Caladieae and Colocasieae (Weiss 1866, Porsch 1911). The most complete study of the systematic occurrence of laticifers is by French (1988 and unpublished observations), who examined leaves and inflorescences of 75 genera. Anastomosing laticifers are limited to the tribes Caladieae, Colocasieae and Zomicarpeae. Articulated, non-anastomosing laticifers occur in Calla, Orontium and almost all Aroideae (except tribes Pistieae, Stylochaetoneae, Zamioculcadeae). Tribe Cryptocoryneae have been previously reported as lacking laticifers, but recent unpublished work has demonstrated their presence in the stems, roots and cataphylls of both Cryptocoryne and Lagenandra, though not in the foliage leaves (Sivadasan, pers. comm.).

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Limited cytological observations of laticifers were made by Molisch (1899, 1901) and Southorn (1964). No clear picture of latex cytology has yet emerged. Schmid (1882) observed nuclei, small dense spheres and larger unidentified elliptical particles approximately the size of a nucleus in laticifers of Caladium. Molisch (1899, 1901) made the most extensive cytological study, using fresh latex. He observed socalled “Blasenkerne”, which represent nuclei having sac-like protrusions, in several genera, including Aglaonema, Philodendron, and Xanthosoma. Molisch also found what he called abundant leucoplasts in the latex of Steudnera colocasiifolia, as well as biconvex particles (7–16 µm in diameter) of unknown identity. In other studies Molisch (1901) noted that in latex of Amorphophallus konjac (syn. A. rivieri), large (16 µm) membrane-bound six-sided crystalloids are found. In the only recent study, Southorn (1964) reports the presence of vacuoles containing particles in Brownian motion in Dieffenbachia. Fox & French (1988) found that abundant terpenoid particles were responsible for the white appearance of the latex of tribe Caladieae, analogous to the white latex of Euphorbiaceae. The clear or cloudy latex of tribe Colocasieae lacks the abundant terpenoid particles. Larger latex particles were also found in the latex of “colocasioids” (i.e. tribes Caladieae and Colocasieae combined), as described by Molisch, and their occurrence was found to be systematically significant (French & Fox, unpublished results).

Resin canals and cavities The presence of resin canals in Araceae has been documented from the mid-nineteenth century (Trécul 1865, 1866, Tieghem 1867). However, Solereder & Meyer (1928) proposed the neutral term “secretion spaces” to refer to these canals in Araceae because in their opinion there was a lack of chemical evidence for the presence of resin. Resin typically contains large amounts of various terpenoids and other lipophilic substances and is thus chemically distinct from mucilage, which is hydrophilic (Fahn 1979). It is synthesized and accumulated by canals and cavities which are of limited occurrence in Araceae (Trécul 1865, 1866, Tieghem 1867, 1872, 1885). Resin is generally considered to be part of the plant’s chemical defence against herbivores. Resin canals are composed of a central cavity surrounded by 1–3 layers of synthetically active epithelial parenchyma. A sclerotic or collenchymatic protective sheath may surround the epithelium (Möbius 1885). Secretory cavities differ from canals principally in their shape which ranges from spherical to irregular. Both canals and cavities are reportedly schizogenous in origin (Leblois 1887, Engler 1920b). Resin canals were first reported in Araceae by Trécul (1865, 1866) who found them in leaves, stems and roots of Philodendron

and Homalomena species. Additional reports of resin canals and cavities in these two genera were made by Tieghem (1867, 1872, 1885), Möbius (1885), Dalitzsch (1886), Leblois (1887), Lierau (1888), Went (1895), Porsch (1911), Engler (1920b) and Pohl (1932a, b). The most complete study is a family-wide survey by French (1987b), which included 91 genera and 250 species and showed that resin canals occur in the roots of only five genera, Culcasia, Cercestis, Homalomena, Furtadoa and Philodendron. In the leaf of Culcasia the canals are easily visible with the unaided eye and exhibit a variety of differences in their length and orientation depending on the species (Knecht 1983). They are also visible without magnification in leaves of Philodendron. Recent studies have also confirmed their presence in the stems of Culcasia, Homalomena (cavities), Philodendron, and Cercestis (French & Tomlinson 1981a, c–d, 1983). Pohl (1932a, b) and Mayo (1986b, 1991) investigated the resin canals that are present in the inflorescence of Philodendron. The resin is released onto the spathe or spadix surface and plays a role in the floral biology in Philodendron by glueing the pollen onto the pollinator’s body (Gottsberger & Amaral 1984, Grayum 1990, Mayo 1991). In species of Philodendron subgen. Meconostigma (e.g. P. bipinnatifidum studied by Pohl 1931a, b as P. selloum), one end of the canal is close to the inner surface of the spathe and releases resin shortly before spathe closure. In Philodendron subgen. Philodendron resin is secreted onto the spadix surface from numerous resin canals located just below the axial epidermis, around the bases of the stamens and staminodes. In these species, the pollen and resin are mixed even before appearing at the spadix surface (Mayo 1986b).

Mucilage cells, canals, and cavities Mucilage contains various hydrophilic substances and can readily be distinguished from resin using appropriate histochemical procedures. It may occur in various idioblasts such as raphide tubes (Solereder & Meyer 1928), together with calcium oxalate crystals, or in large mucilage cells, as in Amorphophallus (Abranowicz 1912, Wakabayashi 1957a, b). Mucilage also occurs in canals and cavities which may form schizogenously, as in Epipremnum pinnatum (Leblois 1887) or lysigenously as in Colocasia (Solereder & Meyer 1928). Mucilage canals and cavities may have a distinct epithelium of smaller, more densely stained cells, and some epithelial cells may project into the locule or grow into it like a tylose (Engler 1920b). Mucilage canals are of relatively limited occurrence in Araceae. They occur in Aglaonema, Alloschemone, Alocasia, Anthurium, Cercestis, Colocasia, Epipremnum, Monstera, Philodendron, Remusatia, Rhaphidophora, and Xanthosoma (Solereder & Meyer 1928, French & Tomlinson 1981a–d, 1983, 1984, Boyce, pers. obs.).

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Extrafloral nectaries and punctations Zimmermann (1932) exhaustively reviewed the occurrence of extrafloral nectaries in angiosperms and listed reports of possible extrafloral nectaries in Arisaema (Knuth 1909), Colocasia (Solereder & Meyer 1928) and Philodendron, but nectar secretion is listed as very doubtful. Madison (1979a, b) reported nectar production by a ring of extrafloral nectaries at the base of the leaf lamina in Philodendron myrmecophilum (= P. megalophyllum), and sweet secretions are produced also by the extrafloral nectaries of prophylls in many species of Philodendron, and on the outside of the spathe in P. goeldii (Bogner, pers. obs.). In a description of the leaf of Araceae, Engler (1920b) made no specific mention of extrafloral nectaries, but “sessile superficial glands” were reported in Alocasia by Gardiner (1883, 1889) and BelinDepoux (1978). The anatomy of the “glandular” structures in Culcasia was described by Solereder (1919) who considered them possibly to be extrafloral nectaries (Solereder & Meyer 1928). Solereder (1919) and others (Tieghem 1867, Bachmann 1880, Dalitzsch 1886, Gentner 1905, Engler 1920b) have described the mature structure of the unusual “punctations” on the leaves of various species of Anthurium. They are often referred to as glands or glandular punctations and Dalitzsch (1886) reported seeing secretions released internally, but this has not been confirmed. Each mature punctation consists of three regions: 1) a basal region of 1–3 curved layers of prismatic cells in files embedded in mesophyll, 2) a central region of large, radially elongated cells lacking dense contents, and 3) an outer cluster of compact cells with dark brown or black contents beneath a stomate. No secretion has been observed to exude from the surface punctations and the chemical identity of the dark contents is unknown. Solereder (1919) pointed out the similarity of these structures to the cork warts of some dicotyledons, also mentioned by Bachmann (1880).

Hydathodes Hydathodes occur in the leaf tips of many Araceae, and have attracted considerable attention because of the high rate of guttation in some species. Guttation has been studied frequently in Colocasia, in which rates of over 100 ml/day/leaf are not unusual (Molisch 1903). The hydathodes of Araceae are often localized in the leaf tip, the “Vorläuferspitze” (precursor tip), which contains numerous stomata on both surfaces (Müller 1919); hydathodes may also be found elsewhere on leaves. The stomata may be relatively normal in size and appearance, or may be larger than surrounding stomata. The hydathode also contains underlying ground and vascular tissue with numerous tracheids (epithem). However, Gardiner (1883) did not regard the epithem of Calla and Zantedeschia

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THE GENERA OF ARACEAE

as specialized or conspicuous, compared with dicotyledons. The leaf apex and hydathodes have been described in various species of Araceae by Müller (1919), who recognized three types of hydathodes: the Philodendron type, with normal or slightly larger stomata; the Alocasia type, with large stomata, and finally the Colocasia type with gigantic macroscopic stomata through which guttation occurs. The Philodendron type occurs in all subfamilies; the Alocasia type is more restricted, and the Colocasia type was found only in Ariopsis, Colocasia and Steudnera. More complete lists have been given by Müller (1919) and Grayum (1984). In Lasia hydathodes are present on leaf emergences. Further details of hydathode structure have been given by Ducharte (1859), Dalitzsch (1886), Minden (1899) and Gentner (1905).

Intravaginal squamules These are glandular, flattened, non-vascularized, multicellular scales of uncertain function, ranging from approximately 1 mm to over 25 mm in length. They occur in the leaf axils of Acorus (Acoraceae), Philodendron, Cryptocoryne and Lagenandra (Irmisch 1858, Engler 1912, 1920a, b, Lawalrée 1945, Ritterbusch 1971, Blanc 1978). Velenovsky (1907) reported squamules in many Araceae, which must be an error. Similar structures also occur in Spirodela (Lemnaceae) and in all Alismatiflorae except Scheuchzeria, where they are replaced by hairs (Tomlinson 1982). In section, squamules of Araceae and Alismatiflorae often have a mucilaginous and glandular appearance (Kaplan 1973a Tomlinson 1982). Kaplan (1973a) reported that in Acorus, two squamules arise from the protoderm of the axillary meristem and expand rapidly before the prophyll is initiated.

Mineral crystals and crystal idioblasts A wide range of calcium oxalate crystals and crystalcontaining idioblasts is present in Araceae, although few detailed studies have been made of their development, distribution or biological significance (see Seubert 1993 for a recent study in Araceae). Mineral crystals have been viewed both as waste products and as agents of plant protection against herbivores (Esau 1965, Madison 1979a). Calcium oxalate crystals occur most commonly as raphides and druses, less commonly as crystal sand, “ styloids” or prisms Histochemical tests by Sunell & Healey (1981) confirmed that the raphides of Colocasia esculenta contain calcium oxalate. X-ray diffraction studies of raphides of Monstera deliciosa (Al-Rais, Myers & Watson 1971), Xanthosoma sagittifolium (Cody & Horner 1983) and Colocasia esculenta (Sunell & Healey 1981) demonstrated monoclinic calcium oxalate monohydrate. Comparable studies have not been made on the other types of presumed calcium oxalate crystals in Araceae.

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No silica or calcium carbonate crystal inclusions have been demonstrated. Unidentified inclusions (probably not calcium oxalate) are present in the parenchyma of tubers of Amorphophallus konjac (syn. A. rivieri) (Abranowicz 1912). The structure of raphides in Araceae is apparently unusual in angiosperms (Cody & Horner 1983) because they consist of twinned crystals, H-shaped in transverse section and are often barbed laterally (Sakai & Hanson 1974). Similar shaped raphides occur in some Lemnaceae. The systematic significance of these findings is unclear because there have been very few general surveys of raphide structure in angiosperms using the scanning electron microscope, which is necessary for more rigorous comparative studies. Raphide-containing idioblasts are variable in length, wall specialization and manner of raphide release. There are two general types in Araceae: 1) thin-walled raphide idioblasts, and 2) thick-walled idioblasts called biforines. Thin-walled raphide idioblasts range in shape from isodiametric to very elongated (3–5 mm long in Monstera deliciosa, Kovacs & Rakovan 1975). The elongated forms have been referred to as “Raphidenschläuche” (Solereder & Meyer 1928) and may contain many bundles of raphides oriented at various angles and embedded in mucilage. They are of common occurrence in Araceae and have been reported by many authors (e.g. Dalitzsch 1886, Porsch 1911, Solereder & Meyer 1928, Hinchee 1981). The elongated type may intergrade with cells that are much shorter at maturity and tend to grow rapidly by apical intrusive growth, allowing them to exceed the length of surrounding cells. There are reports of raphide idioblasts anastomosing during development in Anthurium scandens, A. scherzerianum and A. triphyllum (Samuels 1923) and in other genera, such as Monstera and Dieffenbachia (Solereder & Meyer 1928). Samuels challenged the idea that the long tubes in many Araceae are single cells, proposing instead that they form by the anastomosis of cells in files. No subsequent authors have addressed this issue and Samuels’ interesting findings are unconfirmed. Raphides are reportedly absent from Acorus (Acoraceae). The second type of raphide idioblast is the biforine, a term coined by Turpin (1836) to refer to spindleshaped or cylindrical cells with thick, unlignified walls (Middendorf 1983). No specific function for these cells has been demonstrated, but some authors have described a “blowgun” release of raphides (Middendorf 1983). One or both ends of the cell may be blunt, pointed or papillose, and with much thinner walls. The thinnest part of the end wall in Colocasia esculenta is 0.1 µm thick (Sakai & Hanson 1974). The variable orientation of fibrillar material causes the cell wall to appear layered, except in Alocasia (Sakai & Hanson 1974). The dimensions of some biforine cells are 50 x 150 µm in Colocasia esculenta (Sakai & Hanson 1974) and 40 x 90 µm in Alocasia. Biforines like those of Colocasia esculenta may have one end embedded in tissue and the rest of the cell projecting into an intercellular space.

Alternatively both ends may project into separate spaces with the middle of the cell loosely held by mesophyll cells or vascular tissue (Sunell & Healey 1981). The presence of mucilage in biforines has been recognized since Turpin’s study (1836) and its swelling is generally considered to be the motive force for the expulsion of the contents of the biforine (Sakai & Hanson 1974). In some species raphides are expelled individually (Colocasia) by piercing and subsequent rupturing of the thin papillae. In others expulsion occurs after the papillae are ruptured by other means (Sakai & Hanson 1974), as in Dieffenbachia. By contrast, in Alocasia the cell wall of the biforine breaks at the base of the cell and the entire mass of crystals and mucilage is ejected, followed by swelling and individual dispersal of the raphide crystals. The occurrence of biforines in Araceae has been summarized by Solereder & Meyer (1928), by Nicolson (1959) who added many new observations, and by Grayum (1984). No detailed family-wide survey has yet been made, but existing data (De Bary 1884, Dalitzch 1886, Solereder & Meyer 1928, Grayum 1984) suggest that biforines are characteristic of genera with unisexual flowers and generally absent in bisexually-flowered genera. Among the latter, biforines have so far been observed only in Anthurium, Orontium and Symplocarpus. Calcium oxalate is also the major component of druses, which have a wide systematic occurrence in the family. Druses vary considerably in size, sometimes reaching 90 µm in Zamioculcas (Solereder & Meyer 1928). They also vary in morphology and include atypical forms in leaves of Anthurium, Dieffenbachia, Pistia and other genera (Solereder & Meyer 1928) and may occur in small rounded epidermal cells in Anthurium, Culcasia, Pothos, Schismatoglottis and many other genera. Prisms are of limited occurrence in Araceae, and have been reported only in subfamilies Pothoideae and Monsteroideae (Solereder & Meyer 1928, Nicolson 1959, Seubert 1993). Developmental studies of raphide crystals and crystal-containing idioblasts have been made by Samuels (1923), Becker & Ziegenspeck (1931), Rambour (1965), Mollenhauer & Larson (1966), Rakovan, Kovacs & Szujko-Lacza (1973), Kovacs & Rakovan (1975), and Hinchee (1981). According to Kovacs & Rakovan (1975), raphide idioblasts in Monstera arise by unequal cell division. The smaller cell develops into the raphide idioblast which has densely staining cytoplasm and a larger nucleus than the surrounding cells. Hinchee (1981), however, did not observe asymmetric divisions giving rise to raphide idioblasts in Monstera. Kovacs & Rakovan (1975) noted that raphides develop as soon as the young idioblast cells are the same size as surrounding cells, before the phase of rapid elongation. Seubert (1993) has recently carried out a survey of crystal types occurring in the seeds of Araceae as part of a comprehensive study of Araceae seed anatomy. Her illustrations reveal a fascinating diversity of novel crystal forms.

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Occurrence of vessels Vessels with scalariform or reticulate perforation plates have long been recognized in the roots of some Araceae, such as Monstera deliciosa, and Philodendron and Alocasia species (Solereder & Meyer 1928, Kundu 1942), while other genera like Caladium and Pistia have been considered vesselless (Solereder & Meyer 1928). Solereder & Meyer were apparently the first to identify vessels in the stems of Araceae, in Pothos scandens and P. rumphii. In Culcasia scandens they found no vessels but tracheids with wide lumens. Later Cheadle (1942) examined 7 genera and 8 species and found vessels in the roots only, with scalariform perforation plates in all species. Subsequently Hotta (1971) reported vessels in stems of a few species, but in the roots of nearly all those he examined except Arisaema, Homalomena, Pothos and Rhaphidophora. Hotta’s sample was not very broad taxonomically, but represents the most extensive to date. Vessels occur in stems of several species of Pothos, Epipremnum, Rhaphidophora and Scindapsus. Hotta (1971) emphasized the difficulty encountered in recognizing vessel members in Araceae, principally because of the similarity between scalariform pitting and scalariform perforations on the long, oblique end walls of tracheids. He also referred to some tracheids as “vesselform” tracheids because of their similarity to vessels.

Phloem cytology The phloem of Araceae has been examined using light microscopy in studies by Shah & James (1971) as well as in investigations by Lesage (1891). Recent ultrastructural studies have been made by Behnke (1969, 1981), Bonzi & Fabbri (1975, 1978) and

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THE GENERA OF ARACEAE

Parthasarathy (1980). Behnke’s work on sieve element plastids is of considerable interest. In an extensive survey of the Araceae, Behnke (1995) confirmed the widespread occurrence of the typically monocot plastid type P2 in the family. The Araceae have a relatively rich array of sieve element plastid ultrastructure, with type P2c found in 14 species and P2cfs in one species. A P2 subtype with both cuneate protein and starch grains was found in 110 of the 126 species surveyed. Unexpectedly Pistia was found to have S type plastids thus completely different from all other monocotyledons. The S type has heretofore been found only in dicotyledons. Conversely only two dicotyledons have been found with P2c plastids, Asarum and Saruma. The phylogenetic significance of these findings is discussed at length by Behnke (1995).

Starch grains Starch grain morphology in Araceae has been examined by Reichert (1913), Czaja (1969, 1978a, b) and Seubert (1993). Reichert reported that starch grains in Arum, Arisaema, Dracunculus and Zantedeschia all belonged to the same type, while in Dieffenbachia a remarkably different kind is present. Despite gross structural differences all five genera have starch with similar physical and chemical properties. Reichert (1913) also studied starch grains in Alocasia, Amorphophallus, Peltandra and Colocasia. Those of Peltandra were found to resemble the Dieffenbachia type, while grains in Amorphophallus and Colocasia were of the Arum type. Grains of Alocasia were “peculiar” and not assigned to either type. More recently, Seubert (1993) has surveyed starch grain morphology in Araceae seeds. Using Czaja’s terminology (Czaja 1969), she recognized 11 different types in a survey of over 70 genera.

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4

INFLORESCENCE AND FLORAL MORPHOLOGY

The inflorescence of the Araceae is composed of an unbranched spike bearing flowers, the spadix, subtended by a bract called the spathe. The flowers are usually numerous, very small, sessile in all genera except Pedicellarum, and lack floral bracts. They are generally spirally arranged and usually tightly packed, although in some species of Pothos ser. Goniuri, Pedicellarum, Amorphophallus (male and female flowers), tribe Dieffenbachieae (female flowers) and most species of Arisaema and Arisarum (male flowers), they may be somewhat distant from one another. The spathe is, strictly speaking, the last leaf of a flowering article. It is usually a specialized attractive organ although in a few genera (Gymnostachys, Orontium) is inconspicuous. The internode between spathe and spadix (spadix stipe) is usually very short or absent, while the peduncle – the internode between spathe and last foliage leaf or cataphyll – is much longer. In some primitive genera, however, this arrangement is reversed (Gymnostachys, subfamily Orontioideae, some Pothos species). It seems likely that an important stage in aroid evolution involved the combined development of a relatively long peduncle and

short stipe, modified attractive spathe and continuation shoot development at the second node below the spathe. Gymnostachys and subfamilies Orontioideae and Pothoideae seem to represent earlier, less uniform phases of organization. The typical araceous pattern has given rise to a wide range of variant forms in different genera, which can be seen to represent an evolutionary trend of increasing integration towards a synflorescence or pseudanth. The major phyletic modifications are: 1) loss of perigone in the flowers; 2) specialization of flowers on the spadix into a lower female zone, upper male zone and, often, one or several zones of sterile flowers, entirely naked axial zones and smooth or staminodial terminal appendices; 3) differentiation of the spathe into a lower, convolute tube and an upper, expanded blade. Spathe and spadix modifications are closely related so that the spathe may be seen evolutionarily as becoming increasingly integrated into the inflorescence itself, until in extreme cases, such as tribe Cryptocoryneae, Ambrosina, Pistia and Pinellia, fusion and still more elaborate modifications have brought about division of the spathe into separate chambers.

Spathe blade Spadix

Male zone

Flowers Sterile zone Stipe Spathe constriction Spathe Female zone Spathe tube Stipe Peduncle Peduncle

Figure 7. Inflorescence types: A, bisexual flowered spadix with a simple, undifferentiated spathe; B, unisexual flowered spadix with a spathe divided into a limb (blade) and convolute lower tube.

INFLORESCENCE AND FLORAL MORPHOLOGY

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Other notable specializations of the inflorescence include the wide range of odours found in different genera, colour patterns, especially on the spathe, and the relative persistence of different regions of the spathe. In Philodendron, for example, the entire spathe persists until fruit, but in tribes Colocasieae, Caladieae, Peltandreae and Schismatoglottideae the spathe blade withers or drops off immediately following anthesis while the spathe tube persists. In many Monstereae the entire spathe withers or drops off soon after flowering, a behaviour which is correlated in this tribe with the presence of numerous protective trichosclereids in the style tissue. Terminal appendices of the spadix are found in tribes Areae, Arisaemateae, Colocasieae, Schismatoglottideae, Thomsonieae and Zomicarpeae, sporadically elsewhere in the family. The function of the appendix, where investigated, is to produce odours to attract pollinators (osmophore, Vogel 1963, 1990). The appendix is either clearly composed of staminodes (e.g. Pseudodracontium) or is partially to entirely smooth with no vestiges of floral organs (e.g. Arum). Flowers in Araceae may be 2- or 3-merous. In perigoniate flowers the tepals, when free, are organized in two whorls. The tepals are usually more-or-less fleshy and fornicate apically (except subfamily Pothoideae) and in some genera or sections (Anadendrum, Holochlamys, Pedicellarum, Spathiphyllum sect. Massowia, Stylochaeton) they are fused into a cup-like structure. Stamens in perigoniate flowers and in the naked bisexual flowers of most Monsteroideae have essentially the orthodox structure of distinct (usually flattened) filament, basifixed anther and slender, inconspicuous connective. In the unisexual flowers of many tribes of subfamily Aroideae, however, filaments are typically very short or lacking, and there is a thick, fleshy connective which probably acts as an osmophore (Aglaonemateae, Culcasieae, Homalomeneae, Montrichardieae, Nephthytideae, Philodendreae, Zantedeschieae). Stamens of tribes Anubiadeae, Caladieae, Colocasieae, Dieffenbachieae, and Peltandreae are essentially similar but are always fused into synandria. In tribe Arophyteae the stamens may be fused or not and exhibit a diversity of structure. Large connectives also occur in tribe Spathicarpeae but their different morphology suggests that they are not homologous with those of the other

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THE GENERA OF ARACEAE

tribes of subfamily Aroideae mentioned above. Anthers are almost always extrorse (introrse in Zamioculcas, latrorse in Pedicellarum). Theca dehiscence may be by a longitudinal or rarely transversal slit (most genera with bisexual flowers and some unisexual-flowered genera: Anubias, some Areae, Arisaema, Arisarum, Stylochaeton) or by apical or subapical pores or short slits. In many genera of subfamily Aroideae dehiscence of each theca is by a subapical stomial pore and this morphology is frequently correlated with the extrusion of pollen in strands. Similar structures occur in Amorphophallus and Dracunculus. The gynoecium usually varies between 1- and 3locular, and when unilocular often shows traces of 2or 3-merous origin through the presence of a severallobed stigma (e.g. Typhonodorum) or more than one placenta (e.g. Schismatoglottis). Gynoecia with more than 3 locules are less common but are found in tribe Spathicarpeae (1–8 locular) and in Philodendron (2–47 locular). Placentation varies from axile to parietal, basal, apical or basal and apical (the latter in Dracunculus, Helicodiceros, Heteroaridarum and Theriophonum), with many intermediates. Ovules may be anatropous, campylotropous, orthotropous or intermediate between these types. Funicle trichomes are usually present (French 1987c) and secrete a clear, mucilaginous substance which in many genera (e.g. tribe Monstereae, Philodendron) entirely fills the ovary locules; this secretion appears to play a role in pollen tube growth (Buzgó 1994). The style may be narrowed and elongated (e.g. Dracontium) but in most genera is relatively inconspicuous externally. However, there is very often a thick stylar region between the ovary locules and stigma (e.g. Philodendron, Mayo 1989b). In tribe Monstereae this stylar region is especially well developed and densely filled with trichosclereids. Here the style seems to substitute functionally for a perianth in protecting the sexual organs of the flower. Stigmas are always wet in Araceae and in some genera (Anthurium, Arum, several Lasioideae) produce conspicuous nectar droplets at anthesis. In Amorphophallus, Dieffenbachia and some Spathicarpeae, the lobing of the stigma can be very pronounced, or the stigma relatively massive. In subfamily Monsteroideae stigmas vary from subcapitate to conspicuously elongated, either transversely (e.g. Anadendrum) or longitudinally.

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5

I N F L O R E S C E N C E A N D F L O R A L A N ATO M Y

Inflorescence and flower anatomy of Araceae received little attention after the completion of Engler’s last monograph (Engler 1920b) until very recently. Knoll (1926) made a detailed study of inflorescence anatomy in Arum with particular reference to structures and adaptations concerned with pollinator behaviour. Pohl (1932a, b) and Mayo (1986b, 1989b) studied inflorescence and floral anatomy in Philodendron and demonstrated the existence of spathe tissues adapted for opening and closing movements, resin secretion of various types from resin canals in the spathe and spadix and a wide variety of gynoecial and androecial structures. Eyde, Nicolson & Sherwin (1967) made the first general survey with a study of the flowers of 18 genera (including Acorus), which concentrated more on those with bisexual or bisexual-tepalate flowers. They established that in floral anatomy, as in other characters (Grayum 1987), Acorus differs markedly from the Araceae and confirmed the absence of floral bracts subtending the flowers. Barabé and coworkers (Barabé1982, 1987; Barabé & Chrétien 1985, 1986a, b; Barabé, Chrétien & Forget 1986; Barabé & Forget 1987, 1988a, b, 1993; Barabé, Forget & Chrétien 1986, 1987; Barabé & Labrecque 1983, 1984, 1985; Barabé, Labrecque & Chrétien 1984) have made a series of detailed floral anatomical studies of different genera with particular

emphasis on floral vasculature. One aim of their studies has been to establish whether unilocular gynoecia in Araceae are always pseudomonomerous (see also Eckardt 1937) or if truly 1-carpellate gynoecia occur; thus far no examples of the latter have emerged. French (1985a,b, 1986 a,b, c, 1987c) has made familywide surveys of endothecial thickenings, stamen and ovule vasculature and ovular trichomes. These have further confirmed the distinctness of Acorus and revealed some taxonomically useful character variation within the family. Carvell (1989a, b) has made a detailed survey of floral anatomy in the bisexual and bisexual-tepalate flowered genera. Vogel (1963, 1978) made many fascinating observations, including histological studies, of the terminal appendix in various genera (Alocasia, Arisaema, Arum and others) and the spathe limb of Cryptocoryne, in relation to their biological function as osmophores. Eyde et al. (1967), Vogel (1963, 1990), Mayo (1986b, 1989b) and Chauhan (unpublished results) demonstrated the existence of papillate and sculptured cell surfaces in the epidermis of the spathe, androecia and gynoecia of Amorphophallus, Homalomena, and Philodendron species; this feature is probably very widespread in the family. Ittenbach (1993) has made a very interesting study of the anatomy and micromorphology of some African Amorphophallus species.

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6

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FRUITS AND SEEDS

The fruits of Araceae are typically juicy berries, although rarely drier and leathery. The infructescence is usually cylindric or sometimes globose. The berries are most commonly red or orange (see family description) and are almost always free. Exceptions are Syngonium, in which the berries form an indehiscent syncarp, and Cryptocoryne which has an apically dehiscent syncarp. In Lagenandra the berry actively opens at the base to release the seeds, but aroid berries are otherwise indehiscent. The various mechanisms observed for protection of the developing fruits and seeds have been discussed by Madison (1979a). In the Monstereae, which have bisexual but non-perigoniate flowers, the thick stylar region is filled with trichosclereids which protect the developing seeds. At maturity the stylar region is shed to reveal the seeds. In perigoniate genera like Anthurium the perigone clearly plays a protective role and keeps pace during growth of the developing berry. The latter only becomes fully exposed at maturity by extrusion from the flower. In Lysichiton, also perigoniate, the stylar region and tepal apices protect the young berry, eventually breaking off to reveal the ripe seeds (Hultén & St. John 1931). In many unisexual-flowered genera the protective function is assumed by the persistent spathe or spathe tube. Spathe growth continues around the developing fruits until maturity when the spathe may split open (Alocasia, Dieffenbachia) or absciss at the base (Philodendron), exposing the infructescence of white or coloured berries. In other monoecious genera, however, the spathe is marcescent and plays no role in fruit protection. In such cases (e.g. Arum) protection may possibly be through the presence of toxic chemical compounds in the berries. The seeds are often embedded in mucilaginous pulp (secreted by the ovular and placental trichomes). In Anthurium the inner layer of the pericarp may also be mucilaginous and in other genera the outer integument becomes mucilaginous. This makes the seeds sticky and aids the dispersal of epiphytic and

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THE GENERA OF ARACEAE

hemiepiphytic species to new sites (e.g. neighbouring trees) by birds or mammals. The amount of endosperm in the seed varies considerably within the family and has long been regarded as a useful taxonomic character at tribal level. However, a thorough comparative anatomical study has been lacking until very recently (Seubert 1993). Among a wealth of other interesting new observations, Seubert’s study shows that endosperm may be present in mature seeds of some groups e.g. subfamily Lasioideae, as only a very thin layer. Many intermediate conditions exist between the presence of copious endosperm and absence of endosperm. Absence of endosperm is often correlated with the presence of a well-developed plumule and the largest seeds of Araceae are of this type, e.g. Orontium, Typhonodorum. The seeds are usually straight, but in Lasioideae and Monstereae they are often curved, sometimes strongly so. In a few genera the plumule is also highly developed (Cryptocoryne, Gonatopus, Nephthytis, Orontium, Typhonodorum). In Cryptocoryne ciliata, which grows in freshwater tidal zones, between 20 and 40 cataphylls are formed in the embryo, and these may serve to fix the seed to the substrate, preventing it from being swept away with the ebb and flow of tides. Seeds with a well developed plumule usually lack endosperm and have only a very thin, papery testa or none at all (Nephthytis). Such large embryos generally contain chlorophyll at maturity and are only viable for a short time. The testa may be smooth, rough, verrucose or costate, thin or thick and in subfamily Lasioideae is often very hard and thick with prominent sculpturing. In tribes Ambrosineae, Areae, Arisaemateae and Arisareae, most genera have a prominent fleshy strophiole (aril), and in tribe Colocasieae smaller but distinct strophioles also occur. Arillate seeds have been observed in Philodendron subgen. Meconostigma (Mayo 1986b, 1991). In Zomicarpa the swollen funicle remains connected to the mature seed (Peyritsch 1879, Bogner, pers. obs.). In Pistia a double operculum is formed by both integuments.

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7

SEEDLING MORPHOLOGY

Four different types of seedling can be recognized in the Araceae (Tillich 1985, 1995, Seubert 1993). In Type 1, the seeds have copious endosperm, the cotyledonar hyperphyll functions as a haustorium, the cotyledonar sheath is condensed and short, the hypocotyl and primary root are well developed and first leaf is either a cataphyll or a foliage leaf. This type is quite common in the family, e.g. Arisaema, Arum, Calla, Gymnostachys, Pinellia, Zantedeschia. Type 2 is similar to Type 1, but the cotyledonar sheath is broadened and blade-like, green and assimilatory, whereas the cotyledonar hyperphyll is a minute haustorium, e.g. Colocasia, Philodendron, Xanthosoma. Type 3, which occurs only in Pistia and is very similar to the seedling morphology of Lemna (Lemnaceae), differs from Types 1 and 2 in that the primary root and hypocotyl are undeveloped. In Type 4 the seeds have little or no endosperm.

The cotyledonar hyperphyll functions as a storage organ and perhaps has a very limited haustorial role when some endosperm is present. The cotyledonar sheath, hypocotyl and primary root tend to be reduced or often completely absent. In Acorus (Acoraceae) the seedling bears no similarity to any form found in the Araceae. The cotyledonar hyperphyll is subdivided into a long, cylindrical, assimilatory portion and a small haustorial tip. The first plumular leaves are unifacial and ensiform and the primary root is well developed. Tillich (1985, 1995) has pointed out the resemblance between the seedling of Acorus and those of the Juncaceae, Melanthiaceae, and Typhaceae. In Type 2 and in Acorus, the root collar bears conspicuous, long and densely disposed rhizoids. In Type 1 rhizoids are found only sporadically (e.g. in Cryptocoryne cognata but not in C. ciliata).

SEEDLING MORPHOLOGY

29

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8

C

E M B RYO L O G Y

The embryology of Araceae has not been studied on a broad comparative scale, but excellent reviews have been published by Grayum (1984, 1991b), on which this chapter is based. Thorough treatments of all embryological aspects of a single taxon have been published for Peltandra virginica (Goldberg 1941), Theriophonum minutum (Parameswaran 1959) and Synandrospadix vermitoxicus (Cocucci 1966); other genera are known less completely. Jüssen (1928) made the most important single contribution to date, but generalizations about the family’s embryology are still based on rather fragmentary coverage. Acorus (Acoraceae) has a secretory anther tapetum, thus differing from the Araceae, which have the periplasmodial type (sensu Clausen 1927). Pollen mother cell division is probably always of the successive type in Araceae. The nucellar epidermis of virtually all Araceae divides to form a nucellar cap, usually 2–3 cell layers thick (1 in Pistia), and for this reason Araceae have been treated as

30

THE GENERA OF ARACEAE

crassinucellate. However, in a strict sense most Araceae so far investigated have tenuinucellate ovules, except Symplocarpus and Calla. An endothelium, derived from the inner surface of the inner integument, is reported from most genera investigated. The seed of Acorus has a perisperm derived from the nucellus in addition to the endosperm, and in this differs from all Araceae. Linear megaspore tetrads are the commonest type, but T-shaped tetrads are also found, sometimes both types occurring in the same species. The mature embryo sac is usually of the 8-nucleate type (10–12 nucleate in Nephthytis). Embryogeny is clearly understood in only nine genera. Onagrad and asterad embryogeny are known in monoecious genera and caryophyllad and solanad types are known in bisexual genera. Cellular and free-nuclear types of endosperm development both occur but after a review of the available facts, Grayum (1984, 1991b) concluded that endosperm development in Araceae is best interpreted as a form of the helobial type.

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C

9

C Y TO L O G Y

This chapter is based on Petersen’s (1989, 1993) recent comprehensive review of aroid cytology. The 2n chromosome numbers given in Table 1 and in the generic descriptions are also from her results, supplemented by unpublished data from Dr. Marcelo Guerra (pers. comm.). Reliable diploid (2n) numbers and intraspecific aneuploid derivatives are given, with dubious numbers placed in brackets. The chromosome numbers in Araceae vary greatly between genera, from 2n=14 (Ulearum) to 2n=168 (Arisaema). Within a single genus the diploid number may be highly variable, as in Cryptocoryne (2n=20 to 2n=132), or in Arisaema (2n=20 to 2n=168). Some genera, on the other hand, have very stable diploid numbers. Anthurium, the largest genus of the family, is surprisingly uniform cytologically, with the great majority of species having a diploid number of 2n=30. The distribution and variation of chromosome numbers among the genera suggests that chromosome number has increased in some phyletic lines to a high level of polyploidy and in others has been greatly reduced. The most advanced tribes, such as Areae, Arisaemateae and Cryptocoryneae, tend to have the highest numbers. In the more primitive genera chromosome number tends to be both less variable and less extreme, being neither very high nor very low. Examples of this type are Pothos (2n=24, 36) and Spathiphyllum (2n=30, 60). Petersen (1989) considered that no modern Araceae species constitutes a true primary diploid. In the morphologically most primitive genera the lowest diploid number is 2n=24. In contrast, the highly derived Ulearum sagittatum has the lowest diploid number known in the family (2n=14), evidently the result of phyletic reduction. A primary basic number of x=7 has been proposed by Jones (1957), Larsen (1969) and Marchant (1973).

Petersen (1989) considered that the basic numbers x=14 or x=12 must have been the starting points for the derivation of all the modern chromosome numbers in the Araceae. It is possible, however, that these basic numbers may represent secondary basic numbers which arose by chromosome doubling from hypothetical basic numbers x=7 or x=6. Petersen regarded the widely occurring number x=14 as of ancient origin. She also considered that the number x=12 might be primitive since the diploid number 2n=24 occurs in tribe Potheae. Reduction in diploid number occurs in various morphologically advanced genera. A good example is tribe Areae in which the commonest diploid numbers are 2n=26 and 2n=28 but the genera Biarum and Typhonium both include species with diploid numbers as low as 2n=16. Aneuploid changes at the diploid level followed by polyploidy or aneuploidy at the polyploid level have taken place in some taxa, e.g. Cryptocoryne ciliata (2n=22, 33), Cryptocoryne cordata (2n=34, 68, 85, 102). The size and shape of the chromosomes are also quite variable. Chromosome length varies from 1–17 µm, depending on the genus. The chromosomes have been observed to differ greatly in size and shape within a single genome in certain cases, e.g. the New and Old World species of Homalomena. Homalomena speariae (New World) has 42 chromosomes, of which 40 are small (1–2 µm) and one pair is nearly three times longer (ca. 5 µm). In Old World species there is no large pair of chromosomes. Chauhan and Brandham (1985), in a study of Amorphophallus cytology, showed some appreciable size differences within the karyotypes of certain species. Ramalho (1994) has made a recent study of Araceae chromosomes in Pernambuco, Brazil.

C Y TO L O G Y

31

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Table 1. List of diploid (2n) and assumed basic (x) chromosome numbers in the genera of Araceae.

Chromosome numbers arranged in euploid series are separated by a semicolon; dubious numbers are in brackets (data from G. Petersen, pers. comm., updated from Petersen 1989 and Guerra, pers. comm.). 2n

x

22, 44; 24, 36, 48

11, 12

I. Subfamily Gymnostachydoideae 1. Gymnostachys

48

12

II. Subfamily Orontioideae 2. Orontium 3. Lysichiton 4. Symplocarpus

26 (24, 28) 28 30, 60 (28)

13 14 15

24, 36 no data 24

12

20, 40; 24, 48, 84; 28, 56; 30, 60, 90

10, 12, 14, 15

30, 60 60

15 15

60

15

28

14

60 60, 120 (42, 54, 56) 60 (56, 84) 60 (42, 56, 58, 64, 70, 112) 60 (24, 48, 56, 58, 70) 84 28, 56 28

15 15 15 15 15 14 14 14

26 26 26 26 26 26 26 26 26 52

13 13 13 13 13 13 13 13 13 13

Family Acoraceae 1. Acorus Family Araceae

III. Subfamily Pothoideae Tribe Potheae 5. Pothos 6. Pedicellarum 7. Pothoidium Tribe Anthurieae 8. Anthurium IV. Subfamily Monsteroideae Tribe Spathiphylleae 9. Spathiphyllum 10. Holochlamys Tribe Anadendreae 11. Anadendrum Tribe Heteropsideae 12. Heteropsis Tribe Monstereae 13. Amydrium 14. Rhaphidophora 15. Epipremnum 16. Scindapsus 17. Monstera 18. Alloschemone 19. Rhodospatha 20. Stenospermation V. Subfamily Lasioideae 21. Dracontium 22. Dracontioides 23. Anaphyllopsis 24. Pycnospatha 25. Anaphyllum 26. Cyrtosperma 27. Lasimorpha 28. Podolasia 29. Lasia 30. Urospatha

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THE GENERA OF ARACEAE

12

1-22 Section A Acro 18/7/97 8:02 Page 33

VI. Subfamily Calloideae 31. Calla VII. Subfamily Aroideae Tribe Zamioculcadeae 32. Zamioculcas 33. Gonatopus Tribe Stylochaetoneae 34. Stylochaeton Tribe Dieffenbachieae 35. Dieffenbachia 36. Bognera Tribe Spathicarpeae 37. Mangonia 38. Taccarum 39. Asterostigma 40. Gorgonidium 41. Synandrospadix 42. Gearum 43. Spathantheum 44. Spathicarpa Tribe Philodendreae 45. Philodendron Tribe Homalomeneae 46. Furtadoa 47. Homalomena Tribe Anubiadeae 48. Anubias Tribe Schismatoglottideae 49. Schismatoglottis 50. Piptospatha 51. Hottarum 52. Bucephalandra 53. Phymatarum 54. Aridarum 55. Heteroaridarum Tribe Cryptocoryneae 56. Lagenandra 57. Cryptocoryne Tribe Zomicarpeae 58. Zomicarpa 59. Zomicarpella 60. Ulearum 61. Filarum Tribe Caladieae 62. Scaphispatha 63. Caladium 64. Jasarum 65. Xanthosoma 66. Chlorospatha 67. Syngonium 68. Hapaline Tribe Nephthytideae 69. Nephthytis 70. Anchomanes 71. Pseudohydrosme

36, 54, 72

18

34 34, 68

17 17

28, 56

14

34, 68 34

17 17

no data 34 34 34 34 no data 34 34

17 17 17 17 17 17

28; 30; 32; 34; 36; 48 (26)

15, 16, 17, 18

40 38; 40, 80; 42

20 19, 20, 21

48, 72

24

26, 39, 52 26 26 c. 26 26 24 no data

13 13 13 c. 13 13 12

36, 72 20; 22, 33, 66, 88, 132; 28, 42; 30; 34, 68, 85, 102; 36, 54, 72, 90

18 10, 11, 14, 15, 17, 18

20 26 14 28

10 13 7 7

28 22; 26; 28; 30; 32 22 22; 26, 39, 52 26 28 (24, 26) 26; 28

14 13, 14, 15, 16 11 11, 13 13 14 13, 14

36; 40, 60 40 c. 40

18, 20 20 c. 20

C Y TO L O G Y

33

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Tribe Aglaonemateae 72. Aglaonema 73. Aglaodorum Tribe Culcasieae 74. Culcasia 75. Cercestis Tribe Montrichardieae 76. Montrichardia Tribe Zantedeschieae 77. Zantedeschia Tribe Callopsideae 78. Callopsis Tribe Thomsonieae 79. Amorphophallus 80. Pseudodracontium Tribe Arophyteae 81. Arophyton 82. Carlephyton 83. Colletogyne Tribe Peltandreae 84. Peltandra 85. Typhonodorum Tribe Arisareae 86. Arisarum Tribe Ambrosineae 87. Ambrosina Tribe Areae 88. Arum 89. Eminium 90. Dracunculus 91. Helicodiceros 92. Theriophonum 93. Typhonium 94. Sauromatum 95. Lazarum 96. Biarum Tribe Arisaemateae 97. Pinellia 98. Arisaema Tribe Colocasieae 99. Ariopsis 100. Protarum 101. Steudnera 102. Remusatia 103. Colocasia 104. Alocasia Tribe Pistieae 105. Pistia

40, 60, 80, 100, 120 (70, 110) 40

20 20

42, 84 42

21 21

48

24

32

16

36

18

26, 39; 28 26

13, 14 13

38, 76; 54 (40) 54, 108 54

19, 27 27 27

56, 112 112

14 14

28, 42, 56

14

22

11

28, 42, 56, 70, 84 24; 28 28 56 16 (14, 18) 16; 18, 36, 54; 20; 26, 52, 65; >100 (14) 26, 52, 104 c. 78 16; 20; 22; 24; 26; 32; 36; 74; 96; 98*

14 14 14 14 8 8, 9, 10, 13 13 c. 13 ?

26, 52 20; 22, 24, 48, 72; 26, 39, 52; 28, 42, 56, 70, 112, 140, 168 (64)

13

28, 28 28, 28, 28, 28,

14 14 14 14 14 14

28

84 (80, 86) 42, 42, 42, 42,

56 56 56 56, 70, 84

10, 11, 12, 13, 14

14

* No attempt has been made in this case to arrange the numbers into euploid series in this genus.

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THE GENERA OF ARACEAE

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10 PA L Y N O L O G Y

Thanikaimoni (1969) and Grayum (1984, 1985, 1992a) have given detailed comparative surveys of aroid pollen structure. These studies showed that palynological characters are important for the suprageneric taxonomy. Aperture type is the feature of most general value. Bisexual-flowered genera have monosulcate, extensive-sulcate, meridionosulcate (zonate), diaperturate or forate grains, but monoecious genera have inaperturate grains. The only exceptions seem to be tribe Spathiphylleae (bisexual, inaperturate), Anadendrum (bisexual, inaperturate), and tribe Zamioculcadeae (monoecious, extended-monosulcate to zonate). Ornamentation may be smooth (psilate), scabrate, foveolate, reticulate, spinulose, spinulose-reticulate, spinulose-pilate to papillate, spinose, fossulate, rarely gemmate, verrucate, retiverrucate, areolate, rugulate to tuberculate, striate, striate-verrucate, striate-reticulate, striate to plicate or baculate. Spiny pollen grains are common in Araceae, and have been considered (Grayum 1984) as an adaptation for aiding attachment to insect vectors which have hairy bodies. In many other genera, smooth pollen grains are extruded in strands composed of many grains glued together by pollenkitt. These strands also adhere to insect bodies, sometimes through the aid of sticky secretions within the inflorescence (e.g. Philodendron). On the basis of existing observations, smooth pollen is almost always

associated with beetle pollination and spinose pollen with fly pollination (Grayum 1984, 1985, 1990). Beetles (Phaeochrous camerunensis, Scarabaeidae-Rutelinae) and blowflies (Calliphoridae) were both observed as pollinators in Amorphophallus maculatus which has almost smooth pollen grains (Bogner 1976a). The shape is globose to ellipsoid, boat-shaped or hamburger-shaped (the latter in Gonatopus). The polarity is heteropolar to isopolar or apolar. Usually the pollen grains are found in monads, only two genera (Xanthosoma, Chlorospatha) shed the pollen grains in tetrads, arranged either tetragonally or serially (Chlorospatha longipoda). Grain size (measurements given here and in the generic descriptions are mostly taken from Grayum, 1984, 1992a) varies considerably, from small (12 µm in Homalomena versteegii) to very large (114 (120) µm in Pseudohydrosme gabunensis) but the majority of the genera (68%) are medium-sized (between 25–50 µm diam., mean 37 µm). In 73% of species examined the pollen contains starch although this may vary within a single genus; in Schismatoglottis some species have starchy pollen and others do not. Nine out of ten genera with monosulcate pollen grains were found to be starchless which suggests that this is the primitive type in Araceae.

PALYNOLOGY

35

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11 P H Y TO C H E M I S T RY A N D C H E M OTA X O N O M Y

C

by Robert Hegnauer

Several recent reviews have dealt with chemical aspects of the biology of Araceae (Hegnauer 1963, 1986; Dahlgren & Clifford 1982; Dahlgren et al. 1985; Bown 1988). Many references to the chemical characters and the ethnobotany of the family are available in the two treatments by Hegnauer and in chapters 9 and 10 of Bown’s book. The present review supplies only the most essential bibliography, and for additional references the reader should refer to these three sources.

1. Phytochemistry

Mineral deposits and primary metabolites Oxalic Acid Aroids produce large amounts of oxalic acid, most of it being deposited as crystals of calcium oxalate. Raphide bundles (see also chapter 3), i.e. agglomerations of large, needle-like crystals lying parallel to one another, are the typical crystal form of the family. These raphide bundles usually occur singly, embedded in mucilage within large idioblasts (see section on Irritants in Araceae). Other types of calcium oxalate crystals are found in aroids, such as druses, and in Acorus (Acoraceae), a genus lacking raphides, cells containing a solitary crystal are situated in rows accompanying fibres. For more detailed studies of calcium oxalate crystals in Araceae, see Seubert (1993). According to Molisch (1918), who investigated Amorphophallus rivieri (= A. konjac), Caladium nymphaeifolium (probably a variety of Colocasia esculenta), Monstera deliciosa and Sauromatum guttatum (= S. venosum), aroids also tend to accumulate moderate to large amounts of soluble oxalates in leaves. In this respect they are similar to Lemnaceae, Helobiae (= Alismatiflorae) such as Stratiotes aloides and Vallisneria spiralis, and Zingiberales (Canna, Musa, Maranta). Silica, aluminium and heavy metals are not known to be accumulated significantly by members of the family. Tubers of Eminium spiculatum and Arisarum vulgare were found not to contain soluble oxalic acid; tartaric and citric acid were detected in both species (Ahmed et al. 1968).

Carbohydrates The carbohydrates stored by Araceae have been investigated many times. Pollard (1982) studied the

36

THE GENERA OF ARACEAE

distribution of kestose- and isokestose-types of sucrose fructosides in the basal parts of fresh monocotyledonous stems; no such oligofructans (oligofructosans) were present in Acoraceae (Acorus gramineus) or Araceae (Arisaema atrorubens (= A. triphyllum), Dieffenbachia picta (= D. maculata) and Peltandra virginica were investigated), Lemnaceae, Alismataceae, Dioscoreaceae, Sparganiaceae and Arecaceae (see e.g. Dahlgren et al. 1985: 277). Sakai & Hayashi (1973) studied the distribution of starchy and sugary leaves in monocots. Japanese Acoraceae and Araceae belong to those taxa which temporarily store sugars and non-starchy polysaccharides, but little if any starch, in the leaves. Acorus gramineus, Amorphophallus konjac, eight species of Arisaema, Calla palustris, Colocasia esculenta, Lysichiton camtschatcensis and Pinellia ternata were investigated. Starch was seen occasionally only in Acorus gramineus and in two species of Arisaema. Araceae, which have “sugary leaves” thus differ from the starchy leaved families Alismataceae, Dioscoreaceae and Tricyrtidaceae, but resemble most Japanese members of the Liliiflorae (sensu Dahlgren & Clifford 1982).

Mucilages Vegetative parts of the Araceae mainly store starch (Czaja 1969, 1978a, b; for economically important Araceae see also Brücher 1977, Mansfeld 1986, Palmer 1989, and for taro – Colocasia esculenta – p. 54 and table 4.28 in Sunell & Arditti 1983). In some taxa starch is accompanied by mucilages in considerable amounts, which usually consist mainly of glucomannans (Hegnauer 1963, 1986). Ohtsuki (1967) showed that subterranean parts of most Acoraceae and Araceae contain much starch but only negligible amounts of glucomannans (Acorus calamus, Arisaema atrorubens (= A. triphyllum), A. japonicum (= A. serratum), A. serratum, A. thunbergii, Lysichiton camtschatcensis, Pinellia ternata, P. tripartita, Amorphophallus campanulatus (= A. paeoniifolius) and A. kiusianus were investigated). In certain other species of Amorphophallus starch is partly (A. bulbifer) or largely (A. konjac, A. oncophyllus, A. variabilis) replaced by glucomannans, which are located in giant idioblasts. A procedure to isolate starch-free glucomannans from tubers of Amorphophallus rivieri (= A. konjac) was described by Wootton et al. (1993); this glucomannan had a Gluc : Man ratio of 58 : 42. Literature concerning mucilages of Araceae is partly contradictory (Ahmed et al. 1968; Sunell & Arditti 1983:

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54–55). This is not surprising since mucilages are complex mixtures of heteropolysaccharides and very difficult to obtain in pure form. Crude mucilages are always mixed with variable amounts of proteins and pectic and other substances (see section on Pectins). If large amounts of mucilage are present in large idioblasts it is relatively easy to obtain a fairly pure mucilage, as is the case with the glucomannans of some species of Amorphophallus. If the major storage compound is starch, however, the water-soluble “polysaccharides” will always be of different origin, either as pectic substances, true mucilages such as those contained in raphide idioblasts, some cell wall hemicelluloses or non-mucilaginous substances like proteins. According to Amin (1955) and others (see Sunell & Arditti 1983), mucilages purified from taro tubers (in several cultivars) are essentially branched arabino-galactans with an approximate gal:arab ratio of 8:1 to 11:1; this is perhaps the mucilage of the raphide idioblasts. The mucilages isolated in 3–4% yields from tubers of Eminium spiculatum and Arisarum vulgare by Ahmed et al. (1968) were mixtures of pectic substances, hemicelluloses and true mucilages. Starch grain morphology Czaja (1969, 1978a, b) investigated rhizomes, tubers, corms, stems and seeds of many araceous taxa for the presence and structure of starch grains and observed two main types: large grains of the so-called envelope-layer type (Hüllen-Lage-Stärkekörner) and small granules which are aggregated in compound grains. Two main subtypes of compound grains were distinguished by Czaja: highly compound grains (hochzusammengesetzte Stärkekörner, Czaja 1969: 37–38), composed of 400 and more small granules, and compound grains (höher zusammengesetzte Stärkekörner, Czaja 1969: 34–37) with usually up to 10 granules having a diameter of more than 6µm; these latter granules usually belong to the envelope type (Hüllen-Stärkekörner). Highly compound grains were observed by Czaja (1978a) in all araceous seeds; 54 species from 17 genera were investigated. Rhizomes, tubers and stems are less homogenous (90 species from nearly 40 genera were investigated). Some store large grains and some store compound grains. Czaja’s statements (1969: 36–38, 1978a: 60) concerning the two subtypes of compound grains mentioned above are sometimes discordant (e.g. Anthurium, Colocasia, Pinellia). On consulting the author’s table 1 (description of starch grains) and the figures of starches of Amorphophallus campanulatus (= A. paeoniifolius), A. oncophyllus and A. variabilis given in Ohtsuki’s (1967) paper, it becomes clear that Czaja’s two subtypes of compound starch grains often merge into one another. The highly compound type in pure form is probably less frequent in vegetative parts of Araceae than is suggested by Czaja’s list (1978a,b). For other detailed studies of starch types in Araceae, see Seubert (1993).

Pectins The following remarks concern the occurrence of pectins in non-lignified cell walls. Jarvis et al. (1988) showed that dicots and part of the monocots have primary walls with more than 150 mg of galacturonans per gramme of cell wall preparations (high contents). Grasses and other Commeliniflorae had low (< 50 mg/g) galacturonan contents. Alocasia and Lemna (Ariflorae) and all investigated Alismatiflorae and Liliiflorae belonged to the high-content group; some monocot taxa had intermediate (50–150 mg/g) galacturonan contents (see section on Mucilages).

Secondary metabolites Saponins, phenolic compounds including flavonoids, cyanogenic glucosides, the widespread occurrence of constituents which cause skin irritation and painfully acrid sensations on mucous membranes (mouth, throat, eyes) and calcium oxalate raphides may be considered the key chemical characters of the family (Hegnauer 1963, 1986; Bown 1988).

Saponins Saponins are by no means ubiquitous but were shown to be probably present in a number of taxa. Hegnauer (1963) stressed the lack of chemical knowledge about araceous saponins; the occurrence of steroidal sapogenins was and still remains uncertain (Hegnauer 1986). Nevertheless, the occurrence of steroidal saponins was mentioned for Montrichardia and Pinellia (Dahlgren & Clifford 1982) and for Arales without indicating genera (Dahlgren et al. 1985). This discrepancy is most probably caused by the fact that Altman (1954 in Hegnauer 1963) reported Montrichardia arborescens to be a rich source of steroidal sapogenins. His analytical method, however, was unreliable. Haemolytic and foam-producing substances, generally believed to be saponins, were shown to be present in many species (Clark & Waters 1934, Fontan-Candela 1957, Hegnauer 1963, Schroeter et al. 1966), but authors screening for steroidal saponins (Villar Palasi 1948, Anzaldo et al. 1957, Wall et al. 1954–1961) never obtained indications for this type of saponin nor could they detect or isolate steroidal sapogenins after hydrolysis (Marker et al. 1947, Wall et al. 1954–1961). According to Villar Palasi (1948), saponins are probably present in some parts of Alocasia odora and Arum italicum, according to Anzaldo et al. (1957) in Acorus calamus (Acoraceae), Amorphophallus campanulatus (= A. paeoniifolius) and Colocasia esculenta, and according to Wall et al. (1954–1961) in Acorus calamus (Acoraceae), several taxa of Colocasia, a species of Philodendron and Symplocarpus foetidus (confirmed by Segelman & Farnsworth 1969). All these researchers failed to detect saponin-like substances in

P H Y TO C H E M I S T RY A N D C H E M OTA X O N O M Y

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the investigated plant parts of Arisaema triphyllum, Arisarum vulgare, Amorphophallus bulbifer, several taxa of Colocasia, Dieffenbachia cordata, Dracunculus canariensis, Monstera deliciosa, Montrichardia arborescens, Orontium aquaticum, Peltandra virginica, Pistia stratiotes and Zantedeschia aethiopica. No appreciable amounts of saponins were present in Anchomanes difformis or Cyrtosperma senegalense (= Lasimorpha senegalensis) (Delaude-Hulst 1974), or Typhonium brownii (Simes et al. 1959). Summarizing, it may be stated that saponin-like substances occur occasionally in Araceae but that the chemistry of the saponins is still totally unknown.

Phenolic compounds Phenolic compounds occur in large amounts and are structurally and biosynthetically diverse. Bate-Smith (1968) investigated hydrolysed leaf extracts of 24 araceous plants. Ten of them contained procyanidins (formerly leucocyanidins), seven contained quercetin, six kaempferol, ten caffeic acid, seventeen p-coumaric acid, twelve sinapic acid and eleven contained ferulic acid; in the case of Orontium aquaticum the presence of scopoletin was indicated. A much more comprehensive survey of proanthocyanidins, cinnamic acids, anthocyanins and flavonoids of Araceae was published by Williams et al. (1981). According to these investigators, procyanidins occur in leaves of nearly half the investigated taxa and C-glycoflavones are the characteristic leaf flavonoids of the family. In some taxa they are accompanied or even replaced by O-glycosides of flavonols or flavones. Sulphates (esters of sulphuric acid) of vitexin, isovitexin, vitexin 7-glucoside, chrysoeriolgalactoside and quercetin occur sporadically (Philodendron ornatum, Culcasia saxatilis, Scindapsus pictus). As yet not fully characterized sulphates of esters of caffeic acid (sulphated caffeoyl glucoses?) are much more common, being mainly present in subfamilies Monsteroideae (67%), Philodendroideae (23%) and Pothoideae (20%); non-sulphated caffeic acid derivatives with a free carboxylic group are common in subfamilies Colocasioideae (80%), Lasioideae (38%) and Pothoideae (20%) (subfamily circumscriptions in Williams et al. 1981 follow Bogner 1979a). The results of this investigation have been summarized twice by the authors (Harborne 1982; Williams & Harborne 1988). It is highly probable that the two types of acidic caffeic acid derivatives detected by Williams et al. (1981) are involved in the irritating properties distinguishing most aroids (see Irritants in Araceae). Ellis et al. (1983) showed that the chemistry of the proanthocyanidins of the fruits of Zantedeschia aethiopica (two types investigated) and Z. rehmannii varies with the taxon. They contain afzelechin, catechin and (or) gallocatechin and their epimers as building blocks and consist either of pure procyanidins, mixtures of propelargonidins and procyanidins or mixtures of prodelphinidins and procyanidins. Since p-coumaric acid

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THE GENERA OF ARACEAE

and ferulic acid are common in the family it is worth mentioning that they never seem to be bound to the cell wall polysaccharides. Harris & Hartley (1980) showed that p-coumaric, ferulic and diferulic acids occur combined with non-lignified cell walls in all investigated members of Commelinidae (sensu Cronquist 1981) and in Arecaceae, Philydraceae, Pontederiaceae and Haemodoraceae in the strict sense. Araceae (Arum italicum, Pistia stratiotes and Sauromatum venosum have been investigated) and Acoraceae (Acorus calamus), like Alismatidae and typical Liliidae, lack this character. Acorus calamus has p-hydroxybenzoic acid combined with non-lignified cell walls; this feature occurs erratically in vascular plants. Phenolic amines are dealt with in the section on Biogenic amines and alkaloids. Recapitulating, it may be said that proanthocyanidins and C-glycoflavones and derivatives of caffeic acid belong to the main phenolics of the family, but that glycoflavones may be replaced in certain taxa or populations by flavonol glycosides (Stylochaeton; rutin in middle European Arum maculatum; kaempferol 3,7-bisglycoside in Gymnostachys anceps) or flavoneO-glycosides (tribe Areae sensu Bogner 1979a) as main leaf flavonoids (Williams et al. 1981, Hegnauer 1986, Williams & Harborne 1988). Leaves of Eminium spiculatum yielded the C-glycoflavones vitexin, isovitexin and its 7-glucoside (= saponarin), iso-orientin and its 7-galactoside, vicenin-1, the flavone-O-glycosides luteolin 3’-glucoside, luteolin 7-glucoside and chrysoeriol 7-glucoside and the flavanone glycoside eriodictyol 7glucoside (Shammas & Couladi 1988). A second lignanoid compound (see also acoradin, in Essential oils), a 1-ethyl-2-methyl-3-aryl-indane derivative, was isolated from rhizomes of Acorus calamus (Saxena 1986); it seems to arise by spontaneous dimerisation of asarone (Al-Farhan et al. 1992). A series of glucosylated lignans was recently isolated from subterranean parts of Arum italicum, together with ferulic acid and glycosides of coniferyl alcohol and 4coumaryl alcohol (Della Greca et al. 1993). Whole plants yielded traces of 8-O-3’ and 8-O-4’ neolignans (Della Greca et al. 1994; see also Gellerstedt et al. 1995). Inflorescences of Zantedeschia aethiopica yielded the C-glycoflavones swertisin and swertiajaponin (Sivakumar & Nair 1992).

Cyanogenic glucosides Cyanogenic glucosides have been known since the work of Jorissen, Greshoff and Treub (see Hegnauer 1963, 1977, 1986) to be rather common in the family. The distribution and chemistry of cyanogenesis (the ability to release HCN after injury) in aroids have been thoroughly discussed by Hegnauer (1963, 1973, 1977, 1986). Additional observations have been published by McBarron (1972: one of three tested samples of Gymnostachys anceps was weakly and two were strongly cyanogenic), Kaplan et al. (1983: two out of

1-22 Section A Acro 18/7/97 8:02 Page 39

O Gluc

NH 2 HOOC HOOC

COOH

HO

CN

II

1

I

C

CO2 O Gluc

OH NH 2

COOH

COOH

HO

VII

OH

+ 12_ O2

O

V

2

N

O

OH

IV

CH3 COOH

VI

O HO

NH 2

NH 2

HO

III HO

IX

RO

3

RO

4

1

COOH

HO CO2

VIII

CHO OH 1

XII 8’

HO Y

3

14’

5 1’

XIV

COOH

17’

Me

11’

HO Y

XI

B- ring OH R

O

8 7

1

B

O

A PA L

Gluc

6

A-ring OH

O

XV NH 2

s everal ty pes of amid es

COOH

CH3 COOH or CH2 HOOC

COOH

XIII

X

Figure 8. Some general lines of secondary metabolism of Araceae and some of their key chemical characters I-IX = Tyrosine and some of its metabolites: I = Tyrosine; II = cyanogenic glucoside Triglochinin; III = oxoaporphine Liriodenine; IV and V = Homogentisic acid and its 2-glucoside which are responsible for the so-called egumi-taste according to Hasegawa et al. (1959); VI = Acetic acid generated by total catabolism of tyrosine; VII = Tyramine; VIII = DOPA; IX = Dopamine, one of the possible causes of melanogenesis in dying parts of araceous plants; X-XII = presumably metabolites of phenylalanine:– X = Phenylalanine; XI = p-Coumaric (Y = H), Caffeic (Y = H, OH), Ferulic (Y = H, OMe) and Sinapic (Y = OMe) acids; XII = 3,4-Dihydroxybenzaldehyde (R=H) and its diglucoside, 3,4-Diglucosyloxybenzaldehyde (R = Glucosyl), which are responsible, at least partially, for “Hange” and “Taro” acridity according to Suzuki et al. (1975); XIII and XIV = pure Acetogenins:– XIII = Acetic or Malonic acids (activated forms); XIV = main allergen of Philodendron, 5-Heptadec-8’,11’,14’-trienylresorcinol; XV = the 6-C-glucoflavones Isovitexin (= Saponaretin: R = H) and Saponarin (R = Glucosyl); flavonoids are of mixed biogenetic origin. 1 and 2 = enzymes of tyrosine catabolism; 2 = “Homogentisicase” of Hasegawa et al. (1959). PAL = Phenylalanine lyase ➤ = alternative pathways indicated.

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four tested samples of an Anthurium species and four out of six samples of Philodendron corcovadense were cyanogenic) and Gibbs (1974: cyanogenesis was demonstrated for Anthurium aemulum, A. scandens, Calla palustris). I could not confirm the latter for a Dutch sample of this species, Schismatoglottis sp. (“S. ruttenii”), Xanthosoma lindenii (= Caladium lindenii) or Zantedeschia rehmannii. In the case of Gibbs (1974), only personal observations are taken into consideration because of his rather uncritical evaluation of results published by other investigators; he accepted without comment many highly doubtful cases of cyanogenesis reported in the literature. Triglochinin, a seco-derivative of dhurrin or taxiphyllin, is the cyanogenic glucoside occurring in all Araceae so far investigated for cyanogenesis. It is accompanied by a substrate-specific enzyme (“triglochininase”) which rapidly splits the glucoside after injury. Cyanogenesis is therefore often extremely rapid in Araceae and in many instances the HCN is totally lost during the drying of plant parts. Triglochinin and the corresponding enzyme were shown to be the cause of cyanogenesis in Alocasia macrorrhizos (tribe Colocasieae), Arum maculatum (tribe Areae), Pinellia tripartita (tribe Arisaemateae), Lasia spinosa (subfamily Lasioideae) and Dieffenbachia picta (= D. maculata, tribe Dieffenbachieae) by Nahrstedt and his group (see Hegnauer 1986).

Polyhydroxy alkaloids or alkaloidal glycosidase inhibitors (AGIs) This class of biologically active secondary metabolites has been shown in recent years to comprise the toxic principles of Locoweeds (Astragalus, Oxytropis), Darling Peas (Swainsona) and Moreton Bay Chestnut or Black Bean (Castanospermum australe), all belonging to the Leguminosae. The same and similar compounds occur also in some fungi, a fern, several Euphorbiaceae, Moraceae and Polygonaceae, but had not been reported previously from monocotyledonous plants. A research group at Kew (Sharp et al. 1993) has been screening Araceae for this type of metabolite. So far about 70 species from 47 genera have been investigated, and appreciable amounts of AGIs were detected in leaves of several genera of subfamily Aroideae: Anchomanes, Nephthytis and Pseudohydrosme (tribe Nephthytideae) and Aglaonema and Aglaodorum (tribe Aglaonemateae). Trace amounts of AGIs were present in species of Amorphophallus (tribe Thomsonieae). Accumulation of AGIs thus appears to support the present circumscription of Nephthytideae and Aglaonemateae. The major araceous AGIs were identified as DMDP, a 2,5-dihydroxymethyl3,4-dihydroxypyrrolidine, an N-analogon of fructofuranose and two piperidine derivatives, HNJ (2,6-dihydroxymethyl-3,4,5-trihydroxypiperidine) and DMJ (= deoxymannojirimycine). Several minor com-

40

THE GENERA OF ARACEAE

pounds still await definitive identification. It is rather unexpected that no new AGIs were detected in Araceae; their three main polyhydroxy alkaloids had formerly been isolated from other taxa, e.g. Leguminosae and Euphorbiaceae. Obviously the synthesis of such sugar- and/or pipecolic acid-related compounds is a common feature of a large number of plants and microorganisms, but their accumulation is of much more restricted occurrence.

Irritants in Araceae Peckolt (1893; see Hegnauer 1963) reported long ago that several types of irritants must occur in Brazilian Araceae. A comprehensive summary of irritant aroids and their irritating properties is given by Mitchell & Rook (1979); see also Hegnauer (1963, 1986), and Bown (1988). Brazilian scientists have paid much attention to the irritating and toxic properties of Araceae. Lethal intoxications of children from eating the spathe and spadix of cultivated Zantedeschia aethiopica have been reported (Ladeira et al. 1975). Working with juices expressed from different parts of Dieffenbachia picta (= D. maculata) it could be demonstrated that stems and petioles are much more aggressive than leaf blades and the centrifugation of the stem and petiole juices resulted in an inoffensive supernatant and a highly offensive precipitate. Moreover, it was shown that the toxic principle(s) of this species is (are) not of proteinaceous nature and neither stable to prolonged heating nor to vacuum drying (Ladeira et al. 1975; see also Carneiro 1985). Later these observations were confirmed and raphides present in the precipitates were investigated; they lost their irritating and toxic properties on washing with ether, but not on washing with water (Jesus Neves et al. 1988). Dieffenbachia maculata contains an inhibitor of human salivary amylase and several other types of amylases (Padmanabhan & Shastri 1990). It is, however, improbable that such inhibitors are involved in Dieffenbachia toxicity. Contact dermatitis Contact dermatitis is caused by some species of Philodendron. Their main allergens have been isolated and identified as alkenylresorcinols with one, two or three double bonds; they are accompanied by corresponding tridecyl-, pentadecyl- and heptadecylresorcinols. Philodendron scandens subsp. oxycardium is the most noxious taxon investigated hitherto and may contain much 5-pentadecatrienylresorcinol. Philodendron angustisectum, P. erubescens and P. radiatum are also suspected of being able to cause contact dermatitis because mono- and di-unsaturated alkenylresorcinols have been detected in them. On the other hand, no alkyl- nor alkenylresorcinols were observed in P. bipennifolium, P. fenzlii, P. sagittifolium, P. squamiferum or P. tuxtlanum (Reffstrup et al. 1982, Reffstrup & Boll 1985).

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Non-allergenic skin irritations Non-allergenic skin irritations can occur in persons handling large quantities of horticultural Araceae. They may be caused by the combination of raphides and papain-like proteolytic enzymes like the so-called dumbcain described for taxa of Dieffenbachia. This type of skin lesion is best known from workers in the pineapple industry (Ananas comosus, Bromeliaceae).

metabolite of tyrosine, and its 2-glucoside were shown to be responsible for this taste sensation. It should not be forgotten, however, that large amounts of soluble oxalates as well as certain proanthocyanidins can also produce taste sensations of a tart to slightly bitter, astringent or acrid nature. Moreover, egumi-taste gradually passes into taro acridity (Hasegawa et al. 1959).

Irritation of mucous membranes Painful irritations of mucous membranes are caused by many araceous plants. Long ago, Chauliaguet (1897; see Hegnauer 1963) showed that raphides alone are harmless. Notwithstanding this fact, raphides are still believed to be the main irritating factor of Araceae by many modern authors who are not familiar with the pertinent literature. Raphides, however, are only the vehicles of irritating substances. The construction and contents of araceous raphide idioblasts (Wiley 1903; Safford 1905; Middendorf 1983; Tang & Sakai 1983) and the structure of the individual needles (Sakai et al. 1972; Tang & Sakai 1983) suggest that they are adapted to transport acrid, pain-producing and otherwise irritating substances. According to Suzuki (1969, 1975), 3,4-dihydroxybenzaldehyde (protocatechualdehyde) and its 3,4-bisglucoside are acrid constituents of tubers of Pinellia ternata and Colocasia antiquorum (= C. esculenta). Presumably the sulphated and acidic derivatives of caffeic acid mentioned earlier also take part in araceous acridity. It is still uncertain whether specific pain-producing substances are also involved in painful irritations of mucous membranes caused by Araceae. Pain-producing oligopeptides like moroidin of Laportea moroides in the Urticaceae (Leung et al. 1986) may be part of the “dart” poison of some Araceae. Saponins may also be involved in the acridity of aroid species which produce and store these substances; it is known that several saponins (formerly called sapotoxins) are highly irritating to mucous membranes. According to Tang & Sakai (1983), Suzuki’s (1969, 1975) diglucosyloxybenzaldehyde was a misidentification; they assumed that in fact the isolated compound was 5-(hydroxymethyl)-furfural, a decomposition product of some hexoses under acidic conditions. Notwithstanding the fact that the reports of Suzuki (1969) and Suzuki et al. (1975) contain inaccuracies, it is clear that Tang & Sakai failed to read carefully Suzuki’s (1969) paper and it is highly improbable that their acrid bisglucoside C19H26O13, was a derivative of 5(hydroxymethyl)-furfural.

Biogenic amines and alkaloids

Egumi taste Another effect of Araceae on mucous membranes was described by Hasegawa et al. (1959) for dried tubers of Pinellia ternata (“Hange”). They have a harsh, somewhat bitter and astringent taste similar to the so-called egumi-taste of edible shoots of some cultivars of Phyllostachys edulis (Gramineae subfamily Bambusoideae). In both cases homogentisic acid, a

Biogenic amines and alkaloids occur in many Araceae. True alkaloids seem to be rare but perhaps rather significant from the taxonomic standpoint because they indicate biochemical relationships with Magnoliidae. The aporphine-type alkaloids liriodenine and lysicamine were isolated from roots of Lysichiton camtschatcensis. Tubers of Pinellia ternata yielded ephedrine. A tertiary base, C20H35O2N, was isolated from tubers of Eminium spiculatum (Ahmed et al. 1968). Traces of volatile biogenic amines, but no coniine, were shown to be present in tubers and leaves of Arisarum vulgare, Arum maculatum (here also traces of nicotine were present) and Eminium spiculatum. Just before and during flowering, spadices and spathes of a number of Araceae produce large amounts of volatile amines and indoles, giving them a putrescent odour attractive to their pollinators. The composition of these “perfumes”, which may even contain skatol, is taxon-dependent (Hegnauer 1986; Bown 1988). The temperature of the upper part of the spadix rises considerably just before anthesis; this increases volatilization of amines. The substance triggering heat production in the spadix is known as “calorigen” and was shown recently to be salicylic acid in Sauromatum guttatum (= S. venosum) by Raskin et al. (1987). Spathes and male and female flowers of many Araceae also produce large amounts of non-volatile amides of p-coumaric and ferulic acids with putrescine, spermidine, tyramine and, in Zantedeschia aethiopica, spermine. Large amounts of free tyramine and (or) dopamine were present in these parts of Monstera deliciosa, Philodendron andreanum (= P. melanochrysum), P. erubescens, P. martianum, P. scandens, P. selloum (= P. bipinnatifidum), P. tripartitum, Remusatia vivipara, Rhaphidophora decursiva and Zantedeschia aethiopica, but not in Arisarum vulgare, Arum maculatum, A. italicum, or Dracunculus vulgaris. Up to 4 mg dopamine/g fr. wt were observed in one month-old ovaries of Monstera deliciosa (Ponchet et al. 1982). The tendency for spathes and other plant parts to turn black (melanogenesis) in a number of Araceae may be due to the simultaneous presence of dopamine and phenol oxidases. The starch-rich tubers of North African and Mediterranean Arisarum vulgare are known in Morocco as “Irni” or “Erni”. This name is used, however, for several taxa of subfamily Aroideae with starch-rich edible tubers which also contain toxins, and are suitable as food only after adequate treatments

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such as heating and drying or repeated cooking etc. (Bellakhdar 1978). A pyrrolidine alkaloid called irniine, C20H33N, was isolated from tubers of Arisarum vulgare and shown to be one of the toxic principles of this species (Melhaoui et al. 1992).

Miscellaneous compounds Essential oils Acorus (Acoraceae), with two species, A. calamus and A. gramineus, is a highly aromatic taxon. The roots, rhizomes and leaves produce large amounts of essential oils, which are stored in idioblasts similar to the oil cells of woody polycarps (Magnolianae sensu Takhtajan 1959), aromatic Gramineae and Zingiberaceae. Both species of Acorus comprise several chemodemes with respect to the composition of essential oils. Best known are the taxa which produce the biologically active phenylpropanoid cis-asarone (= ß-asarone), which has insecticidal and antifeedant properties (Koul et al. 1990). This compound occurs together with the less toxic trans-asarone, mainly in triploid European A. calamus and in much larger quantities in Indian tetraploids (Hegnauer 1963, 1986; Röst 1979b). Rhizomes of Indian A. calamus also contain the dimethylbutanoid lignan acoradin (a dimer of cis-asarone, see Saxena & Mukherjee 1985), as well as asar(yl)aldehyde, acoramone, and galangin; for mono- and sesquiterpenoid constituents of Acorus oils, the reader is referred to Röst (1979b), Röst & Bos (1979) and Hegnauer (1986). Stereoisomers of acoradin are andamanicin from Piper sumatranum var. andamanicum, heterotropan from Asarum taxa and magnosalin from Magnolia taxa (Malhotra et al. 1990). A chemotype of Acorus gramineus cultivated in Italy accumulates a number of phenylpropanoids, including ß-asarone and its epoxide, which suppress growth of several microalgae (Della Greca et al. 1989). Another aromatic genus of Araceae is Homalomena (Wealth of India 1959; Bown 1988: 223). H. aromatica yielded 1.2% essential oil containing mainly monoterpenoids (Hegnauer 1986). Rhizomes of Homalomena aromatica contain an essential oil with up to 80% of linalool and several sesquiterpenoids of which the homalomenols C and D are new compounds with the rather rare carbon skeleton of mintsulphide and the aphanomols (Sung et al. 1992). Rhizomes of Homalomena occulta, a Chinese medicinal crude drug, yielded 0.79% of essential oil with much linalool and lesser amounts of other monoterpenoids and sesquiterpenoids (Zhou et al. 1991). Pistia stratiotes produces similar allelochemicals, one of which was shown to be asarone (Aliotta et al. 1991). Peltandra virginica is pollinated by the chloropid fly Elachiptera formosa, which also breeds in the inflorescences of this plant. This peculiar symbiosis between a plant species and its pollinating insect seems to depend, at least in part, on

42

THE GENERA OF ARACEAE

odoriferous compounds, which are synthesized by spathes and spadices of Peltandra and which function as a lure for the pollinator (Patt et al. 1992). Notwithstanding the toxicity of asarone to many insects, it seems to have become, together with other volatile phenylpropanoids, asarylaldehyde and the sesquiterpenoid acarogermacrone, a trigger for certain behavioural activities of fruit flies of the genera Ceratitis and Dacus (Jacobson et al. 1976). Larvae of fruit flies are often serious orchard pests and substances attracting males and (or) females may become useful as lures for capturing adults of pest-causing insects. An example of such a use is Spathiphyllum cannifolium, which is planted in Northern Thailand around orchards. Its flowering inflorescences do not emanate foetid amines, but a mixture of benzyl acetate, methyleugenol, methylchavicol, p-methoxybenzaldehyde, propyltetradecanate and yet other compounds, which are highly attractive to several species of fruit flies; Lewis et al. (1988) report the same attraction for the fruit fly Dacus musae in Northern Queensland. Another fascinating observation was published by Seidel et al. (1990) as follows. Anthurium gracile, A. ernestii and Philodendron megalophyllum belong to the so-called ant garden plants found in lowland Amazonian Peru. Workers of the ant Camponotus femoratus collect seeds of these and some other, taxonomically unrelated plants and store them in brood chambers where they later germinate. Volatile aromatic compounds may be cues which initiate collection of the seeds of these epiphytic plants by ants. In the case of the aroid species, 6-methyl methyl salicylate, benzothiazole and vanillin were detected in seed coats and adhering fruit parts.

Phytosterols and triterpenes Sitosterol-type phytosterins are ubiquitous in vascular plants. ß-sitosterol palmitate was isolated from tubers of Amorphophallus campanulatus (= A. paeoniifolius) together with other phytosterols, triacontane, lupeol and betulinic acid by Chawla & Chibber (1976). New di- and trioxigenated sterols have been isolated from tubers of Colocasia esculenta (Ali 1991) and from Pistia stratiotes (Aliotta et al. 1991; Monaco & Previtera 1991). As far as I am aware, there exist only a few other reports concerning the occurrence of triterpenes in Araceae. Taraxerol acetate was isolated together with phytosterins, lignoceric acid, hentriacontane, hentriacontanol and hentriacontanone from rhizomatous stems of Alocasia fornicata (Sharma et al. 1972). Air-dried aerial parts of Xanthosoma robustum collected in Oaxaca, Mexico, yielded four antibacterial hydroperoxy derivatives of 4,14-dimethyl-cholesta-8en-3ß-ol and of cycloartenol (Kato et al. 1996).

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Epicuticular waxes Behnke & Barthlott (1983), Fröhlich & Barthlott (1988) and Barthlott (1990) showed that the presence and architecture of epicuticular waxes is a character useful for the classification of monocotyledons. Their so-called Strelitzia-type and Convallaria-type seem to be restricted to monocots and to characterize major taxa, i.e. either parts of the Zingiberiflorae, Commeliniflorae, Areciflorae and Bromeliiflorae, or parts of the Liliiflorae. The Ariflorae, like the Alismatiflorae, Triuridiflorae and Dioscoreales (Liliiflorae), lack these characteristic types of epicuticular waxes (compare Dahlgren et al. 1985: 65, 96, 98–99).

2. Chemotaxonomy It is advisable not to put too much weight on chemical characters with regard to estimating phylogenetic relationships of the Araceae as long as the chemical structures of araceous saponins remain unknown. As stated here and in an earlier publication (Hegnauer 1986), there are similarities between the primary and secondary metabolism of the Araceae and a rather large number of other monocotyledonous taxa. The most striking chemical attributes of the

family are metabolites derived from tyrosine (triglochinin and tyramine, dopamine and their amides; egumi-taste principles (Hasegawa et al. 1959) and the presumably related, acrid constituents). However, the participation of tyrosine in the production of secondary metabolites such as the cyanogenic glucosides dhurrin, taxiphyllin and triglochinin (Hegnauer 1973, 1977, 1986: 356–357) and the amaryllidaceous and colchicinoid types of alkaloids (Hegnauer 1986: 316, 576), is widely scattered in monocotyledons. Ultimately, one point cannot be overemphasized. Use of chemical characters for taxonomic purposes can only be meaningful if all available facts are checked and evaluated carefully. Authors who neglect to exploit information made readily available in handbooks (Wehmer 1929, 1931, 1935; Hegnauer 1963, 1986 (for monocotyledons)) do not do a good job as far as the chemotaxonomic part of their work is concerned. Furthermore, the need to consult as many original papers as possible can be illustrated by two curious examples. First, the cause of the painful acridity of most araceous plants is still ascribed solely to the presence of raphides by many authors. Secondly, the summary in Hegnauer (1986), based on the original paper, of the results of the phenolic research carried out by Williams et al. (1981) is actually more accurate than a summary later given by the authors themselves (Williams & Harborne 1988).

P H Y TO C H E M I S T RY A N D C H E M OTA X O N O M Y

C

43

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12 E C O L O G Y A N D L I F E F O R M S

C

Croat (1990, 1992a) has published a more detailed review of araceous life forms and ecology. The growth of Araceae is dependent on abundant available water and prevailing atmospheric humidity. Structurally and physiologically they are not well adapted for growth in arid or cold conditions, and hence do not occur in the most extreme environments. Araceae are most diverse and abundant in the humid tropics and it is there that the richest variety of their life forms is found. Relatively few genera inhabit temperate regions of the world and these are either geophytes (e.g. Arisaema, Arum, Pinellia) or helophytes (Calla, Lysichiton, Orontium, Peltandra, Symplocarpus, ). The very few genera found at high altitudes exist in a warm temperate climatic regime and are also geophytic. Gorgonidium has been found at around 3000m in the Andes, while Arisaema occurs at up to 3000m in Africa (A. ruwenzoricum in the Ruwenzori) and 4400–4500m in the Himalaya (A. flavum, A. jacquemontii, A. lobatum).

Taxonomic considerations From the taxonomic and evolutionary viewpoints there are some generalizations that can be made concerning life forms. Hemiepiphytes are commonest in the more primitive tribes and subfamilies. Most genera of subfamilies Pothoideae and Monsteroideae are hemiepiphytes and among more advanced genera this life form occurs only in tribe Culcasieae, Philodendron and Syngonium, all belonging to subfamily Aroideae. These latter genera show marked structural adaptations in their habit and in these features must be considered derived. Aquatic, subaquatic and helophytic genera are scattered throughout the family from very primitive groups such as subfamily Orontioideae to very advanced ones such as tribe Cryptocoryneae and Pistia in subfamily Aroideae. The least specialized mesophytic habit is shown by some rainforest terrestrial herbs. In these the stem is aerial and erect or decumbent, with short but distinct green internodes. This habit type has been judged primitive in the family by some previous authors (e.g. Grayum 1990), but in fact is found predominantly in more advanced genera. Typical examples are Aglaonema (tribe Aglaonemateae), Dieffenbachia (tribe Dieffenbachieae), Homalomena (tribe Homalomeneae) and Schismatoglottis (tribe Schismatoglottideae), all of subfamily Aroideae. Among primitive groups only tribe Spathiphylleae and terrestrial Anthurium species have

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THE GENERA OF ARACEAE

this habit. Many other genera, however, seem to represent relatively small adaptive shifts from a mesophytic norm. Within subfamily Lasioideae for example, Lasia, Podolasia and many species of Cyrtosperma have habits which are intermediate between the helophytic/mesophytic and geophytic/mesophytic categories. Nephthytis, in which the rhizomes normally grow superficially, has a considerably more mesophytic habit than the strongly tuberous stems of the other two genera of tribe Nephthytideae. Culcasia has many terrestrial species, spanning the hemiepiphytic/mesophytic categories, and Philodendron is similar. Anubias is predominantly helophytic but Dieffenbachia and Spathiphyllum, while typical of wetter habitats, also occur on drier ground within a humid tropical habitat. In the predominantly geophytic tribes Caladieae and Colocasieae, the genera Alocasia, Colocasia and Xanthosoma have mesophytic species with decumbent to erect, arborescent stems, while Steudnera and Chlorospatha are exclusively of this type. The most primitive Araceae, subfamilies Gymnostachydoideae and Orontioideae, are geophytes, rhizomatous helophytes or aquatics, and largely extratropical. While their habits are doubtless a prerequisite for survival in a more demanding climate, and therefore could have evolved from a mesophytic common ancestor, it is nevetherless equally possible that the mesophytic habit has evolved various times within the more advanced subfamilies from geophytic or helophytic ancestors. The geophytic habit is strongly represented in the relatively primitive subfamily Lasioideae and particularly common in the most advanced subfamily Aroideae. The rheophytic habit is characteristic of tribe Schismatoglottideae, the genera being almost exclusively rheophytic except for Schismatoglottis, which consists mainly of terrestrial mesophytic herbs.

Hemiepiphytes Humid tropical forests are the characteristic habitat of hemiepiphytic genera. The species vary considerably in size, from shortly climbing plants found on the major branches or trunks of trees (some Anthurium species) to huge plants with attached stems growing high into the forest canopy and producing enormously long, pendent flowering stems (e.g. Philodendron scandens). Hemiepiphytes can be divided into primary and secondary hemiepiphytes. Primary hemiepiphytes begin growth above ground level but produce feeder roots which eventually grow

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down to the forest floor. Secondary hemiepiphytes germinate on the forest floor, grow up tree boles, become detached from the ground by rotting of the juvenile stem but then become reconnected later by feeder roots which grow down from the upper internodes. Hemiepiphytic aroids typically have anchor roots as well, and are thus often called “root climbers”. Flagelliform shoots, heteroblastic leaf development and shingle plants (see chapter 2) are characteristic features of hemiepiphytic Araceae, though not present in all species of each genus. Highly developed heteroblasty coupled with skototropism, a specific growth strategy for seeking host tree boles, has been described in Monstera (Madison 1977a, Strong & Ray 1975). In certain species the seedling is a very slender, non-photosynthetic plant with long internodes and minute scale leaves. Having germinated on the forest floor it seeks the defined area of shadow represented by the nearest tree bole. Once the tree has been reached the plant transforms itself into the shingle form and later, higher up, into a mature flowering plant. Vegetative reproduction may then take place by the production of flagelliform shoots. Seed size is almost certainly an important element in the growth strategies adopted by hemiepiphytes. In Monstera seeds are relatively large and lack endosperm, which probably increases the efficiency and duration of the seedling’s nutrient supply until it has reached a suitable habitat for photosynthesis. Other hemiepiphytes which produce flagelliform shoots have numerous very small seeds and endosperm (e.g. Philodendron fragrantissimum, Rhodospatha latifolia) and probably have a different kind of seedling development. More observations are needed, especially in tropical Asia. To date the most important ecological observations of hemiepiphytic aroids have been made in tropical America (Blanc 1977a, b, 1978, 1980; Madison 1977a, Ray 1986, 1987a–c, 1988, 1990) and tropical Africa (Knecht 1983). Cercestis (e.g. C. mirabilis) and Philodendron (e.g. P. linnaei), among other genera, have species with another growth strategy aptly termed “rhythmic growth” by Blanc (e.g. 1977a). The mature flowering region of the stem is short with abbreviated internodes and more-or-less rosulate foliage leaves. The continuation shoot climbs upwards and is slender and flagelliform with cataphylls instead of foliage leaves. After an interval it produces another rosulate-leaved mature zone. The repetition of this pattern produces a series of connected rosulate plants one above the other on a single tree trunk.

Epiphytes True epiphytes, which never become connected to the ground by feeder roots, are found in Anthurium, Arophyton, Philodendron, Remusatia, Scindapsus and Stenospermation. The seeds presumably germinate directly on the host tree after dispersal by birds or other animals. Many species of Anthurium sect. Pachyneurium (e.g. A. hookeri) and some

Philodendron species (e.g. P. insigne) are litter basket epiphytes. The large leaves form an inverted cone in which leaf litter and other debris accumulate and into which the roots grow and ramify in a dense mass. Remusatia vivipara, which has a tuberous stem, is a widespread epiphyte, due to the dispersal of hooked bulbils which are probably transported by birds and primates high in the forest canopy. Arophyton buchetii appears to occur only as an epiphyte in leaf litter accumulated within large Pandanus crowns.

Lithophytes Many hemiepiphytes, epiphytes and geophytes are also found as lithophytes in suitable conditions. Certain groups, such as the Anthurium coriaceum complex in eastern Brazil, are characteristically lithophytic. Different species of this complex grow in humid coastal forests (e.g. A. coriaceum) where they are common on exposed areas of outcropping rocks, and in the semi-arid interior (A. erskinei), where they survive exposure during the prolonged dry season. Vining hemiepiphytes frequently grow on rocks in forest regions wherever shade and humidity are sufficient, the rock surface providing much the same conditions for attachment as tree boles. A number of geophytes are characteristically found growing in the eroded, litter- or humus-filled cavities of limestone outcrops; examples are Amorphophallus albispathus, Colocasia gigantea, and Typhonium albispathum in S.E. Asia and Amorphophallus hildebrandtii Carlephyton and Colletogyne in Madagascar. Rheophytes are also typically lithophytic.

Geophytes This category includes all genera with tuberous, rhizomatous, subterranean or partly subterranean stems. Geophytic aroids characteristically have periodic dormant periods when no leaves are present and these normally correspond to the dry season (or winter) of their habitat. However, rainforest geophytes exhibit growth periodicity and dormancy even in nonseasonal climates, e.g. Amorphophallus maculatus, A. titanum, Asterostigma riedelianum, Dracontium prancei, Zomicarpella amazonica. Several genera occur in more than one kind of climatic regime. In Stylochaeton, the rainforest species S. zenkeri is evergreen with unthickened roots and the inflorescence appears with the leaves. Other species, such as S. natalensis, grow in areas with a strongly marked dry season during which they are dormant. This species has thick, fleshy roots and usually flowers before the leaves or just as they emerge. The genera Amorphophallus and Dracontium are similarly diverse ecologically, with species in rainforest or in seasonal evergreen forest, deciduous forest, savannas or grasslands (A. abyssinicus, D. margaretae).

ECOLOGY AND LIFE FORMS

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Geophytes from deciduous forests, savannas or strongly seasonal grasslands flower without the leaves at the end of the dry season, mostly after the first rains fall. Leaf and fruit development take place during the rainy season. Variations occur in this basic phenological pattern. Biarum davisii (Crete and Turkey) flowers in the autumn (± November) after the rainy season has started, whereas B. ditschianum (Turkey) flowers at the end of the rainy season (± May) and the fruit development then takes place over a year (Bogner & Boyce 1989). The Mediterranean species of Arum (e.g. A. dioscoridis, A. italicum) grow during the relatively warm winter rainy season, whereas the more northerly A. maculatum grows from spring to summer and is dormant during the cold winter (Boyce 1993a). No Araceae occur in true deserts except Eminium spiculatum subsp. negevense, from the Negev desert (Koach 1988). Some species, however, grow in very dry areas, e.g. Arum and Eminium in central Asia, Arum and Biarum in North Africa and Asia Minor, Arisaema and Sauromatum venosum in the Arabian Peninsula and East Africa and a few Stylochaeton species of the Sahel zone of Africa. All these regions normally have some rain each year during which the plants grow vegetatively, or they may occur in places with a ground water supply. Zamioculcas zamiifolia is a succulent plant which stores water in its thick petioles and is sometimes found in very dry habitats, but it is more common in evergreen seasonal forests and savannas.

Rheophytes Rheophytes are flood-resistant plants, usually of tropical rainforests, growing in or along swift-running streams or rivers up to the flood level. They are characterized by narrow, leathery leaves and a firmly attached, usually epilithic stem. In addition to tribe Schismatoglottideae, in which the majority of genera have this habit, rheophytes are also found in Homalomena, Anubias and Holochlamys, and rarely in Anthurium.

Submerged or periodically submerged aquatics Jasarum steyermarkii and many Cryptocoryne species are permanently submerged plants (hydrophytes, sensu Cook 1990). Either the inflorescence as a whole (Jasarum) or its upper portion (Cryptocoryne) is held above the water surface while all other parts are completely submerged. Cryptocoryne is the largest genus of aquatic aroids and merits more detailed consideration. There are a number of species which are usually submerged but which are emergent at times of exceptionally low water (C. affinis, C. aponogetifolia, C. purpurea). The submerged leaves of such species are relatively large, whereas the emergent leaves are quite small, indicating that such conditions are unfavourable to their growth. The submerged and emergent leaves of the same

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THE GENERA OF ARACEAE

species generally look very different in shape, size, colour and structure. Submerged leaves are softer and emergent ones more coriaceous. Many species occur in the freshwater tidal zone where there is a daily cycle of exposure and submersion. Some species are found only in freshwater, like C. ferruginea, C. lingua and C. pontederiifolia, while others can grow both in fresh and brackish water (C. ciliata). A few species are helophytes, preferring swampy conditions and growing during the dry season completely emergent in normal soil, like C. spiralis, a weed of rice fields in India. Usually Cryptocoryne species flower at low water level when the plants become emergent. Cryptocoryne consobrina occupies a somewhat more specialized niche. The leaves are present only during the monsoon season when the streams are in flood. Flowering occurs after the monsoon rains when the streams have dried up and the leaves are shrivelled. During the dry season the plants are completely dormant with the rhizomes buried in the soil. C. nevillii flowers at the beginning of the monsoon season before the leaves appear or as they emerge.

Helophytes About 38 genera are helophytic or have at least some helophytic species, i.e. plants which grow in swampy habitats or along river and stream margins. Four of these genera (Gearum, Mangonia, Scaphispatha, Spathicarpa) are geophytes which habitually or frequently grow in seasonally flooded sites, Nineteen genera are strictly helophytes or are aquatics with some helophytic species (e.g. Cryptocoryne). This life form is thus widespread throughout the family in many different taxonomic groups, and in both temperate and tropical genera. There is little constancy in habit type. The stem may by tuberous (e.g. Caladium, Typhonium), rhizomatous (Typhonodorum), rhizomatous and arborescent (Montrichardia), semi-prostrate to aerial (Lasia), erect and arborescent (Philodendron) or merely shortly erect and aerial (Homalomena). The helophytic life form may be considered relatively unspecialized in the majority of genera which exhibit it. Tuberous or rhizomatous stems may be associated with seasonally flooded habitats and a marked dry season. Rhizomes may, on the other hand, be adaptations for colonizing muddy riverine margins as in the case of the strict helophytes Typhonodorum and Montrichardia. Genera such as Dieffenbachia, Homalomena and Spathiphyllum exhibit no special adaptations in their helophytic species, which appear to take advantage of wetter habitats for more vigorous growth rather than because of a strict requirement for a flooded substrate.

Free-floating aquatics The only free-floating species of Araceae is the pantropical Pistia stratiotes.

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13 P O L L I N AT I O N B I O L O G Y

Recent reviews of pollination biology and flowering phenomena have been given by Grayum (1984, 1986b, 1990) and Meeuse & Raskin (1988). Few thorough studies have yet been made of araceous pollination biology, though this is clearly a field of the utmost interest; most of the unusual and important taxonomic characters of aroid inflorescences and flowers are probably linked in one way or another to floral biological adaptations. Detailed studies of tropical genera have mostly been published only in recent years: Williams & Dressler (1976) for Spathiphyllum, Ramirez & Gomez (1978) for Monstera, Silva (1981) for Pistia, Shaw & Cantrell (1983) for Alocasia macrorrhizos, Gottsberger & Amaral (1984) for Philodendron and Young (1986) for Dieffenbachia. For temperate genera there are the classic studies of Arum by Knoll (1926) and Prime (1960), Vogel’s observations of fungus gnat pollination in Arisaema and Arisarum (Vogel 1978) and various studies of Arisaema (e.g. Barnes 1934, Bierzychudek 1982). Only Young (1986) and Bierzychudek (1982) have studied pollination biology in relation to entire populations of plants of the same species. There is a large literature which reports less critical studies or observations of insect visits to Araceae inflorescences. The phenology and behaviour of the spathe, spadix and flowers during anthesis are also subjects which in general have not been studied critically. Notable exceptions are the studies of Knoll (1926, Arum), Croat (1980, Anthurium), Gottsberger & Amaral (1984, Philodendron) and Meeuse & Raskin (1988, Sauromatum). Araceae inflorescences are almost always insect pollinated, although “wind tunnel” pollination has been proposed for Pinellia (Uhlarz 1985). The pollinators of Acorus (Acoraceae) are still unknown. Pollinators so far reported for Araceae (see Table 2) include trigonid bees, euglossine bees, beetles, flies, possibly thrips and very doubtfully mites. Some species of drosophilid flies are known to breed on the inflorescences of Alocasia, Colocasia and Homalomena (Okada 1986). They also exhibit specialization in their behaviour even on the spadix of a single species: stamenicolous species lay their eggs in the male zone while pistilicolous species lay their eggs in the female. Furthermore, several different pairs of fly species, one stamenicolous and the other pistilicolous, are known to breed in association on one aroid species (synhospitality). The trap mechanisms of genera with unisexual flowers and relatively complex spathes (e.g. Arum, Arisaema, Arisarum, Philodendron) have attracted most attention in pollination studies. Much less is

known about pollination mechanisms in genera with bisexual flowers and simpler or spreading spathes (e.g. Anthurium, tribe Monstereae, many Lasioideae). While “trap” seems an accurate description in Arum, Arisaema and Cryptocoryne, it is less clear in other genera (e.g. Amorphophallus, Philodendron) whether the pollinators are unable or merely unwilling to leave the inflorescence, once they have entered it, kept there perhaps by possible attractants such as stigma secretions, food bodies or sites for reproduction. Odour is evidently a prime factor in attracting pollinators and while Araceae are famous for foul inflorescence odours, which have been compared to dog faeces, horse dung, rotten fish, old socks, sulphurous gas, dead cow, mushrooms, cheese, etc., many others are not foul-smelling. Floral odours in Philodendron, Spathiphyllum and Xanthosoma are heavy and spicy and in Anthurium range from spicy to the smell of decaying fruit. At least two species of Amorphophallus are known to have a pleasant floral odour (A. galbra, A. manta). The wide range of odours must be correlated with different kinds of pollinator but though some studies of odour chemistry have been made (e.g. Meeuse 1966b, 1978, 1985), this fascinating field is largely unworked from a comparative standpoint. The colour of the spathe, and to a lesser extent of the spadix, varies considerably within the family, ranging from inconspicuous greens (e.g. Anthurium, Nephthytis) to elaborate patterns (e.g. Colletogyne, Sauromatum) or striking “flags” (e.g. Anthurium andraeanum). In fly pollinated species the spathe colours and patterns are known to be important in attracting pollinators (myriophile colours). Differentiated colour zones are frequent: in Philodendron many species have purple zones inside the spathe tube, while the blade is white or pale green. In Arisaema and Arisarum, the reverse situation is found, with white spathe tubes and dark purple blades which are often striped. The foul odours of such species as Amorphophallus konjac are very often associated with flesh-coloured or livid spathes, resembling carrion. By contrast, Zantedeschia aethiopica and many Spathiphyllum species have pure white spathes and perfumed inflorescences from which euglossine bees collect fragrances. This also occurs with naturalized Zantedeschia in tropical America (G. Gerlach pers. comm.). Trigonid bees collect pollen which also attracts certain beetles, e.g. Nitidulidae, that eat the fertile male flowers of Aridarum nicolsonii (Bogner, pers. obs.)

POLLINATION BIOLOGY

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Inflorescence heating (thermogenesis) in connection with flowering has been studied more, particularly in Arum (Rees et al. 1976, 1977), Sauromatum (Meeuse 1966a–b, 1975, 1978, 1985, Meeuse & Raskin 1988) and Philodendron (review by Mayo 1991). Thermogenesis in other genera has also been studied (e.g. Symplocarpus), but the only general comparative surveys are by Leick (1914, 1916, 1921 and, briefly, by Engler (1920b). While thermogenesis is very common in Araceae it is by no means universal. Its function is generally agreed to be to volatilize odour compounds,

and, more speculatively, thus to give precision both to the duration of the period of pollinator attraction (length of heating period), type of odour compound utilized (molecular weight) and distance over which pollinators can be attracted (maximum temperature). In other genera, however, odours are produced apparently without heating. Odour production and thermogenesis (when present) occur mainly in terminal appendices or in the male zones of the spadix in those genera that lack appendices (e.g. Philodendr on, Xanthosoma).

Table 2. Pollinators of Araceae. Data mainly from Grayum (1984, 1990).

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euglossine bees

Anthurium, Spathiphyllum, Xanthosoma

trigonid bees

Monstera, Spathiphyllum, Amorphophallus

beetles: Asilidae

Amorphophallus

beetles: Cetoniidae

Amorphophallus

beetles: Curculionidae

Anthurium, ?Pistia

beetles: Dermestidae

Dracunculus

beetles: Nitidulidae

Alocasia, Amorphophallus, Anchomanes, Anubias, Aridarum, Cercestis, Culcasia, Cyrtosperma, Nephthytis, Typhonium, Urospatha, Xanthosoma

beetles: Ptiliidae

Typhonium

beetles: Scaphidiidae

Pseudohydrosme

beetles: Scarabaeidae

Alocasia, Amorphophallus, Anubias, Caladium, Dieffenbachia, neotropical Homalomena, Philodendron, Syngonium, Xanthosoma

beetles: Scydmaenidae

Typhonium, Zantedeschia

beetles: Silphidae

Amorphophallus

beetles: Staphylinidae

Alocasia, Amorphophallus, Anthurium, Chlorospatha, Dracunculus, Lysichiton, Piptospatha, Pseudohydrosme, Typhonium

flies: Anthomyiidae

Alocasia

flies: Calliphoridae

Amorphophallus, Dracunculus, Helicodiceros

flies: Centropogonidae

Arum, Cryptocoryne

flies: Chloropidae

Peltandra

flies: Chorideae

Pseudohydrosme

flies: Drosophilidae

Alocasia, Colocasia, Culcasia, Homalomena, Nephthytis, Schismatoglottis

flies: Ephydridae

Cryptocoryne

flies: Neurochaetidae

Alocasia

flies: Phoridae

Cryptocoryne

flies: Psychodidae

Arum

flies: Sciaridae

Arisaema, Arum

flies: Simuliidae

Arum

flies: Sphaeroceridae

Arum, Pseudohydrosme

flies: Syrphidae

Peltandra

flies: Mycetophilidae

Arisaema, Arisarum

mites

Ambrosina (reported in literature but very doubtful as pollinators: fruit-set is extremely rare and possibly the pollinators are extinct).

thrips (Heterothrips arisaemae)

Arisaema (reported in literature but doubtful)

THE GENERA OF ARACEAE

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Unpleasant odours from the spathe have been observed in Dracontium and in the spathe blade of Lagenandra (Bogner pers. obs.) and Cryptocoryne (Vogel 1963, 1990, Bogner pers. obs.). In the spathe tube “kettle” of Lagenandra a different, fruity odour is produced which emanates mainly from the anthers and occasionally also from the so-called “olfactory bodies” above the female flowers (Buzgó pers. obs.). These structures are present only in Lagenandra and Cryptocoryne and their function is still unclear. The thickened connectives of stamens or synandria commonly found in several tribes of subfamily Aroideae (e.g. Philodendreae, Homalomeneae, Anubiadeae, Caladieae, Colocasieae) probably represent adaptations for osmophore function (see Vogel 1963, 1990 for an important survey of osmophores). In taxa with well-developed terminal appendices (e.g. tribes Thomsonieae, Areae, Arisaemateae) thickened connectives are usually absent. Odour production in genera with bisexual flowers is much less well understood. In Spathiphyllum the stigma plays this role (Vogel 1963, 1990, Williams & Dressler 1976) and in Anthurium it is probable that the thickened tepal apices are involved. Araceae are always protogynous and the female (stigma receptivity) and male (anther dehiscence) phases usually do not overlap, so that obligate out-

crossing seems to be the general rule. Some cases of self-pollination or apomixis are known or suspected (Amorphophallus bulbifer, Anthurium bakeri, Pinellia, some Arum). Manipulation of pollinator behaviour within the inflorescence may be the basis for many of the specialized features of the spadix, particularly in unisexual-flowered genera. Spathe constrictions may act as “skid zones” (Arum) or as “brooms” to eject pollinators from the female chamber after pollination (Philodendron). The various types of hairs, scales and warts found on the inner base surface of the spathe (e.g. Amorphophallus) or the wide range of staminodial or pistillodial structures on the spadix (e.g. tribe Areae, Bucephalandra) have less obvious functions. In Arum the filamentous pistillodes and staminodes are thought to exclude inappropriately large insect visitors while in Dieffenbachia the protein-rich staminodes have been shown to be food bodies for scarab beetles (Young 1986). It is conceivable that at least some of these structures are also osmophores which create odour gradients within the inflorescence itself. Certain structures, like the filamentous projections on the spadix appendix and spathe of Helicodiceros, mimic characteristics of animal corpses normally used by the pollinating insects for egg deposition or breeding.

POLLINATION BIOLOGY

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14 D I S P E R S A L

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The seeds of most Araceae do not remain viable for long. Those lacking endosperm and with large embryos cannot withstand dessication and the genera with fleshy testas are similarly vulnerable. In certain species, however, the seeds can withstand dessication for longer periods, e.g. Philodendron bipinnatifidum, which can remain viable for over 12 months (Mayo, pers.obs.). Seeds with a leathery testa, such as in Arum and Arisaema, also have longer viability. One consequence of short viability is that Araceae seeds are unlikely to survive long distance dispersal by natural vectors. This makes it unlikely that generic disjunctions over major ocean basins have resulted from long distance dispersal. Islands located far from continental areas tend to be very poor in native Araceae or lack them altogether, e.g. most islands of the Pacific. In Sauromatum venosum and Remusatia vivipara, however, long distance dispersal may well be the cause of their widely scattered Old World distribution patterns. Little is known of the viability of Sauromatum seeds, but its purple berries are very probably dispersed by birds, since in upland forests it often occurs as epiphyte. Remusatia vivipara, also epiphytic in upland forests, is almost certainly distributed mainly by means of its peculiar bulbils, which are burr-like with recurved scales. Another important generalization that can be made is that animal dispersal, and more specifically, ornithochory (bird dispersal), must be the dominant mode, due to the universality of berried fruits in Araceae. Reliable data on dispersal is very scarce, a recent exception being that of Barbara and David Snow (1988) for bird dispersal by blackbirds (Turdus merula) and robins (Erithacus rubecula) of Arum maculatum in England. Peckover (1985) observed in Papua New Guinea that captured birds of paradise (magnificent riflebird: Ptiloris magnificus) fed on fruits of Amorphophallus paeoniifolius and regurgitated seeds about four hours later. Glossy-mantled manucode bird of paradise (Manucodia atra) was also observed on an infructescence of Amorphophallus paeoniifolius and was presumed to be feeding rather than merely perching. If a similar delay between feeding and regurgitation also occurs in the wild a wider dispersal could thereby be achieved. Indigenous people (the local Batak population) have observed hornbills (Bucerotidae) eating berries of Amorphophallus titanum, and the bulbul bird (Pycnonotus zeylanicus) feeding on the berries of A. brooksii in Sumatra (Hetterscheid, pers. comm.). T. Lamb (pers. obs.) has added support to this evidence by his own observations of bulbul birds eating berries of A. lambii in

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Sabah. Shaw et al. (1985) reported Lewin’s honeyeater birds (Meliphaga lewinii) and regant bowerbird (Sericulus chrysocephalus) eating the ripe berries of Alocasia brisbanensis (as “A. macrorrhiza”) in eastern Australia. It is not known if the seeds are regurgitated, destroyed in the gizzard or stomach, or voided intact in the faeces. Circumstantial evidence for other genera points to birds (Anthurium) and mammals, including primates (Anchomanes, Philodendron) and bats (?Philodendron), as the commonest vectors. The tawny-capped euphonia (Euphonia anneae) was reported to feed heavily on fruits of Anthurium (Loiselle & Blake 1990). Wheelright et al. (1983) observed three different birds (resplendent quetzal: Pharomachrus mocinno, long-tailed manakin: Chiroxiphia linearis, common bush tanager: Chlorospingus ophthalmicus) feeding on three unidentified species of Anthurium. A fecal sample of the wood thrush (Hylocichla mustelina) was observed containing seeds of an unidentified species of Dieffenbachia by Blake & Loiselle (1992). The common palm civet (Paradoxurus hermaphroditus) is reported to disperse the seeds of Colocasia esculenta in Indonesia (Hambali 1980). Marks were left on the peduncle by the claws and teeth of this mammal and germinating seeds of two cultivars of C. esculenta were seen in its excreta. This is the only report seen that records the passage of viable aroid seeds through the digestive system of an animal. Hambali (1980) confirmed an earlier report by Leeuwen (1932) that seeds of Colocasia gigantea were dispersed by the common palm civet, and Hambali (1980) also stated that ripe fruits of Homalomena pendula are usually eaten by the same animal. The fruits of all three aroid species are odoriferous, which may attract the civet (Hambali 1980). Presentation of the fruits for dispersal is normally a rather sudden event. In Philodendron and Dieffenbachia the spathe falls off or splits at maturity to reveal the infructescence, whereas in genera with non-persistent spathes the berries remain inconspicuous during maturation and take on their bright colours in a final rapid flush. In Anthurium the berries are extruded from the perigone at maturity and in most species dangle by tiny threads of tepal epidermis. This mode of presentation has also been observed in Cyrtosperma by Hay (1988). The bright colours, sticky gelatinous mesocarp and mode of presentation in such species strongly suggests bird dispersal. In other, terrestrial or rupicolous species, e.g. Anthurium erskinei, the inconspicuous greenish berries merely fall into a heap onto the ground, sug-

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gesting some other kind of vector. Genera of subfamily Monsteroideae display the seeds by simultaneous abscission of the stylar region of each flower. The resulting compound mature fruit has the seeds embedded in sticky, often sweet and mucilaginous material. T. Croat (pers. comm.) has suggested that these fruits may be dispersed by monkeys. Philodendron subgen. Meconostigma has fruits with a similarly sweet, pineapple to mango flavour and some of its Brazilian vernacular names suggest mammal dispersal agents (“monkey’s banana” or “bat’s banana”). Black spider monkeys (Ateles paniscus) have been observed eating the infructescences of Philododendron goeldii (Bogner, pers. obs.). In Anthurium, Dieffenbachia, Nephthytis and other genera, the berries are often sufficiently distant from one another within the infructescence to be dispersed individually. Mature berries in Arum and Nephthytis have been observed to remain for a relatively long time awaiting dispersal. In contrast the

dense infructescences of Caladium, Colocasia, Philodendron, Syngonium, may disappear very soon after exposure. Dispersal by ants (myrmecochory) has been observed in Biarum, in which the strophiole of the seed is probably implicated. In the tropics it is likely that ant dispersal is involved in the occurrence of certain aroids in Amazonian root gardens (e.g. Philodendron megalophyllum (syn. P. myrmecophilum), Anthurium ernestii, A. gracile; see Ule 1905). Anthurium gracile is a characteristic plant of ant-gardens (Benzing in Huxley & Cutler 1991). T. Croat (pers. comm.) has observed ants dispersing seeds of Philodendron megalophyllum. Dispersal by water (hydrochory) is very probable in the helophytic genera Montrichardia and Typhonodorum, which have very large berries and floating seeds. Lagenandra, Cryptocoryne and Pistia are certainly water-dispersed and have much smaller fruits and floating seeds.

DISPERSAL

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15 G E O G R A P H Y

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Reviews of araceous geography have been published recently by Croat (1979), Grayum (1990), Hay (1992b) and Mayo (1993). Individual generic distributions are given in the maps, generic treatments and country lists (Appendix). The genera of Araceae are concentrated in the tropics of America, Southeast Asia and the Malay Archipelago (we include under this designation Malaysia, Indonesia, the Philippines, Papua New Guinea, Singapore and Brunei). Continental tropical Africa is the next richest region, followed in order of decreasing diversity by temperate Eurasia, southern Africa, Madagascar and the Seychelles, and North America (including northern Mexico). Australia has two endemic genera (Gymnostachys, Lazarum). The other native Australasian genera (in northern Australia) are essentially extensions of the tropical Asian and Malay Archipelago flora. The vast majority of genera are endemic to the major regions (as given above), but some extend further. Pistia is pantropical and Calla is circumboreal in the northern hemisphere, reaching as far as the subarctic zone in northern Scandinavia. Arisaema is the most widespread genus of more than one species. It is most diverse in south-west China, and extends west as far as Tanzania and Burundi, and east to eastern North America and northern Mexico. Amorphophallus, Remusatia, Rhaphidophora and Sauromatum are shared between tropical and subtropical Africa and Asia and the Malay Archipelago. Amorphophallus and Remusatia also occur in Madagascar and northern Australia. Pothos is found in Madagascar as well as in tropical Asia, Malay Archipelago, northern Australia and the western Pacific. Lysichiton and Symplocarpus occur in northeastern Asia as well as in temperate North America,

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Lysichiton reaching subarctic latitudes in Alaska. An unusual pattern is shown by the close taxonomic relationship of Peltandra (eastern North America) and Typhonodorum (Madagascar). Certainly the most interesting feature of araceous geography is the occurrence of three genera (Homalomena, Schismatoglottis and Spathiphyllum) with ranges disjunct between the Malay Archipelago or Melanesia and tropical America. These are all rainforest herbs for which long-distance dispersal by water or suitably far-ranging animal vectors is probably impossible. It is also difficult to imagine these genera being rafted by the southern Gondwanic route (Chile, Antarctica, Australia, New Zealand) or via the Bering Straits because of their intolerance of even subtropical conditions, much less temperate climate. Their patterns of diversity are dissimilar. While Homalomena and Schismatoglottis are most diverse in southeast Asia and Malesia, Spathiphyllum is richest in tropical America. No really plausible historical explanation has yet been proposed for these disjunctions, nor for those of other taxa with this kind of range, such as Sloanea (Elaeocarpaceae) and Heliconia (Heliconiaceae). Nevertheless, it is tempting to speculate that these patterns are the relicts of a once-continuous distribution in Gondwanaland during the Cretaceous period. A number of species which are important as food (Colocasia esculenta, Cyrtosperma merkusii, Xanthosoma sagittifolium complex) or ornamental plants (Alocasia macrorrhizos, Epipremnum pinnatum ‘Aureum’, Monstera spp., Philodendron spp., Syngonium spp., Zantedeschia aethiopica) have been widely dispersed throughout the tropics by man and have become naturalized as well. Some Typhonium species (Nicolson & Sivadasan 1981) have become widely dispersed and weedy in many parts of the tropics.

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16 U S E S

Bown (1988) gives an excellent general account of useful aroids to which the reader is referred for greater detail.

Food plants The most important food aroids are from tribes Colocasieae and Caladieae, i.e. Colocasia and Xanthosoma. The great majority of Araceae are poisonous when fresh and in almost all cases, edible species must be cooked or processed in some way before they can be used as food. Recent technical reviews of edible aroids, especially taro, are given by Wang (1983) and Chandra (1984). Colocasia esculenta, the taro plant, originated in tropical Asia and has been cultivated there for more than 2000 years. Its original geographical range is obscure, but Assam and Burma are likely possibilities. Today it is an important root crop in most humid tropical countries, especially in the Caribbean, Africa, Madagascar, Asia and the Pacific Islands. The tuberous stem is a rich starch source (13% to 29% by weight, depending on the cultivar), and the leaves of certain cultivars are widely eaten as a spinach. Taro tubers are also richer in protein than most other major starch crops and provide very nutritious food (Chandra 1984). When fresh, all parts of the plant are poisonous and must be cooked, roasted or heated in some way to become edible. The uncooked tissues contain an irritant toxin which can burn the skin (see chapter 11). Colocasia esculenta is known by a plethora of local vernacular names which correspond to the many different land races and cultivated varieties that have evolved by human selection. The species is most widely known today as taro, its name in the Pacific Islands. In some varieties the flesh of the tuberous stem is white or yellow while in others it is violet; the latter are sometimes preferred for their stronger flavour. Some cultivars are used as animal food, especially for pigs. Cultivars with tubers rich in mucilaginous material tend to be used for animal feed while those with a lower mucilage content are used for human consumption. The neotropical genus Xanthosoma also contains species which are very important food plants, particularly Xanthosoma sagittifolium. This and other species are widely cultivated, not only in tropical America but also in tropical Africa and Asia. As in Colocasia esculenta, both leaves and starch-rich tubers are used and can only be eaten after cooking. Plant remains from ancient Peruvian graves have been iden-

tified as Xanthosoma, showing that Xanthosoma must have been cultivated as a food plant in precolumbian times (Costantin & Bois 1910, Towle 1961). Xanthosoma sagittifolium is most widely known as cocoyam. There are many other names associated with the other species used for food, X. violaceum, X. atrovirens, X. mafaffa (used especially in Nigeria, Okeke 1992), X. brasiliense, X. caracu and X. robustum. These are all more-or-less closely related to X. sagittifolium, but much taxonomic confusion currently reigns in this group of taxa and needs to be clarified urgently (S. Thompson, pers. comm.). Alocasia macrorrhizos, the giant taro, was widely used for animal fodder in the nineteenth century and more rarely for human food. The stems have relatively little mucilaginous material and are eaten after roasting. They are said to be tasty when warm but irritant and unpleasant when cold. The stems may grow to as long as 5m and contain abundant latex. The starch-rich tubers of the elephant yam, Amorphophallus paeoniifolius (syn. A. campanulatus), are commonly used as food in tropical Asia, especially in India, where the species is widely cultivated. The tubers may reach 10 kg in weight and are eaten after roasting or boiling, like potatoes. In the Malay Peninsula the tubers of Amorphophallus prainii are also eaten after cooking, but must be sliced and soaked in water beforehand. Amorphophallus konjac (syn. A. rivieri), the konjac plant, is widely cultivated in Japan. The mannan-rich tubers (see chapter 11) are harvested after 1 to 3 years. They are boiled in water and then treated with lime milk to make a flour from which noodles (chira take) or cakes (chiroko) are prepared. A jelly (nama konjaku) is also made from the lime milk preparation and the gumlike juice can also be used to make glue. Konjac flour is traditionally prepared by slicing the raw tubers into 5mm thick pieces and then drying them in the sun for a week or so until the water content is reduced from about 90% to about 15%. The dried material is then pounded into a flour. Modern methods to extract konjac mannan involve washing the flour with 70% ethanol in a nylon filter or by acetylation. The quality of the flour differs according to the plant variety, the area in which it is grown and the method of preparation. The rigid gels prepared from the flour also vary according to the source of the raw material. The three main varieties of konjac cultivated in Japan are: ‘zairai-shu’ (traditional cultivar), ‘shina-shu’ (Chinese cultivar) and ‘bicchu-shu’. The thick, starch-rich rhizomes of Cyrtosperma merkusii (swamp taro) are used for food in Southeast Asia and the Pacific Islands where the plant is grown in

USES

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nutrient-poor sites unsuitable for the main starch staple, Colocasia esculenta. The tubers of various species of Dracontium have been recorded as a food source for Amerindians; preparation is by roasting. The fresh, ripe infructescences of Monstera deliciosa are eaten and used to flavour ice cream in Mexico; the taste recalls that of pineapple. The trichosclereids in the rind of the infructescence are troublesome and the wider use of this delicious fruit is hampered by lack of better cultivars. In Brazil, the infructescences of Philodendron bipinnatifidum are occasionally eaten by man (Crisci & Gancedo 1971); they have a mango-like flavour and a slightly irritant after-taste. The seeds of Typhonodorum lindleyanum and Montrichardia linifera are recorded as having been eaten by indigenous peoples of Madagascar and tropical South America respectively, after cooking or roasting. The seeds of Orontium aquaticum were eaten after drying and boiling by North American indigenous people. Theophrastos recorded the use of tubers of Arum italicum as a source of food starch in ancient Greece. In the Middle Ages, tubers of Arum maculatum were used as famine food, especially in England. North American Amerindians are known to have made a flour from the tubers of Arisaema triphyllum, but neither Arum nor Arisaema appear to be used for these purposes today.

family as a whole. The various medicinal uses of aroids described by Bown include external healing of stings, wounds, skin complaints and arthritis, expectorants and decongestants, contraceptives, parasite insecticides, anticancer agents, sedatives and hallucinogens. Acorus (Acoraceae) has a long history of use as a medicinal plant for problems of the digestive system and the scientific literature that exists on the subject is extensive (see Röst 1978, 1979a, b, Röst & Bos 1979) Toxic effects in Araceae are widely known and have received some attention from chemists (see chapter 11). The effects of the highly toxic Dieffenbachia are the best known and most completely studied. Araceae are used in arrow poisons and fish toxins (Bown 1988). Magical and ritual uses of aroids are known, but little studied. The use of Dieffenbachia and Caladium to ward off the “evil eye” is widespread in Brazil.

Fibres The roots of Heteropsis and Philodendron have been used for fibres in tropical South America. Heteropsis spruceana is still today an important source of twine and basket-weaving material in Brazilian Amazonia. The stems of Montrichardia linifera provide a fibre which is suitable for paper-making. Bown (1988) reports the use of fibres from species of Anthurium, Cercestis, and Gymnostachys anceps.

Medicinal, toxic and magical uses Ornamental uses The ethnobotany of the Araceae appears to be diverse and fascinating, judging from the many circumstantial reports which are scattered through the scientific literature. No comprehensive modern review exists and none is attempted here. However, valuable contributions have been made by Plowman (1969) and Croat (1994) for the New World and particularly by Bown (1988), who gave an excellent modern account for the

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Araceae are best known as ornamental plants and commercially are among the most important foliage ornamentals cultivated and sold for display. Their beautiful and unusually diverse leaf forms and textures form an essential part of any tropical plant display. In the tropics they are universally seen in private and public gardens. Further details are given in chapter 17.

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17 C U L T I VAT I O N

Crops The major crop species are Colocasia esculenta (taro), Xanthosoma sagittifolium (cocoyam), Cyrtosperma merkusii (swamp taro), Alocasia macrorrhizos (giant taro), Amorphophallus paeoniifolius (elephant yam) and Amorphophallus konjac (konjac). The cultivation methods of major aroid crop species are dealt with in some detail by Wang (1983) and Chandra (1984), which also give extensive bibliographies of the subject; Bown (1988) gives a very readable and informative account. Colocasia esculenta, Xanthosoma sagittifolium and Cyrtosperma merkusii are generally grown in humid conditions, although some varieties of Colocasia esculenta are also suitable for drier sites. Their preference for moisture makes these species especially suitable for humid tropical subsistence farming on sites unsuitable for other kinds of food crop. Amorphophallus paeoniifolius and A. konjac are grown in shade and on well-drained soils.

Ornamental Araceae Temperate genera Many hardy Araceae do best in shady and half shady sites with humus-rich soil and are therefore very suitable for temperate woodland gardens. This group includes species of Arisaema, Arisarum (with winter protection by leaf litter), Arum, Dracunculus and Pinellia. In cultivation they prefer leaf litter on the soil surface and this also protects the tubers or rhizomes from frost in winter. Growth starts in the spring or wet season and dormancy sets in during the autumn or dry season. They are suitable for growing with ferns and other shade-loving plants. Smaller species can also be grown in shady parts of rock gardens. Half hardy species can be grown in pots and overwintered in frost-free conditions. Pinellia ternata easily becomes weedy by vegetative propagation from the bulbils which form on the petioles. Biarum, Eminium and some Arum species thrive best in the rock garden, with stony, drier soils and sunny sites on walls or between rocks, but they require some protection in winter. Helicodiceros, a Mediterranean genus, is not quite hardy and the tubers need to be well protected during a temperate winter. One way to do this is to remove them from the soil and keep them frost-free and dry in sand. Other genera are best grown in wet places, like bog gardens, along streams, in pools or on the margins of a pond. These include Acorus (Acoraceae), Calla,

Orontium, Lysichiton, Peltandra and Symplocarpus. They do well in a nutrient-rich, loamy soil in a sunny or half shaded position. Calla prefers a more acid habitat in shaded or partially shaded sites. Orontium does best in full sun. Propagation is mostly by seed in Lysichiton, Orontium and Symplocarpus. Acorus and Calla are easily propagated by dividing the rhizomes. In Europe, naturally occurring Acorus calamus is triploid and cannot set seed. Division of Lysichiton, Orontium and Symplocarpus is best not attempted because their rhizomes are situated rather deeply and should not be disturbed. Tropical genera In tropical countries Araceae have an important role as ornamental plants. They are cultivated in public parks as well as in private gardens or as house plants. Under these favourable conditions they are easily grown and need little care beyond regular watering in regions with a drier climate. Their varied life forms provide magnificent climbers, bedding and large, sculptural terrestrial plants. Commonly cultivated aroids in tropical countries include species of the genera Aglaonema, Alocasia, Anthurium, Caladium, Colocasia, Dieffenbachia, Epipremnum, Homalomena, Monstera, Philodendron, Rhaphidophora, Schismatoglottis, Scindapsus, Spathiphyllum, Syngonium, Xanthosoma and Zantedeschia (especially at higher altitudes). Lasia spinosa, Typhonodorum lindleyanum, Montrichardia linifera and M. arborescens are often cultivated in or beside ponds where because of their large size they form impressive and unusual-looking stands. Pistia stratiotes is often grown in ponds and lakes. The geophytic (tuberous and rhizomatous) tropical Araceae are only seldom cultivated in gardens, perhaps because they require more care. In temperate regions, tropical genera are very important as house plants and are raised and sold on an industrial scale. Nearly every apartment or house in Europe and the Americas has at least one aroid as an ornamental plant. These, the best known and most widely grown aroids, tolerate low light levels (especially important for surviving the winter), dry air from central heating, irregular watering and general neglect, all of which makes them suitable for cultivation as house plants. A variety of Araceae is also grown in hydroculture in large offices and public buildings. Particularly well known house plant aroids belong to the genera Monstera (M. deliciosa), Philodendron (P. scandens, P. erubescens, P. bipinnatifidum and many other species and hybrids), Dieffenbachia (many cultivars of D. seguine and D. maculata), Aglaonema

CULTIVATION

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(many cultivars of A. commutatum, A. nitidum and other species), Syngonium (cultivars of S. podophyllum, often with variegated leaves), Zantedeschia (especially Z. aethiopica and hybrids), Spathiphyllum (especially S. floribundum, S. cannifolium and other species and hybrids), Epipremnum (E. pinnatum ‘ Aureum’, also known as Pothos aureus or Scindapsus aureus), Anthurium (particularly A. andraeanum and A. scherzerianum and cultivars) and Caladium (C. bicolor and cultivars). Many species of Cryptocoryne are prized by aquarists and species of Anubias are also widely grown as aquarium plants. Commercial nurseries that raise Araceae as house plants are common in many parts of the world, perhaps the most important being the USA and the Netherlands. Anthurium and Zantedeschia are the major sources of aroid cut flowers and are cultivated on a large scale. Anthurium andraeanum has a very large number of cultivars differing in spathe colour with various shades of red, pink, white and green and in spathe texture and the attitude of the spadix. Zantedeschia aethiopica and other species and their hybrids have a range of spathe colours which includes white, yellow, and various shades of red, and the spathes vary widely in size and shape. Plants for the cut flower market are cultivated in beds. Some are grown in gravel for hydroculture (hydroponic) or in ordinary, standardized or peat soils. Propagation is usually vegetative by means of cuttings but for certain cultivars tissue culture is also employed. Tropical aroids are mostly evergreen and are forest herbs, climbing hemiepiphytes or epiphytes in their natural habitats. Anthurium species grow best in conditions of high humidity and a minimum temperature of 18–20°C, with a soil mixture of sphagnum moss and fibrous peat. Dieffenbachia, Philodendron, Monstera and Spathiphyllum can be grown in standardized soil mixtures or specially made up composts composed of sterilized soil, treated leaf mould or bark chips, peat and sand with some balanced fertilizer. As a base, sandy loam with added peat and sand can also be used. Zantedeschia prefers a heavier, more loamy mixture. Climbers like Philodendron, Monstera and Epipremnum need a support on which to develop properly, such as a bamboo stick, moss-covered stick, roughened wooden billet or a suitable wall. Much more care is required to successfully grow seasonally dormant genera with tubers or rhizomes and these are consequently less commonly cultivated, e.g. Anchomanes, Amorphophallus, Caladium, Gonatopus, Dracontium, Remusatia, Synandrospadix, Taccarum, Theriophonum, Typhonium and Xanthosoma sect. Acontias. These plants must be kept dry during their dormant period and they are best grown in pots with a rather sandy compost in order to control soil humidity better. They require regular and abundant water during the growing period but watering must be reduced and very carefully controlled at the beginning and end of dormancy. While dormant the tubers are best kept dry

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in the old soil and left undisturbed, but they may also be removed from the soil and stored in a shady, humid place. Repotting should be carried out as the new roots begin to develop, i.e. near the end of the dormant period before the shoots appear. For example, in Europe the tubers of Caladium bicolor are repotted in the early part of the year (February, March) and grow during the summer until the end of August or September when they re-enter dormancy. They must then be kept dry in the old compost or removed entirely from the pot until the following year. Amorphophallus konjac, Colocasia esculenta and Sauromatum venosum can be grown outside from spring to autumn in central Europe. Propagation of these genera is by seed or daughter tubers. Cultivated tropical aquatics and helophytes include Cryptocoryne, Cyrtosperma, Dracontioides, Lagenandra and Urospatha. Most of these are grown only in a few botanic gardens. The most difficult species to grow, such as Jasarum and some species of Cryptocoryne, are from nutrient-poor black water rivers. These species require an acid soil and very soft water. Compost made from siliceous rock (granite or gneiss) or acid peat or leaf mould (especially from Fagus sylvatica) can be used. For less demanding species a mixture of loam, sand and peat or just sand and loam is sufficient. Some Araceae which do not become completely dormant need a drier period during which growth more-or-less ceases. At this time, however, they should not be allowed to dry out completely. Zantedeschia aethiopica, for example, needs a resting period in Europe between spring and summer (July) when it should be kept in rather dry and sunny conditions. Following this the plant should be repotted and watering gradually increased as the plant comes into more active growth. Other species (e.g. Z. albomaculata) have a completely dormant period without leaves. Some Alocasia species, especially those with beautifully coloured foliage (e.g. A. lowii, A. sanderiana), can be difficult to grow and are only suitable for “stove” conditions (high temperature, high humidity and shade). They also have a dormant period during which they should be watered only very sparingly, while never being allowed to dry out completely. Special attention must be given to the plants during this critical period. When the shoot starts into growth the plant should be repotted or replanted in a mixture of sphagnum moss and peat as used for Anthurium; chopped and partly decomposed wood or bark chips can also be used in the mixture. Alocasia macrorrhizos, A. odora and A. portei, on the other hand, are evergreens with no resting period. Propagation of Alocasia is by division of the stem, from the tuber-like stolons or by seed. True rheophytes like Aridarum, Bucephalandra, Hottarum and Piptospatha are rather difficult to maintain in cultivation because of their special requirements. They are best grown in a mixture of sphagnum moss and fibrous peat, if possible on rocks with a little humus; they require high temperature and humidity.

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18 C O N S E R VAT I O N

The main threat to the long term survival of many Araceae is the loss and reduction in quality of their natural habitats, especially in the rainforest regions of Asia, the Malay Archipelago, Africa and tropical America. Some Araceae are highly adapted to specific habitats and cannot survive in changed conditions; e.g. Chlorospatha and rheophytic species of tribe Schismatoglottideae. These taxa have proved difficult to cultivate and are unlikely to survive in the long term in botanic gardens. The same is also true of forest aquatics such as Jasarum (from blackwater rivers) and several species of Cryptocoryne. Currently, not one species of Araceae is listed in CITES (Convention on International Trade in Endangered Species). Formerly Alocasia zebrina and A. sanderiana from the Philippines were in Appendix 2. The overcollection of Araceae for commercial trade is unlikely to be a major cause of extinction, although some Cryptocoryne species have been collected in large quantities for the aquarium trade, much reducing their natural populations. Removal of tropical forest, however, eliminates most terrestrial, climbing and epiphytic species, many of which are shade-dependent. Other habitats are also affected. In the case of the aquatic genus Cryptocoryne, reduction in tree cover

results in faster flowing streams, which in turn gouge out the stream bed and deposit sediment on the plants along and near the banks. In such a situation the original plants soon die out. Fire is also very destructive, both in its immediate effects and in favouring the invasion of weedy plants which prevent the regeneration of the great majority of forest Araceae. Only a small minority of species can establish themselves in secondary regrowth or in open areas, e.g. some of the large species of Xanthosoma sect. Xanthosoma, Alocasia, Typhonium blumei, Zantedeschia aethiopica and certain races of Colocasia esculenta with a vigorous stoloniferous habit, which have become naturalized widely in the tropics. Some species of Arum have established themselves in parks and plantations, and Cryptocoryne spiralis is a rice field weed. Arisaema mooneyanum appears to have spread vigorously in open highland fields in Ethiopia, although it was probably originally a forest edge species. These, however, are exceptions; the great majority of Araceae disappear along with the forest habitat. Restricted endemic species are at particular risk. The determining ecological or historical factors for such narrow ranges remain completely unknown and this makes the future prospects for such species bleak.

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19 F O S S I L R E C O R D

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Gregor & Bogner (1984, 1989) have reviewed the fossil record of Araceae and their papers should be consulted for further details, particularly for the specialist literature. No fossils are known earlier than the Eocene which can definitely be ascribed to the Araceae, although Spathiphyllum-like pollen has been described from the Palaeocene and Limnobiophyllum scutatum from the late Cretaceous (see below).

Fossils consisting of different parts of the plant The genus Acorus (Acoraceae) is well documented from the Miocene of Spitzbergen by the fossil A. brachystachys, in which at least the leaf, inflorescence axis and spadix are connected (Heer 1870). However, the fossils from North America treated under this name by Lesquereux (1878) belong to the Coniferae. Acorites heeri (syn. Acorus heeri), from the Eocene of North America, is also close to Acorus. The fossils of this species consist of one single spadix and one spadix with a piece of its inflorescence axis (Crepet 1978). Kvaček (1995) has recently drawn attention to Limnobiophyllum Krassilov, a fossil of extraordinary interest in connection with the Araceae/Lemnaceae relationship. This fossil genus consists of free-floating, stoloniferous plants with one or two suborbicular to reniform leaves of different size, and simple as well as branched roots on a reduced main axis. There are (9)10–14 curved primary veins in the leaf blade, and these join the margin or run into the apex. The second order veins are reticulate between the primaries (higher order venation is absent in Spirodela and Lemna). There is no sign of the lateral pouches which occur in Spirodela and Lemna but aerenchyma and pigment cells are present. Two species are recognized: Limnobiophyllum scutatum (Dawson) Krassilov and L. expansum (Heer) Kvacek. Turion-like bodies are associated with L. scutatum. No seeds were found directly connected to these fossils, but numerous, isolated, ribbed seeds were associated with the same strata. These seeds resemble those of Spirodela, Lemna and similar fossil remains were described by Nikitin (1976) as Lemnospermum pistiforme Nikitin from Siberia. Generally, Limnobiophyllum most resembles Spirodela but is larger and the branched roots are more like those of Pistia. The fossils are less similar to Limnobium Rich. (Hydrocharitaceae), a relationship suggested by some authors. Limnobiophyllum scutatum is known from the latest Cretaceous to the Oligocene of western North America and the Palaeocene of east Asia. Limnobiophyllum expansum is known from the

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Miocene of Europe. Limnobiophyllum may be considered as standing between Pistia (Araceae) and Spirodela (Lemnaceae).

Leaf fossils Leaf fossils from North America have been assigned to the genera Peltandra, Philodendron and Pistia while South America leaf fossils have been placed in the genera Stenospermation (S. columbiense) and Caladiosoma. Caladiosoma miocenica from the Miocene of Trinidad belongs either to the genus Caladium or to Xanthosoma (Berry 1925). A North American Eocene leaf fossil with leaf venation typical of Peltandra was described by Hickey (1977) as P. primaeva. A remarkably large fossil leaf, also from the Eocene of North America, was described in great detail by Dilcher & Daghlian (1977) as Philodendron limnestis. However, Mayo (1991) suggested that the leaf venation indicates that this fossil would be better placed in the tribe Peltandreae, near Typhonodorum. Two leaf remains from Sumatra were described by R. Kräusel (1929): Araceophyllum engleri is probably from the Pliocene and resembles Pothos, while Araceophyllum tobleri is probably from the Upper Miocene and belongs to the tribe Monstereae. Two other species of Araceophyllum and the genus Anthuriophyllum are leaf fossils which only doubtfully belong to the Araceae. Weyland (1957) described Anthuriophyllum spectabile based on cuticular analysis from Tertiary brown coal deposits along the lower Rhine. More material showing the venation type is required to confirm that it belongs to the Araceae. The descriptions of Araceophyllum striatum (Weyland 1957) and A. tarnocense (Rásky 1964) are too incomplete to be accepted confidently as Araceae fossils. Several leaf impressions of the genus Pistia (P. claibornensis, P. corrugata, P. mazelii, P. nordenskiöldii, P. wilcoxensis) have been described, but some are not aroids at all, while others are only doubtfully so. P. wilcoxensis Berry (1916), for instance, from the Eocene of North America, has a venation type not found in the Araceae and should be excluded altogether. Chondrophyllum nordenskiöldii was transferred by Berry (1910) to Pistia but it represents dicotyledonous leaves. Depape & Gauthier (1953) described fossil leaves from the Eocene of Morocco and suggested them to be close to extant Pistia stratiotes. They also gave a survey of reputed Pistia leaf fossils, but none of these can be convincingly assigned to this genus or to any other aroid genus.

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Recently, three different leaf fossils have been described from the Eocene of the Grube Messel (Germany), although not formally named (Wilde 1989). One belongs to tribe Colocasieae and another to tribe Monstereae. The third resembles Aglaonema or Homalomena (i.e. subfamily Philodendroideae in the sense of Engler 1920b) in having a coriaceous leaf blade with parallel-pinnate venation and laticifers. Leaf-like fossils were described as Arisaema cretacea (Lesquereux 1892) and as Arisaema dubia (Hollick 1897) which were interpreted as portions of a spathe, but both are very doubtfully araceous and are best excluded from the family.

Fossil spadices and infructescences Acoropsis eximia (syn. Carex eximia, Acoropsis minor), from the Eocene Baltic Amber, represents a well-preserved infructescence of the tribe Monstereae (Bogner 1976c). This infructescence lacks tepals and is therefore not related to Acorus as suggested by Conwentz (1886). Fructus polyspermus from the European Miocene could belong to the Araceae but cannot be assigned with certainty to the family (Engelhardt 1877). Crepet (1978) considered the Eocene spadix Araceaeites fritelii from North America to be too incomplete to be included in the Araceae and the same must be said for the impressions of Lysichiton washingtonensis from the Miocene of western North America (Berry 1931). Lysichiton nevadensis is another impression from the Miocene of western North America and could be araceous (MacGinitie 1933). An infructescence from the Cretaceous of eastern North America was described as Arisaema mattewanense, but no internal structure was preserved and so this fossil remains very doubtfully ascribed (Hollick 1897). A spadix from the Miocene of North America was described as Orontium fossile (Cockerell 1926). Araceaeites parisiense, from the Palaeocene of France, was described as a spadix by Fritel (1910) and was compared by him to Spathiphyllum, but it is very incomplete and thus doubtful. Knowlton (1926) described a very questionable infructescence from the Miocene of North America as Arisaema hesperia. Apart from Acoropsis, Acorites and Acorus brachystachys, all these fossil spadices or infructescences, described under Araceaeites or an extant genus, are too incomplete to be ascribed with confidence to the Araceae and their taxonomic assignment must therefore remain highly dubious. Better preserved material of these fossil taxa is needed to clarify their position. Infructescences from the latest Eocene to earliest Oligocene from Egypt described as Teichosperma spadiciflorum (Renner 1907, Kräusel & Stromer 1924) in the Pandanaceae and new material studied by Tiffney & Wing (unpubl.) show that these fossils, with their smooth reniform seeds, belong to the genus Epipremnum of the Araceae (E. spadiciflorum).

Fossil fruits and seeds Nikitin described Acorus procalamus based on fruits and seeds from Quaternary deposits in the former Soviet Union and this has been well illustrated by Katz et al. (1965). Fossil seeds of Calla palustris and Acorus calamus are also described by Katz et al. (1965), while seeds typical of Pistia were described by Dorofeev (1963) as P. sibirica from the Oligocene of Siberia. Among the best-known aroid fossils are seeds of Monstereae and Lasioideae described from European brown coal deposits of Oligocene, Miocene and Pliocene age. These were formerly considered to belong to Epipremnum, but are today included within three genera: Epipremnites (type: E. ornatus) and Scindapsites (type: S. crassus) of tribe Monstereae, and Urospathites (type: U. dalgasii) of subfamily Lasioideae (Gregor & Bogner 1984, 1989). Keratosperma, with the single species K. allenbyense, is an Eocene fossil seed from Canada and belongs to the Lasioideae (CevallosFerriz & Stockey 1988). Scindapsites has reniform seeds with a smooth testa of variable thickness covered with scattered foveolae. There are distinctive outgrowths on each side of the raphe which are sometimes partly coherent. Epipremnites has curved seeds with the foveolae arranged in rows on an otherwise smooth testa; no outgrowths are present on the raphe. Urospathites has somewhat curved seeds and the testa is warty, spinose or tuberculate.

Fossil pollen Some fossil monosulcate pollen grains have been compared with Araceae, but this pollen type also occurs in many other families of monocots and dicots and these fossils cannot be accepted as araceous in the absence of associated floral remains. Mtchedlishvili and Shakhmundes (1973) described the pollen of four species in a new genus, Jugella (type: Jugella sibirica) from the Lower Cretaceous of the former Soviet Union. The authors compared them to Spathiphyllum pollen because of their striate exine, but Spathiphyllum has inaperturate pollen grains whereas in Jugella they are monocolpate which suggests that this genus should be excluded from the Araceae. Wodehouse (1933) described North American pollen from the Eocene as Peltandripites davisii and compared it with Peltandra, describing it as “...without germinal furrows or pores...”. However, his rough drawing shows something like a furrow and since the pollen of Peltandra is inaperturate this argument seems doubtful. Peltandripites dubius probably belongs to the Asteraceae (Sah & Dutta 1966, Thanikaimoni 1969). Biswas (1962) described fossil pollen from the Eocene of Assam as Colocasioideaepites, but these probably represent palm pollen grains (Nypa type). Monsteroideaepites eospathiphyllum is inadequately

FOSSIL RECORD

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described and figured and is therefore only doubtfully araceous (Muller 1981). Inaperturate pollen grains with a striate exine were reported by Graham (1976) from the Miocene of Veracruz, Mexico and assigned to Spathiphyllum. This is probably correct because other genera of Araceae with the same pollen type do not occur in Mexico. Pollen similar to that of Spathiphyllum was described from the Palaeocene of Colombia by Hammen & Garcia de Mutis (1966) as Ephedripites vanegensis. Pollenites tranquillus from the Eocene was compared with Acorus (Potonie 1934) and was later transferred to the form-genus Monocolpopollenites as M. tranquillus by Thomson & Pflug (1953); this pollen probably belongs to the Arecaceae (Nichols, Ames & Traverse 1973).

Fossils excluded from the Araceae Aracispermum Nikitin (type: A. canaliculatum), a seed fossil, may confidently be excluded from the Araceae, because it is characterized by a very large micropylar aperture, probably the site of an aril. All aroid seeds known have only a very small micropyle. Gregor & Bogner (unpubl.) and Mai (1995) consider that the seeds of Aracispermum canaliculatum and related species belong to the Zingiberaceae. Aracispermum johnstrupii belongs to the dicot genus Myrica (Myricaceae, Kirchheimer 1957). Aracispermum jugatum was transferred by Mai to Caricoidea (Cyperaceae) as C. jugata (Mai & Walther 1978, see also Friis 1985). Aracispermum hippuriformis was considered by Mai (in Mai & Walther 1978) as closely related to the genus Alpinia (Zingiberaceae). The fossil “strobilus” of Aracistrobus dravertii is in fact an old infructescence of Platanus (Platanaceae), showing the scars where the fruits were previously situated (Gregor & Bogner 1989). The “strobilus” remains of Araceites hungaricus are impressions of the shoot tips of a species of Pinus in which the needles are missing (Gregor & Bogner 1989). The fruits and seeds of Campylospermum hordwellensis (syn. Cyrtospermites hordwellensis) belong to the genus Visnea (Theaceae), according to Mai (1971). The monocotyledonous infructescence Viracarpon hexasper mum from the early Eocene Deccan Intertrappean beds of India has been critically reinvestigated by Bande & Awasthi (1986), based on new well preserved material, and a new reconstruction made. Four species of the genus Viracarpon (including Shuklanthus) have been described but only two (V. hexaspermum and V. elongatum) should be recognized (after Bande & Awasthi 1986). A relationship with Araceae, Cyclanthaceae and Pandanaceae has been suggested by various authors. New studies show that these fossils do not belong to the Araceae or Cyclanthaceae and also that they cannot be included in the Pandanaceae (B.C. Stone pers. comm.).

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Perhaps Viracarpon represents a genus of an extinct family. However, it would be helpful if its as yet unknown vegetative parts were found. The fruits of V. hexaspermum are sessile on an unbranched axis and are bractless. The fruit is hexangular with six locules around a central core which is slightly longer than the locules. Each locule contains a single seed. The outer wall of the ovary extends upwards in six perianth-like lobes, forming a cup-like structure (interpretation by Bande & Awasthi, but questionable) free at the apex and connate below with a vertical ridge running the entire length of the middle of the inner surface of each lobe. The inside of the lobes are densely covered with long hairs. The peduncle has monocotyledonous bundles. Porosia verrucosa (Hickey 1977) of the North American Late Cretaceous and Palaeocene has thick, reniform leaves and has been compared to Araceae, Lemnaceae and Hydrocharitaceae ; more complete material is necessary to resolve its relationships (Gregor & Bogner 1989, Crane et al. 1990, Kvaček 1995). Fossil flowers, probably of Eocene age, have been described from the Indian Deccan Intertrappean series as Sahnipushpam shuklai (syn. Sahnipushpam glandulosum) and were assigned to the Araceae by Prakash & Jain (1964). However, following further examination of the original material, Gregor & Bogner (1989) have excluded this fossil from the Araceae. The flowers were always single and abscissed individually (unknown in Araceae). The pollen and the structure of the ovary locules and style are different from those found in Araceae. Although the true taxonomic position of Sahnipushpam is still unclear, it may nevertheless be excluded from the Araceae without reservation. Ettinghausen (1870) described a fossil aerial root from the Miocene of Croatia as Aronium extinctum and compared it with Anthurium, but this fossil is best regarded as “incertae sedis”. Cyclanthodendron sahnii from the Deccan Intertrappean beds, considered to be of early Eocene age, was first described as Palmoxylon sahnii. Later a separate genus, Cyclanthodendron, was established, mainly on the basis of the compound vascular bundles (absent in Arecaceae) and assigned to the Cyclanthaceae. This fossil has also been compared with the Pandanaceae and Araceae, in which compound bundles were observed by French & Tomlinson (1986). New material from the same locality showed that this fossil is very close to the Strelitziaceae (Biradar & Bonde 1990). Some other fossils which have been thought to belong to the Araceae have been transferred in previous palaeobotanical literature to other families: Arecites trabucci (Arecaceae), Aronites dubius (Coniferae, partly indeterminable), Aroites tallyanus (Coniferae), Pistites loriformis (Cycadaceae), Aroides stutterdii (Stichoporella in the Dasycladaceae), Pothocites grantonii (Calamites s.l. in the Calamariaceae). Aroides crassispatha is a very doubtfully assigned fossil from the Permian.

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20 P H Y L O G E N E T I C R E L AT I O N S H I P S W I T H I N T H E M O N O C OT Y L E D O N S The phylogenetic relationships of Araceae to other monocots have never been clear. The fossil record provides no clues and we have to rely on the comparative study of modern taxa. Some influential earlier authorities, such as Engler & Gilg (1919) and Wettstein (1935), grouped the Araceae together with the Arecaceae and Cyclanthaceae. This view was based on the common tendency in these families to floral reduction, “spathe” development, the condensation of the inflorescence into one or several spadices and an associated tendency to pseudanthial inflorescences. For similar reasons, the Araceae have also been associated in the past with Pandanaceae and Typhales. The link with Typhales was also suggested because it had been observed that Sparganium (Sparganiaceae) and Acorus (Acoraceae) are infected by the same rust, Uromyces sparganii. Whether there is a close phylogenetic relationship between Acorus and Sparganium remains unclear, but since Grayum’s (1987) widely accepted removal of Acorus from the Araceae, the case for an Araceae-Typhales relationship has been seriously, if not fatally, weakened. As regards the Arecaceae, Cyclanthaceae and Pandanaceae, their floral and vegetative structure differ so strikingly from the Araceae that any resemblances must surely be due to parallel evolution rather than sister group relationship. The Dioscoreales (in the sense of Dahlgren et al. 1985, consisting of Dioscoreaceae, Petermanniaceae, Smilacaceae, Stemonaceae, Taccaceae, Trichopodaceae, Trilliaceae) is a group sometimes linked to the Araceae because of a superficial similarity in leaf form and venation. The differences, however, are also impressive. The Dioscoreales differ from the Araceae in the usually twining habit, different type of reticulate venation (with numerous densely arranged, scalariform cross veins connecting the major veins), inferior ovaries, usually winged, capsular fruit and usually winged seeds. Furthermore, the Dioscoreales seem unlikely to be monophyletic, which further complicates an assessment of their relationship to the Araceae. The concept of an origin from a common ancestor of the Liliiflorae (sensu Dahlgren et al. 1985), or even from within this taxon has been widely held (e.g. Hallier 1912, Bessey 1915, Hutchinson 1934, 1959, Novak 1954, Kimura 1956, Takhtajan 1959). Hutchinson (1934) went so far as to suggest that the Araceae could be derived directly “... from the stock of the tribe Aspidistreae of Liliaceae in which the flowers are arranged in dense spikes (Tupistra, Rohdea, Gonioscypha).” This latter similarity must surely be, however, the result of parallel evolution. Lying behind the Liliiflorean origin hypothesis was the very general view that this group

represents the “type” or “Bauplan” of the monocots, or to put it another way, that the Liliiflorae has a basal phylogenetic position within the monocots as a whole. Engler (1920b) recognized clear evidence of the “monocot type” in the floral structure of some genera of Araceae (e.g. tribe Potheae) and consequently he considered these to be primitive aroids. He even stated that his subfamily Pothoideae differed from the Liliaceae essentially only in having a fleshy outer seed integument. In his classification however, Engler (e.g. Engler & Gilg 1919) did not associate Araceae and Liliaceae closely and it is reasonable to suppose that he saw the similarities between them in terms of shared primitive characters (plesiomorphies); i.e. Araceae must have arisen by independent evolution from primitive monocot forms. Characters which may be considered primitive in Araceae by outgroup comparison to Liliiflorae are: absence of laticifers, inconspicuous spathe, bisexual flowers with a perigone of two whorls of 3 free tepals and two whorls of 3 free stamens, basifixed anthers on more-or-less elongated filaments, thecae dehiscing by longitudinal slit, monosulcate pollen, syncarpous, 3-locular, superior ovary, several anatropous ovules per ovary locule, axile placentation, and presence of endosperm in the mature seed. However, many of these characters can be regarded as primitive in Araceae by comparison with the monocots as a whole, which weakens the concept of Liliiflorae as a sister group for the Araceae. A further problem is that the Liliiflorae are almost certainly paraphyletic or polyphyletic. Recent molecular studies (e.g. Duvall et al. 1993, Chase et al. 1995) have placed the genus Pleea (Melanthiaceae) near the base of the monocot clade, together with the Araceae, Alismatiflorae and Acorus. These studies also indicated that Lemnaceae belong within the Araceae, a result recently further supported by a comprehensive analysis of cpDNA characters of Araceae and Lemnaceae (French et al. 1995). In the following discussion, therefore, the possible sister group relationships of the Araceae are considered only in relation to the Acoraceae, the Alismatiflorae and the Lemnaceae, together with an overview of the proposed primitive character states of Araceae.

Relationships with the Acoraceae Acorus has long been considered a member of Araceae, but recently Grayum (1987, 1990) has presented a convincing case for its removal to a separate

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family, a view which has been widely accepted. The long list of significant characters by which Acoraceae and Araceae are distinguished (Table 3) also strongly suggests that they are not even sister taxa.

Table 4. Proposed synapomorphies and plesiomorphies of the Araceae. POSSIBLE SYNAPOMORPHIES • Lack of vessels in stems and leaves (stem vessels of climbing Araceae derived?)

Table 3. Characters separating Acoraceae from Araceae.

• Presence of tannins

• Ethereal oil cells

• Distichous leaves

• Lack of raphides

• Presence of spathe

• Unifacial ensiform leaves

• Presence of spadix

• Unique pattern of bud trace insertion

• Flowers lacking floral bracts

• Separate vascular systems in peduncle

• Protogyny

• Introrse anthers (in Araceae known only in Zamioculcas)

• Anthers extrorse

• Stellate endothecial thickenings

• Tapetum periplasmodial

• Tapetal cells with 2–4 nuclei

• Tapetal cells 1-nucleate

• Secretory anther tapetum

• Placentation basal (equivocal)

• Exclusively axile vascular supply to placentae • Location and structure of ovular trichomes

• Helobial endosperm development with cellular development in micropylar chamber

• Presence of perisperm

• Embryogeny caryophyllad or solanad

• Dicot-type cellular endosperm development

• Berries (dehiscent in Lagenandra)

• P2cs sieve tube plastids

• Pollen 2-nucleate when shed

• Lack of endosperm (equivocal)

The presence of ethereal oil cells, absence of raphides, presence of secretory anther tapetum, presence of perisperm and dicot-like cellular endosperm development are characters which could suggest a link to monocot-like dicot families such as Piperaceae and Aristolochiaceae. The molecular studies of Duvall et al. (1993) and Chase et al. (1995) now indicate that Acorus could be considered the most primitive living monocot taxon.

POSSIBLE PLESIOMORPHIES • Presence of raphides • Absence of silica • Absence of tricin • Absence of ethereal oils • Absence of laticifers • Leaves bifacial with petiole and lamina • Flowers actinomorphic • Tepals 4–6 in two whorls of 2 or 3

Primitive characters of the Araceae

• Anthers basifixed • Stamen filaments ± elongate

The removal of Acorus from Araceae has made it easier to reassess the hypothetical primitive character set for the family. The basic evolutionary trend within Araceae has always been viewed as the evolution of unisexual flowers in monoecious pseudanthoid inflorescences from simpler inflorescences with bisexual flowers and a lesser degree of pseudanthial development. This hypothesis is supported by our cladistic study (Mayo et al., in prep.). Once this trend is accepted, the evolutionary polarity of certain other important characters can be defined by correlation. Thus most bisexual genera lack laticifers and have aperturate pollen. Based on our cladistic analyses and a reassessment of some other characters not used in our analysis but discussed by Dahlgren and colleagues (Dahlgren & Clifford 1982, Dahlgren et al. 1985), the primitive character states shown in Table 4 are proposed (see Grayum 1990 for a similar but not identical list).

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• Pollen mother cell cytokinesis successive • Pollen monosulcate (or at least aperturate) • Ovary syncarpous • Ovary superior • Ovules crassinucellate • Perisperm absent

Relationships with Alismatiflorae Dahlgren and co-workers (Dahlgren & Clifford 1982, Dahlgren & Rasmussen 1983, Dahlgren, Clifford & Yeo 1985) argued strongly for a sister relationship of Ariflorae (Araceae, Lemnaceae) to Alismatiflorae, pointing out many important differences between Araceae and Arecaceae. Grayum (1984, 1990, 1991b, 1992a)

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agreed with Dahlgren’s views; his re-interpretation of araceous endosperm development as helobial further emphasized a relationship with Alismatiflorae. The characters that link Alismatiflorae to Araceae, based on our assessments of primitive character states in Araceae, are given in Table 5. Among the clearest of these are periplasmodial tapetum, 1-nucleate tapetal cells and caryophyllad embryogeny, but data for these are based on a very patchy coverage of the two taxa. Many of the other characters could be interpreted as plesiomorphies for the monocots as a whole.

Table 5. Proposed synapomorphies (or plesiomorphies) of the Alismatiflorae and Araceae.

Certain other characters adduced by Dahlgren et al. (1985) to support the Alismatiflorean link must be ruled out in the light of our cladistic study, since they emerge as derived within Araceae – presence of intravaginal squamules, presence of laticifers and presence of 3nucleate pollen grains. The sister group relationship with Alismatiflorae remains equivocal on this evidence but is a hypothesis which continues to merit serious consideration, and appears stronger than any other yet proposed. The molecular studies of Duvall et al. (1993) and Chase et al. (1995) are not completely consistent in grouping Araceae and Alismatiflorae together, but as with the morphological and anatomical data, it also remains one of the strongest available hypotheses based on molecular evidence.

• Vessels lacking in stems and leaves (stem vessels probably derived in climbing Araceae)

Relationships with Lemnaceae

• Leaves distichous (primitive in Araceae, present in Scheuchzeriaceae, Potamogetonaceae)

The most widely accepted sister group relationship of Araceae is that with the Lemnaceae, but this is by no means universally accepted and has been strongly challenged recently by Grayum (1984, 1990, 1991b, 1992a). The Lemnaceae are considered by most authors to be at least closely related to Araceae, and to have evolved from them by neotenous reduction of leaf and inflorescence. This view is based, among other characters, on the similarity between seedlings of Pistia and Spirodela, embryological characters (Maheshwari 1956, 1958, Maheshwari & Khanna 1956) and the putative homology between the aroid spathe and the “spathe” of Spirodela and Lemna. Other similarities adduced are the presence of grooved raphides and operculate seeds in Pistia and Lemnaceae. An important difference, however, is that the pollen of Lemnaceae is ulcerate and spinose while in Pistia it is inaperturate and plicate (or ± striate); ulcerate pollen is unknown in Araceae (for Limnobiophyllum see chapter 19). The three hypotheses considered in more detail here are shown in Figure 9. This illustrates the need to assess possible synapomorphies (shown as thicker lines) for the following sister group pairs: Lemnaceae-Araceae, Lemnaceae-Pistia, and Lemnaceae-monoecious Araceae. A fourth possibility is a trichotomous monophyletic group composed of Lemnaceae-Araceae- Alismatiflorae. In the following discussion, character states in Spirodela are regarded as representing the primitive condition in Lemnaceae.

• Petiolate and laminar leaves with sheathing base (also in Liliiflorae) • Presence of spadix (Aponogetonaceae, Juncaginaceae, Potamogetonaceae, Zosteraceae) • Stamen filaments distinct (plesiomorphy in monocots) • Anthers terminal (plesiomorphy in monocots) • Anthers extrorse (also in Liliiflorae, doubtful in Lemnaceae) • Tapetum periplasmodial (unusual in monocots) • Tapetal cells 1-nucleate (unusual in monocots) • Ovules several per locule (plesiomorphy in monocots?) • Ovules anatropous (plesiomorphy in monocots) • Placentation basal (equivocal as primitive state in Araceae) • Ovule with nucellar cap (occurs elsewhere in monocots) • Embryo sac of Polygonum type (plesiomorphy in monocots) • Endosperm development with helobial type of chalazal chamber (helobial endosperm development occurs elsewhere in monocots, e.g. Liliiflorae) • Embryogeny of caryophyllad type (onagrad and asterad types commonest in monocots) • ?Embryo chlorophyllous (data scarce) • ?Endosperm absent (equivocal as primitive state in Araceae)

Lemnaceae-Araceae as sister groups

• ?Unique type of seedling development (linked to previous character; a similarity between Scheuchzeriaceae, Aponogetonaceae and Araceae with endospermless seeds; equivocal as primitive state in Araceae)

Possible synapomorphies are listed in Table 6. The strongest of these characters is that of endosperm development, which typically combines cellular development in the micropylar chamber with a haustorial chalazal chamber containing a single hypertrophied nucleus (Grayum 1991b).

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Lemnaceae

Table 7. Possible synapomorphies of the Lemnaceae and monoecious Araceae.

Araceae

• Flowers unisexual (controversial interpretation in Lemnaceae)

Alismatiflorae

• Pollen 3-nucleate • Exine spinose (but pollen aperturate and ulcerate in Lemnaceae)

Lemnaceae

• Asterad embryogeny (primitively caryophyllad in Araceae and Alismatiflorae)

monoecious Araceae

• Endosperm starchy (no endosperm in Alismatiflorae)

other Araceae

• Base chromosome number x=10 (this number occurs in monoecious Araceae but not in more primitive bisexual genera)

Alismatiflorae

Lemnaceae Pistia other Araceae Figure 9. Alternative sister groups of the Lemnaceae

Table 6. Possible synapomorphies of the Lemnaceae and Araceae. • Flavonols present (rare in Alismatiflorae, absent in Pistia) • Raphides present (widespread in monocots, absent in Alismatiflorae and monocot-like dicots) • Spathe present (controversial interpretation in Lemnaceae, bract subtending inflorescence widespread in monocots) • Outer integument overtopping inner (?distribution in other monocots) • Ovary syncarpous (controversial interpretation in Lemnaceae, possible symplesiomorphy with Liliiflorae) • Helobial endosperm development with cellular development in micropylar chamber (unique in monocots) • Endosperm present in ripe seeds (possibly derived in Araceae, general in monocots)

Lemnaceae-monoecious Araceae as sister groups Possible synapomorphies, additional to those linking Lemnaceae to Araceae as a whole, are listed in Table 7. A number of lemnaceous characters which do not fit with primitive Araceae agree better with the

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advanced subfamily Aroideae. Unisexual flowers, 3nucleate and spinose pollen and asterad embryogeny are all more characteristic of advanced rather than primitive Araceae. A base chromosome number of x=10 occurs in monoecious Araceae but not in primitive ones. However the aperturate pollen of Lemnaceae is ulcerate and this type is unknown in Araceae. Almost all monoecious Araceae have inaperturate pollen and so this character is problematic were Lemnaceae to be inserted well within the monoecious aroid clade (subfamily Aroideae). In a sister group relationship, however, there is less difficulty. The tribe Zamioculcadeae, which is basal in subfamily Aroideae, has aperturate pollen (and so, occasionally, does tribe Stylochaetoneae – also near basal), and so a position for Lemnaceae near the base of the Aroideae clade is at least conceivable. The molecular data, on the other hand, suggests that Lemnaceae are embedded well within the Aroideae clade (French et al. 1995). On this hypothesis, some further explanation will be required for the evolution of the pollen characters.

Lemnaceae-Pistia as sister groups The major character conflicts and the possible synapomorphies of these two taxa are shown in Table 8. The six character differences are a major obstacle to linking Lemnaceae and Pistia in a sister group relationship, particularly the pollen and nucellar characters. The latter is also problematic in grouping Lemnaceae within Araceae since although few genera of Araceae have been studied, none are yet known certainly to have crassinucellate ovules. On the other hand, the joint possession of operculate seeds and practically identical seedling development appear to be strong evidence for a close relationship. It is nevertheless tempting to regard these as convergences resulting from their highly specialized niche as floating aquatics. The molecular studies

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of French et al. (1995) place the two taxa in widely separate positions within subfamily Aroideae (as defined in our classification). This hypothesis of sister group relationship thus remains very doubtful. The Lemnaceae are best considered for the present as a derived offshoot of Araceae with a sister group relationship to the major advanced clade represented by subfamily Aroideae. There is a great need for more comparative embryological studies of Araceae which could help to shed further light on this problem.

Are the Araceae paraphyletic? The weight of evidence, especially the new data from molecular studies, now strongly favours the inclusion of Lemnaceae within Araceae as merely one subclade of an overall monophyletic group. We have not, however, included Lemnaceae in our classification nor in our taxonomic treatment. Our reasons are pragmatic. The molecular evidence emerged at a very late stage in the preparation of this book. In addition, Landolt (1986) and Landolt & Kandeler (1987) have provided a recent comprehensive taxonomic treatment of Lemnaceae which we certainly could not improve upon. It would also be desirable to carry out a combined analysis of molecular and morphological data to establish the position of Lemnaceae more precisely within Araceae. With these qualifications in mind it is nonetheless important to emphasize that the family Araceae is almost certainly paraphyletic, while the order Arales (Araceae including Lemnaceae) is very probably monophyletic.

Table 8. Character conflicts and possible synapomorphies of the Lemnaceae and Pistia. CHARACTER CONFLICTS Lemnaceae • Stamens free • Pollen ulcerate • Ovules crassinucellate • Exine spinose • Apiose present • Tannins absent

• • • • • •

Pistia Stamens connate Pollen inaperturate Ovules tenuinucellate Exine plicate Apiose absent Tannins present

POSSIBLE SYNAPOMORPHIES • Grooved raphides • Floating aquatic (rare in monocots) • Stolons arising in lateral pouch • Operculate seeds (also in Commelinales and Zingiberales) • Seedling development: median cotyledonar lobe, no primary root, operculum persistent over root pole, first adventitious roots arising around root pole. • Fruit not a berry • Laticifers absent • Root hairs absent

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Previous work on the classification of Araceae has reached a reasonable consensus on the circumscription of the tribes and subtribes. The major difficulties revolve around the subfamily concepts. These were introduced by Engler (1876b) and have been found very useful by subsequent authors. It is much easier to think taxonomically in terms of 8 subfamilies rather than 30 tribes. The reluctance of modern authors to abandon the subfamily concept, despite the obviously superior taxonomic quality of the tribal groups, is shown by their constant use in aroid literature. A stage has now been reached in which confusion abounds. Current classifications (Grayum 1990, Bogner & Nicolson 1991, Hay & Mabberley 1991) differ radically in the composition of several subfamilies and it is no longer possible to speak, for example, of subfamilies Aroideae, Lasioideae or Philodendroideae without citing the author of the system being used. We therefore carried out a series of cladistic analyses using all genera as terminal taxa, and without assuming any higher groupings at the outset. Our classification (chapter 23) is based on the results. The details of these analyses will be given in a separate publication (Mayo et al., in prep.). The purpose of this chapter is to explain the phylogenetic basis of the classification we have adopted. The cladistic study used 63 morphological and anatomical characters and 109 taxa (including 3 outgroups). The character data was gleaned from the literature, our own studies of herbarium, spirit and living material and from unpublished observations generously supplied by various colleagues (see acknowledgements). The main outgroups used were Tofieldia (Melanthiaceae-Liliiflorae) and Scheuchzeria (Scheuchzeriaceae-Alismatiflorae), representing Liliiflorean versus Alismatiflorean sister group relationships respectively. Figure 2 shows one of the 100 equally parsimonious trees found in an analysis using Tofieldia as the outgroup. In producing a classification from the cladistic results we have taken the view that groups which emerged consistently from the analysis should be considered seriously as named taxa in the classification. In cases where traditionally recognized groups failed to emerge in the cladogram it was necessary to decide whether this was a sufficient reason to reject them in the classification. In the case of tribe Zomicarpeae, for example, we have recognized the tribe despite the fact that it usually failed to emerge as a monophyletic group. We justify this because they possess some “good” characters in common (e.g.

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anastomosing laticifers, female zone of spadix fused to spathe, etc.) and especially because this taxon emerged as a monophyletic group in the molecular analysis of French et al. (1995). In contrast, no support was found for the concept of subfamily Philodendroideae in the sense of Engler (1920b), Grayum (1990) or Bogner & Nicolson (1991). Most currently accepted tribal and subtribal groups stood up well to analysis, thus confirming the prevailing view of aroid taxonomists that they are mostly monophyletic (or natural) groups. The tribal groups held together even when chromosome base number, generally considered an important tribal character, was excluded from the analysis. The results of a comprehensive cpDNA study by French et al. (1995) give a strikingly similar result to ours in the basic structure of the cladogram. We have therefore incorporated into our classification the strongest features of these two independent analyses and we have not hesitated to modify our major taxon concepts in the light of their results. We hope this will help to achieve long term stability in aroid classification. With few exceptions, the suprageneric groups recognized and named in our classification represent monophyletic taxon concepts resulting from the cladistic analyses. We have deliberately chosen not to overemphasize the internal topologies of these clades, preferring to concentrate on the “main skeleton”. New studies and analyses will be needed to give a more reliable picture of the internal phylogeny of the various tribes and subfamilies recognized here. Figure 11 shows the cladistic relationships of the seven subfamilies we recognize, based on a consensus tree of 100 equally parsimonious trees, one of which is shown in Figure 10.

A. Major Group Proto-Araceae This clade consists of subfamilies Gymnostachydoideae and Orontioideae and is defined by the following characters – medium sized pollen, condensed, non corm-like thickened stem, subterranean stem, usually unilocular ovaries and locules with 1–2 ovules. These are rather weak and highly homoplasious synapomorphies, which suggests that the group may be paraphyletic rather than monophyletic. It is noteworthy, however, that French et al. (1995) independently and consistently found the same group using cpDNA data.

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1. Subfamily Gymnostachydoideae Among other peculiar characters, Gymnostachys has linear leaves with parallel venation and a flowering shoot of unique structure. We therefore prefer to keep it in its own monospecific subfamily (following Bogner & Nicolson 1991).

clade. The tribe Spathiphylleae failed to group consistently with these three tribes but does so in the cpDNA analysis of French et al. (1995). Our subfamily Monsteroideae thus differs from Engler’s only by the addition of Anadendrum and Heteropsis.

5. Subfamily Lasioideae 2. Subfamily Orontioideae This group corresponds to tribe Orontieae of previous classifications. The synapomorphies are:– leaf blade expanded not linear, anatropous or hemianatropous ovules, endosperm sparse to absent, base chromosome number x=13.

B. Major Group True Araceae This is a previously unrecognized group and is supported by the following synapomorphies:– conspicuous or flag-like spathe, major internode of inflorescence between spathe and next leaf below, continuation shoot in axil of penultimate leaf before spathe, leaf blade expanded not linear, basal or nearbasal placentation. These characters are strong and less homoplasious which suggests that the group is indeed very probably monophyletic.

3. Subfamily Pothoideae Tribe Potheae is a consistent group defined by the following synapomorphies:– monopodial shoot architecture, lack of endosperm, chromosome base number x=12. The genus Anthurium failed to group consistently either with tribe Potheae or any other group in our analysis. French et al. (1995), however, found that Anthurium consistently grouped with tribe Potheae and we have adopted this to form the subfamily Pothoideae. Figure 10 is an example of one family of cladograms which show the two taxa as sister groups, and in this case they share a single synapomorphy – fine leaf venation with secondary and tertiary veins forming mostly cross veins to primaries; the plesiomorphic condition in Anthurium was assumed to be that shown in Anthurium sect. Digitinervium.

4. Subfamily Monsteroideae In our analysis the genera of the tribes Monstereae, Heteropsideae and Anadendreae form a single consistent clade. The synapomorphies are:– spathe undifferentiated into tube and lamina and soon deciduous or marcescent with distinct basal abscission, perigone connate. The latter character occurs only in Anadendrum, the perigone being lost further up the

Our subfamily Lasioideae corresponds to tribe Lasieae of earlier systems and is a very stable and consistent clade. The synapomorphies are:– monosulcate pollen (derived by reversal from the inaperturate state), absence of pollen starch, basal ribs of primary veins very well developed, dracontioid leaf margin development, spadix with basipetal flowering sequence, anthers dehiscing by oblique pore-like slits and very often unilocular ovaries. Monosulcate pollen is normally regarded as primitive in the family and it is possible that its occurrence as a reversal here may be an artefact contingent on the topology of this particular cladogram (see discussion under subfamily Aroideae).

6. Subfamily Calloideae The genus Calla consistently emerges as a single clade and was usually among the basal branches in our analysis. French et al. (1995) also found that Calla emerged consistently as an independent clade, but in their analysis it occurred further up the tree. The autapomorphies are:– perigone absent, pollen diaperturate, pollen globose, laticifers simple, petiole sheath long-ligulate, ovary unilocular, chromosome base number x=18. Calla seems to be highly autapomorphic and its sister relationships remain obscure.

7. Subfamily Aroideae The most striking feature of the analysis is the large clade which contains all the monoecious genera. This group corresponds to Schott’s “Diclines” (Schott 1860) and is not recognized in the classifications of Engler (1876b, 1920b), Grayum (1990), Bogner & Nicolson (1991) and Hay & Mabberley (1991), which all embody the idea that monoeicy and associated advanced spathe and spadix characters must have evolved several times from bisexual-flowered ancestors. Strong support for our concept comes from the DNA work of French et al. (1995) which also produces a single clade for all monoecious taxa. On the basis of both studies, we therefore feel confident in advocating the taxonomic recognition of this group as subfamily Aroideae since it represents a major advance and simplification in our understanding of aroid phylogeny.

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Tofieldia ACORACEAE GYMNOSTACHYDOIDEAE ORONTIOIDEAE CALLOIDEAE ANTHURIEAE POTHEAE

SPATHIPHYLLEAE ANADENDREAE HETEROPSIDEAE

MONSTEREAE

LASIOIDEAE

ZAMIOCULCADEAE STYLOCHAETONEAE

SPATHICARPEAE

Dieffenbachia AGLAONEMATEAE ZANTEDESCHIEAE PHILODENDREAE HOMALOMENEAE ANUBIADEAE Bognera

Figure 10. Cladogram of the genera of Araceae. One of 100 equally parsimonious trees (Mayo et al., in prep.). Terminal taxa are shown as subfamilies or tribes when monogeneric or when all genera emerged consistently as a single clade. Generic names are given where the genera of recognised tribes failed to form monophyletic groups in the analysis (see chapter 23 for a full synopsis of the classification).

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MONTRICHARDIEAE CULCASIEAE THOMSONIEAE

NEPHTHYTIDEAE CALLOPSIDEAE Zomicarpella Ulearum AROPHYTEAE ARISAREAE AMBROSINEAE PISTIEAE Filarum Pinellia Zomicarpa Arisaema

AREAE

CRYPTOCORYNEAE

SCHISMATOGLOTTIDEAE

PELTANDREAE Protarum Alocasia Steudnera Colocasia Remusatia Gonatanthus Scaphispatha Ariopsis Caladium Hapaline Jasarum Syngonium Xanthosoma Chlorospatha

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Major Groups

PROTO-ARACEAE

Subfamilies GYMNOSTACHYDOIDEAE 1 genus ORONTIOIDEAE 3 genera POTHOIDEAE 4 genera MONSTEROIDEAE 12 genera

TRUE ARACEAE

Flowers bisexual

LASIOIDEAE 10 genera CALLOIDEAE 1 genus AROIDEAE 74 genera

Flowers unisexual

Figure 11. Cladogram of the subfamilies of the Araceae, based on a consensus tree (Mayo et al., in prep.), showing the Major Groups and subfamilies recognised in the classification, and the distribution of floral sexuality amongst these taxa.

There is a question as to precisely where to draw the boundary of the subfamily. Our cladogram offers two possibilities which are supported by strong characters. A subfamily Aroideae which excluded the tribes Zamioculcadeae and Stylochaetoneae would be defined by absence of perigone, presence of simple laticifers, thick stamen connectives and porose anther dehiscence. By contrast, subfamily Aroideae including these tribes is defined by unisexual flowers, clear differentiation of the spathe into a tube and blade, and spadix differentiated into male and female zones. The latter, more inclusive concept is a better fit with the DNA cladogram of French et al. (1995). A further consideration is that these characters are of more practical use for distinguishing subfamily Aroideae, since unisexuality and gross morphology of the inflorescence is a much more obvious combination of features than absence of a perigone, presence of laticifers or small floral characters. This is therefore the concept we have opted for. Despite our strong advocacy of this taxon, it should be pointed out that the distribution of certain characters on the cladogram, especially inaperturate pollen, is unsatisfactory. Inaperturate pollen arises between tribe Spathiphylleae and the other monsteroid clade low down on the stem of the cladogram, requiring the

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re-evolution of monosulcate pollen from inaperturate in subfamily Lasioideae. This seems highly implausible. A much more likely arrangement would have inaperturate pollen evolving between Stylochaeton and tribe Spathicarpeae, in association with loss of perigone. Inaperturate pollen in tribe Spathiphylleae and Anadendrum would then be homoplasic. A further point here is that V. Tarasevich (pers. comm.) has suggested on the basis of TEM studies that Spathiphyllum pollen is in fact multi-aperturate. The internal topology of our subfamily Aroideae concept remains largely unresolved above the tribal level and this is a problem to which future phylogenetic studies should devoted. We have, for convenience, recognized an informal paraphyletic “Perigoniate Aroideae” and a monophyletic “Aperigoniate Aroideae”, the latter supported by strong characters as noted earlier. Our analysis has thus not made very much further progress in clarifying the relationships of the tribes of the Englerian subfamilies Aroideae, Philodendroideae and Colocasioideae. Under the influence of the molecular studies of French et al. (1995), we have adopted Grayum’s (1984,1990) device of grouping certain tribes into informal Alliances (see chapter 23). The Dieffenbachia

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Alliance is taken directly from their results. Our analysis never associated tribes Spathicarpeae and Dieffenbachieae, but we think this is a very interesting possibility and the molecular results give strong support to this clade. Recognition of the Philodendron Alliance reflects the fact that both morphological and molecular analyses gave a similar result. The Schismatoglottis Alliance is a strong clade in the molecular analysis and although not present in our analysis with Tofieldia as outgroup, it was commonly found in analyses with other outgroups. The Caladium Alliance is a novel group which emerged strongly in the molecular analysis of French et al. (1995). In our morphological analysis tribe Zomicarpeae was problematic, failing to emerge as a monophyletic group and on the whole being associated with the clade including tribe Areae. The results of French and colleagues reconcile the known presence of anastomosing laticifers in tribe Zomicarpeae with their occurrence in tribe Caladieae, the neotropical distribution of both tribes and the possibly intermediate status of Scaphispatha between them. On this basis we have adopted their result for our classification. One of the consequences of this is that the old Englerian subfamily Colocasioideae is no longer recognized in the classification, despite the fact that it emerges consistently in our morphological analysis. This clade is associated in our analysis with the tribes Peltandreae and Ariopsideae and this larger clade (Peltandreae–Chlorospatha in Figure 10) is defined by the following synapomorphies:– short distinct leaf basal

ribs, presence of a sympodial leaf submarginal vein, thickened stamen connectives, connate stamens, presence of staminodes in the female spadix zone, and base chromosome number of x=14. The other tribes are not arranged into Alliances and their sequence reflects only a generalized affinity as we prefer to remain non-committal. In our analysis a consistent group is formed by Arisarum, Ambrosina and Pistia. With Tofieldia as outgroup, tribe Arophyteae also associated consistently with these three genera, although with other outgroups it emerged in a different position. French et al. (1995) found a somewhat similar result, except that tribe Pistieae grouped with the tribes Areae and Colocasieae. We have followed their results in keeping Pistieae separate. Clearly there are many possibilities for alternative topologies within subfamily Aroideae. The tribes of subfamily Aroideae are likely to remain, more-or-less as circumscribed here, as reliable taxonomic units, but it may be expected that in the future they will undergo much rearrangement. The cost of our approach is measured in redundancy in the classification since there are still many monotypic tribes. A desirable future objective would be to find partners for these solitary genera since in our opinion a classification is more useful if it emphasizes sister group relationships rather than degree of anagenesis. Monotypic tribes are a way of roughly indicating that a genus has no obvious sister group relationships. These should therefore be priority targets for improving the classification in the future.

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22 P R E V I O U S C L A S S I F I C AT I O N S

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Excellent surveys have been published by Nicolson (1960a, 1983,1984a, 1988b) and Croat (1992c). We have discussed only those systems which have, in our opinion, exercised most influence on the progress of araceous systematics. Schott, the founding father of the systematics of Araceae, published his first classification in the Meletemata botanica (Schott 1832). He grouped 35 genera into the Araceae and placed Gymnostachys and Acorus in a separate family “Ordo Acoroideae”. Within his Araceae he divided the genera with unisexual, naked (i.e. perigone absent) flowers into subfamily (“Subordo”) Androgynanthae, and those with bisexual, perigoniate flowers into subfamily Hermaphroditanthae. The order of genera progressed essentially from those which, according to our modern paradigm, had the most derived inflorescence and floral types, to those with the least, beginning with Cryptocoryne and Ambrosina and ending with Symplocarpus and Orontium. Although twenty eight years elapsed before Schott (1860) published his final classification in the Prodromus systematis Aroidearum, now greatly expanded to include 107 genera, the fundamentals of his classification remained the same (see Appendix, Table 10). The “Araceae” have now become the “Aroideae”, Gymnostachys and Acorus are included as a subtribe at the end of the system, and the two subfamilies are renamed as rankless taxa, the Diclines, containing those with unisexual, naked flowers and the Monoclines, containing those with bisexual, perigoniate flowers. A notable feature of Schott’s Prodromus treatment is the extremely detailed family description which shows that Schott was fully aware of the great variety of vegetative characters of the Araceae, including latex (“succo decolori l. lacteo”) and, probably, trichosclereids (“rhaphidibus”). His suprageneric taxa are, nevertheless, based almost entirely on floral characters. Only the tribes and subtribes are ranked. The higher, unranked taxa have names which describe their most important diagnostic character, e.g. “Efilamentatae”: filaments absent, “Pachyzeugmaticae”: large anther connectives, “Stenozeugmaticae”: slender anther connectives, “Orthotropooae”: orthotropous ovules, “Anatropooae”: anatropous ovules, “Gymnogoneae”: gynoecium naked, and “Peristatogoneae”: gynoecium with associated staminodes. Schott (1860) used vegetative characters occasionally in the diagnoses of his tribes and subtribes and much more extensively in the generic descriptions. There is no evidence that Schott used evolutionary principles in his work. His philosophy was clearly that of classification by “natural affinities”, i.e. grouping together those taxa which have the largest number of taxonomi-

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cally important characters in common. For Schott, it may be supposed that the most significant taxonomic characters were the floral ones he had so painstakingly elucidated and magnificently illustrated in the Genera Aroidearum (Schott 1858), where the plates presented each genus as an analysis of its floral structure. Engler, however, considered Schott’s classification to be artificial because it was based primarily on floral characters. His new approach was strongly influenced by the earlier morphological and anatomical studies of Irmisch (1858, 1874) and Tieghem (1867, 1872) and by his own original research on the anatomy and shoot organization of the family (Engler 1877, 1878). In his first complete system Engler (1876b) presented the first classification based on phylogenetic principles (see Appendix, Table 11). The order of the genera was reversed so that those with the least derived flowers and inflorescences came first and those with the most derived came last. Ambrosina and Cryptocoryne, for example, were now placed almost at the end of the system. He recognized 10 subfamilies, one of which was the Lemnoideae, corresponding to the modern Lemnaceae. Subfamily Pothoideae was the group which represented the most primitive forms, i.e. those with the most obvious connections to other monocot families through the joint possession of such common characters as 3-merous, perigoniate, bisexual flowers. The subfamilies were defined by a combination of vegetative and floral characters, with a strong emphasis on the presence or absence of laticifers and trichosclereids, life form, shoot organization, leaf venation and phyllotaxis. His subfamily classification embodied the idea of the evolution of unisexual-flowered genera from bisexual-flowered ones in several independent phylogenetic lines, or clades as we would say today. Engler’s attitude to Schott’s system and the phylogenetic method he applied to his own classification of the Araceae are summarized explicitly in his 1876 paper, which includes detailed dendrograms showing presumed phylogenetic relationships (orig. “Verwandtschaft”) between suprageneric taxa. The introductory paragraph is worth quoting in full (freely translated from German) for the light it sheds on Engler’s thinking:– “Natural system of the Araceae. In the following overview of the Araceae, which is systematic rather than analytical, the [suprageneric] groups and genera are arranged so as to convey an idea of the gradual reduction in floral parts [which is seen in the family]. This system makes it possible to

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perceive with particular clarity that reduction in the floral parts must have occurred in various [suprageneric] groups and thus that Schott’s classification, in which the primary subgroups are based on the floral structure, is unnatural. My system of the Araceae would have been quite different had I been aiming to provide an aid to identification for botanists less familiar with the family. My intention is otherwise: to make as clear as possible all the phylogenetic relationships [orig. “verwandtschaftlichen Beziehungen”] existing between the individual [suprageneric] groups. The number of suprageneric groups is consequently larger than might at first sight seem necessary. The Araceae is a poorly represented family in herbaria. Our knowledge of its forms continues to be incomplete, as is shown by the almost annual discovery of new genera, and it may be expected that some of the suprageneric groups comprising only a few genera will later be enriched by the inclusion of one or several [new] ones. All [literature] citations are here omitted since these will be found in my monographic systematic treatment of the Araceae [he must have been referring to his forthcoming monograph in de Candolle’s Monographiae Phanerogamarum – see Engler 1879] and are here of little interest. In those cases where the origin of one genus from another is [considered] highly probable the names are arranged one above the other. Where only a common origin [for the genera] is assumed, their names are placed side-by-side. Generic names in parentheses refer to taxa considered as genera by Schott but which most probably should be considered only as subgenera.” This earliest classification represents the most creative phase of Engler’s Araceae work. It was produced during the period when he was most competely immersed in his studies of the family and was not yet burdened by the immensely ambitious projects of his later career. The dendrograms, published again in de Candolle’s Monographiae Phanerogamarum (Engler 1879), were by far the most detailed cladistic statement that he ever published. Later on, following the discovery and description of many new genera and innumerable new species, he modified his system somewhat (Engler 1887-1889, 1920b), in particular by the exclusion of Lemnaceae, but the essential structure continued unaltered. In his final classification, Engler (1920b) reduced the number of subfamilies to eight (see Appendix, Table 12). These are based on a rather broad range of taxonomic characters but for the most part are not very sharply defined. As Engler had made clear from the outset, they were concepts intended to convey phylogenetic meaning rather than ease the path of identification. In a rough and ready manner subsequent specialists of the family learned to recognize subfamily Pothoideae by their complete lack of laticifers, subfamily Monster oideae by their trichosclereids and mostly aperigoniate bisexual flowers, subfamily Calloideae by their temperate Northern

hemisphere distribution and preference for swampy habitats, subfamily Lasioideae by their frequent possession of deeply sagittate or dracontioid leaves, subfamily Philodendroideae by their unisexual flowers and parallel-pinnate leaf venation, subfamily Colocasioideae by their unisexual flowers, anastomosing laticifers and special type of leaf venation (“colocasioid”; see glossary), and subfamily Aroideae by their unisexual flowers, mostly geophytic habit and frequent possession of a smooth terminal spadix appendix. This classification has been the basis for most subsequent taxonomic work on the family and it is only in recent years that it has undergone substantial alteration. To begin with, only minor modifications were made. Bogner (1979a) published a synopsis in which new genera were inserted into Engler’s framework and new generic synonymy accounted for. Shortly afterwards, however, French and French & Tomlinson began to publish a series of anatomical studies which suggested that some parts of Engler’s classification were unnatural, particularly his subfamilies Pothoideae and Lasioideae. Engler’s concept of araceous phylogeny allowed for the evolution of unisexual-flowered genera from bisexual-flowered ones within these two subfamilies and thus he saw no difficulty in including unisexual-flowered genera such as Culcasia and tribe Zamioculcadeae within subfamily Pothoideae, and likewise Amorphophallus, Anchomanes and others within subfamily Lasioideae. The new anatomical evidence, however, began to reveal that such placements did not reflect real relationships. Two earlier classifications should be mentioned which essentially followed the principles of Schott’s system, those of Hooker (1883) and Hutchinson (1934, 1959, 1973). Hooker (1883) presented a classification which was essentially an updated version of Schott’s (1860) system in the light of Engler’s new studies. Like Schott’s (1860), the classification starts with the unisexual-flowered genera and ends with the bisexual-flowered ones. Hooker recognized 11 tribes but no subfamilies, and placed all the most primitive genera (Orontium, Lysichiton, Symplocarpus, Lasia, Podolasia, Urospatha, Anaphyllum, Ophione (= Dracontium), Cyrtosperma, Spathiphyllum, Anthurium, Pothos, Pothoidium, Acorus and Gymnostachys) in the final tribe, Orontieae, divided into seven subtribes. The 98 genera recognized have similar circumscriptions to those of Schott and Engler. Hutchinson (1934, 1959, 1973) based his system on Hooker’s but reversed the order of the genera to reflect a phylogenetic sequence, starting with the bisexualflowered genera and ending with the unisexual-flowered ones. The first tribes are Acoreae, Orontieae and Spathiphylleae and the last is Areae. He recognized 18 tribes and 126 genera but no subfamilies. The major contribution of his classification was the publication of a key to all the genera in English. Hutchinson’s system was used, notably, by Thanikaimoni (1969) and

PREVIOUS CLASSIFICATIONS

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Marchant (1970, 1971a, b, 1972, 1973), but in general it made little impact on the prevailing use of the Engler system. There are a number of unnatural tribal circumscriptions, for example, the separation of Symplocarpus from Lysichiton and Orontium, of Holochlamys from Spathiphyllum and of Pycnospatha from Dracontium and related genera. The treatments of Hooker and Hutchinson were parts of general classifications of the Flowering Plants rather than specialist studies as in the case of most other systems mentioned here. Hotta (1970) published a classification of Araceae of east Asia and the Malaysian region which was based on Engler’s system. Among other original ideas, it is notable for the recognition of the subfamily Acoroideae (Acorus and Gymnostachys) and the sinking of subfamily Monsteroideae into subfamily Pothoideae. He transferred Lysichiton and Symplocarpus (as tribe Symplocarpeae) to subfamily Lasioideae and Calla to subfamily Philodendroideae. All these concepts were adopted, at least in part, by later authors, such as Grayum (1984, 1990) and Bogner & Nicolson (1991). Grayum (1984,1990) published a comprehensive survey of aroid taxonomic characters, an original comparative survey of pollen morphology and a cladistic analysis of the entire family. He presented a very different-looking picture of the family’s taxonomy to that of Engler (see Appendix, Table 13). In the first place he argued forcefully for the exclusion of Acorus from the family altogether, which strengthened considerably the homogeneity of the Araceae as a result. Major alterations of all the subfamilies were proposed, including those suggested by Hotta (1970). Engler’s subfamily Pothoideae and subfamily Monsteroideae were merged. Subfamily Calloideae was split, with Calla going to join the genera of the old subfamily Philodendroideae and the three genera of Engler’s tribe Symplocarpeae being transferred to the subfamily Lasioideae. Engler’s tribe Amorphophalleae (= tribe Thomsonieae) was shifted to subfamily Aroideae, and his tribes Nephthytideae and Montrichardieae were moved to the subfamily Philodendroideae (= subfamily Calloideae). The Lasioideae also acquired Stylochaeton from tribe Aroideae. Subfamily Philodendroideae was renamed Calloideae, because of the inclusion of Calla, and enlarged yet further with the inclusion of tribes Spathicarpeae (Engler’s Asterostigmateae), Callopsideae, Arophyteae and Culcasieae. Subfamily Colocasioideae changed only slightly with the addition of Zomicarpa. Subfamily Aroideae was modified by the inclusion of Pistia as a tribe, tribe Thomsonieae and Ariopsis. These changes eliminated certain characters which had helped to preserve the integrity of Engler’s subfamilies. In particular, parallel-pinnate leaf venation ceased to be a useful character for defining subfamily Philodendroideae (= Grayum’s subfamily Calloideae). However, Grayum preserved the essentially Englerian concept of independent evolution of unisexual-flowered groups from bisexual-flowered ancestors in various

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different clades. His subfamilies Pothoideae and Lasioideae both include unisexual-flowered genera (tribe Zamioculcadeae and Stylochaeton respectively). His subfamilies Calloideae (Philodendroideae), Colocasioideae and Aroideae consist almost entirely of unisexual-flowered genera as in Engler’s system. Grayum’s work has been crucial to the re-evaluation of the family classification. Not only was it based on a thorough review of the available taxonomic data from many character fields, both old and new, but he employed modern analysis to produce his classification and was bold in proposing major changes backed up by cogent arguments. Grayum presented extensive discussion and a masterful revision of the literature but did not summarize his conclusions in the form of a detailed taxonomic presentation with diagnoses and keys to his generic and suprageneric taxa. Bogner & Nicolson (1991) presented a classification which adhered more closely to Engler’s subfamily concepts but which likewise took account of new data, especially that of French (see Appendix, Table 14). Their primary aim was to provide a detailed synoptical key to the genera, in which the diagnostic characters were clearly presented. The mass of new data on which their changes were based was not detailed in the form of literature citations. They followed Grayum in eliminating Acorus, but in addition they elevated Gymnostachys to subfamilial rank. Subfamily Pothoideae was greatly altered. All unisexual-flowered genera were ejected to other subfamilies and Anadendrum and Heteropsis were transferred to subfamily Monsteroideae, leaving only three closely related genera (Pothos, Pedicellarum and Pothoidium). Subfamily Monsteroideae acquired the circumscription accepted in our treatment by the addition of Anadendrum and Heteropsis. Subfamily Calloideae was reduced to include only Calla, while subfamily Lasioideae now included tribes Orontieae (= Engler’s tribe Symplocarpeae), Anthurium, Zamioculcadeae, Callopsideae, Nephthytideae, Culcasieae and Montrichardieae, as well as the “core” tribe Lasieae. Subfamilies Philodendroideae, Colocasioideae and Aroideae, however, remained very much as Engler had left them, apart from the inclusion of more recently described genera and the transfer of Protarum to subfamily Colocasioideae, and tribe Thomsonieae to subfamily Aroideae. Pistia remained in its own subfamily as proposed by Engler. Bogner & Nicolson’s paper was the first detailed taxonomic synopsis down to the level of genera published by specialists of the family since Engler’s time. It was the result of many years of study and a profound and first-hand knowledge of the taxonomic characters of the genera. Although these authors proposed no explicit phylogenetic scheme, their simplification of subfamilies Pothoideae and Calloideae reduced the number of subfamilies which included both unisexual-flowered and bisexual-flowered genera to one, their much-expanded subfamily Lasioideae.

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Hay & Mabberley (1991) took a completely different approach to the question of araceous phylogeny, starting out from the precepts of the durian theory of Corner (1949, 1952, 1953, 1954a, 1954b). In a long paper bursting with fascinating if unorthodox ideas, they drew a picture of an ancestral araceous type and recognized many features of the Lasieae (our Lasioideae) as primitive, partly through their correspondence with Corner’s hypotheses of primitive angiosperm characters. One point they bring out is that an argument for adaptation in a particular character state does not necessarily support a hypothesis that it is derived in the taxon under consideration. Thus, there is no reason in principle why the ancestral aroid should not have had a tuberous stem and unilocular ovaries with a single basally inserted ovule. They proposed that the araceous inflorescence may have been the result of homoeotic saltation from Nymphaealean ancestral forms and therefore that the typically monocotyledonous features observed in some genera (i.e. those that we consider plesiomorphic in Araceae) are really derived convergences. They

presented a classification based largely on Grayum (1990) but differing in a number of significant respects. The Lasioideae are placed first, and the Pothoideae and Monsteroideae are recognized as separate subfamilies, the former including Anthurium, Anadendrum, Heteropsis and the Zamioculcadeae and the latter with only the Monstereae and the Spathiphylleae. The Aroideae retains the Spathicarpeae, as in Engler’s system, but also includes Ariopsis, Thomsonieae and Pistia. The classification presented here (chapter 23), together with the results of the molecular studies of French et al. (1995), differs from other modern systems and that of Engler in grouping all unisexual-flowered genera together, as Schott did. Unlike Schott, however, our proposal is explicitly phylogenetic. In essence, it appears that the configuration of the available taxonomic data favours a simpler hypothesis than Engler proposed, and indeed modern results flatly contradict Engler’s view of araceous phylogeny. It is not necessary to hypothesize the multiple evolution of unisexual flowers to arrive at a natural classification of Araceae.

PREVIOUS CLASSIFICATIONS

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B

TAXONOMY

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23 S Y N O P S I S O F T H E C L A S S I F I C AT I O N O F A R A C E A E

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Family Araceae Jussieu

MAJOR GROUP PROTO-ARACEAE a. Flowers bisexual I. Subfamily Gymnostachydoideae Bogner & Nicolson 1. Gymnostachys R. Brown II. Subfamily Orontioideae Mayo, Bogner & P.C. Boyce 2. Orontium L. 3. Lysichiton Schott 4. Symplocarpus Nuttall

V. Subfamily Lasioideae Engler 21. Dracontium L. 22. Dracontioides Engler 23. Anaphyllopsis A. Hay 24. Pycnospatha Gagnepain 25. Anaphyllum Schott 26. Cyrtosperma Griffith 27. Lasimorpha Schott 28. Podolasia N.E. Brown 29. Lasia Loureiro 30. Urospatha Schott VI. Subfamily Calloideae Endlicher 31. Calla L.

MAJOR GROUP TRUE ARACEAE a. Flowers bisexual

b. Flowers unisexual

III. Subfamily Pothoideae Engler

VII. Subfamily Aroideae

Tribe 5. 6. 7.

PARAPHYLETIC GROUP: PERIGONIATE AROIDEAE (perigone present)

Potheae Engler Pothos L. Pedicellarum M. Hotta Pothoidium Schott

Tribe Anthurieae Engler 8. Anthurium Schott IV. Subfamily Monsteroideae Engler Tribe Spathiphylleae Engler 9. Spathiphyllum Schott 10. Holochlamys Engler

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Tribe Zamioculcadeae Engler 32. Zamioculcas Schott 33. Gonatopus Engler Tribe Stylochaetoneae Schott 34. Stylochaeton Leprieur MONOPHYLETIC GROUP: APERIGONIATE AROIDEAE (perigone absent)

Tribe Anadendreae Bogner & French 11. Anadendrum Schott

Dieffenbachia Alliance

Tribe Heteropsideae Engler 12. Heteropsis Kunth

Tribe Dieffenbachieae Engler 35. Dieffenbachia Schott 36. Bognera Mayo & Nicolson

Tribe Monstereae Engler 13. Amydrium Schott 14. Rhaphidophora Hasskarl 15. Epipremnum Schott 16. Scindapsus Schott 17. Monstera Adanson 18. Alloschemone Schott 19. Rhodospatha Poeppig 20. Stenospermation Schott

Tribe Spathicarpeae Schott 37. Mangonia Schott 38. Taccarum Schott 39. Asterostigma F.E.L. Fischer & C.A. Meyer 40. Gorgonidium Schott 41. Synandrospadix Engler 42. Gearum N.E. Brown 43. Spathantheum Schott 44. Spathicarpa W.J. Hooker

THE GENERA OF ARACEAE

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Philodendron Alliance Tribe Philodendreae Schott 45. Philodendron Schott Tribe Homalomeneae M. Hotta 46. Furtadoa M. Hotta 47. Homalomena Schott Tribe Anubiadeae Engler 48. Anubias Schott

Schismatoglottis Alliance Tribe Schismatoglottideae Nakai 49. Schismatoglottis Zollinger & Moritzi 50. Piptospatha N.E. Brown 51. Hottarum Bogner & Nicolson 52. Bucephalandra Schott 53. Phymatarum M. Hotta 54. Aridarum Ridley 55. Heteroaridarum M. Hotta Tribe Cryptocoryneae Blume 56. Lagenandra Dalzell 57. Cryptocoryne Wydler

Caladium Alliance Tribe Zomicarpeae Schott 58. Zomicarpa Schott 59. Zomicarpella N.E. Brown 60. Ulearum Engler 61. Filarum Nicolson Tribe Caladieae Schott 62. Scaphispatha Schott 63. Caladium Ventenat 64. Jasarum Bunting 65. Xanthosoma Schott 66. Chlorospatha Engler 67. Syngonium Schott 68. Hapaline Schott

No Alliance Tribe Nephthytideae Engler 69. Nephthytis Schott 70. Anchomanes Schott 71. Pseudohydrosme Engler Tribe Aglaonemateae Engler 72. Aglaonema Schott 73. Aglaodorum Schott

Tribe Culcasieae Engler 74. Culcasia Palisot de Beauvois 75. Cercestis Schott Tribe Montrichardieae Engler 76. Montrichardia H. Crüger Tribe Zantedeschieae Engler 77. Zantedeschia K. Sprengel Tribe Callopsideae Engler 78. Callopsis Engler Tribe Thomsonieae Blume 79. Amorphophallus Decaisne 80. Pseudodracontium N.E. Brown Tribe Arophyteae Bogner 81. Arophyton Jumelle 82. Carlephyton Jumelle 83. Colletogyne Buchet Tribe Peltandreae Engler 84. Peltandra Rafinesque 85. Typhonodorum Schott Tribe Arisareae Dumortier 86. Arisarum P. Miller Tribe Ambrosineae Schott 87. Ambrosina Bassi Tribe Areae 88. Arum L. 89. Eminium (Blume) Schott 90. Dracunculus P. Miller 91. Helicodiceros K. Koch 92. Theriophonum Blume 93. Typhonium Schott 94. Sauromatum Schott 95. Lazarum A. Hay 96. Biarum Schott Tribe Arisaemateae Nakai 97. Pinellia Tenore 98. Arisaema Martius Tribe Colocasieae Engler 99. Ariopsis Nimmo 100. Protarum Engler 101. Steudnera K. Koch 102. Remusatia Schott 103. Colocasia Schott 104. Alocasia G. Don Tribe Pistieae Blume 105. Pistia L.

C SYNOPSIS OF THE CLASSIFICATION

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24 FA M I L Y D E S C R I P T I O N O F A R A C E A E

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Family Araceae Araceae Juss., Gen. Pl. 23 (1789, “Aroideae”), nom. cons. ANATOMY: Calcium oxalate raphides and druses abundant, raphides always present, laticifers commonly present, either simple and articulated or more rarely anastomosing, trichosclereids present (Monstereae, Spathiphylleae, rarely in Potheae), tannin cells common, resin canals sometimes present (Culcasieae, Homalomeneae, Philodendreae). HABIT: evergreen to seasonally dormant herbs, perennial, sometimes gigantic, climbing or subshrubby hemiepiphytes, epiphytes, lithophytes, terrestrial, geophytes, helophytes, sometimes rheophytes, true aquatics, rarely free-floating (Pistia). STEM: aerial and erect to climbing or creeping with very short (plant rosulate) to very long (plant scandent) internodes, or subterranean and consisting of a subglobose to depressed-globose tuber (sometimes turnip- or carrot-like or irregular in shape) or horizontal to erect rhizome; terrestrial plants and helophytes sometimes arborescent with massive stem and terminal rosette of leaves (Xanthosoma, Alocasia, Montrichardia, Philodendron) or arborescent with a pseudostem of petiole sheaths (large in Typhonodorum, small in many Arisaema); geophytes often with solitary leaf. SHOOT ORGANIZATION: the mature, flowering stem is almost always a sympodium composed of a series of articles, rarely the stem is monopodial (Potheae, Heteropsideae); each article begins with a 2-keeled (except Orontioideae) prophyll followed by a series of leaves and terminates with an inflorescence; leaf number per article may be determinate or indeterminate, and from one or very few to very many; the leaves of each article normally consist of a mixture of foliage leaves with partially to fully developed blades and cataphylls; the sympodial leaf is that subtending the inflorescence and may be a foliage leaf or a cataphyll; the prophyll is almost always a cataphyll (except Orontioideae); subsequent articles (continuation shoots) normally arise at the second node below the spathe node (except Orontioideae); juvenile shoots and flagelliform branches are usually monopodial; terminal inflorescences may be solitary or may form a floral sympodium of several inflorescences; the articles of floral sympodia normally consist of a single 2-keeled prophyll and an inflorescence; the subsequent article of a floral sympodium normally arises in the axil of the preceding prophyll, i.e. at the first node below the spathe node. VEGETATIVE PROPAGATION: climbing hemiepiphytes frequently form flagelliform

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shoots, which are stolon-like branches with leaves reduced to small cataphylls and very elongated internodes, adapted to seek out new host trees; other adaptative forms are subterranean stolons (e.g. Spathiphyllum, Lasimorpha) and tubercules (e.g. Dracontium), bulbils with recurved scales borne on specialized shoots (Remusatia), bulbils on petioles (Pinellia ternata) or leaf blade (Amorphophallus bulbifer), or formation of new plants from abscissed leaflets (Zamioculcadeae). ROOTS: always adventitious, primary root withering soon after germination, sometimes dimorphic (climbing hemiepiphytes) with anchor roots and larger feeder roots, sometime contractile roots present (geophytes), rarely roots very fleshy, water-storing (Stylochaeton). LEAVES: usually spirally arranged, sometimes distichous; normally differentiated into petiole and expanded blade (except e.g. Gymnostachys, some Biarum spp.), usually glabrous, rarely pubescent, tomentose, villous or with small to large and complex trichomes or papillae (e.g. Philodendron squamiferum) on the petiole; ptyxis usually convolute, rarely involute (e.g. Anthurium sect. Pachyneurium, Lagenandra); blade and petiole often variegated or mottled with spots, bands, blotches or irregularly shaped patches and zones of various colours, usually shades and mixtures of green, yellow and silver. CATAPHYLLS: caducous, marcescent, deciduous or persistent, sometimes beautifully mottled and patterned (e.g. Arisaema, Asterostigma), when persistent sometimes a conspicuous feature of plant and either membranous or forming fibrous mass (e.g. many spp. of Anthurium, Philodendron). PETIOLE: often as long as or longer than blade, usually smooth, sometimes hairy, papillose, warty, prickly or aculeate (e.g. Lasioideae), occasionally covered with large multicellular processes (e.g. Philodendron squamiferum), rarely massively succulent and water-storing (e.g. Zamioculcas, Philodendron martianum), often geniculate (pulvinate) apically (e.g. Anthurium), basally or rarely centrally (e.g. Gonatopus boivinii); sheath normally well-developed, often at least half as long as entire petiole, sometimes ligulate apically, often very reduced in sympodial leaves (especially in Anthurium, most Philodendron spp.). LEAF BLADE: simple to compound, extremely variable in shape – rarely filiform (e.g. Cryptocoryne consobrina), linear (Jasarum), most commonly elliptic, ovate, oblong, sagittate, hastate, less commonly trifid to trisect, pedatifid to pedatisect, radiatisect, dracontioid (i.e. trisect with each primary division further much divided), pinnatifid to pinnatisect (Zamioculcas), bipinnatifid, tripinnatifid to quadripinnatifid (Gonatopus), fenestrate (Monstera) or laciniate

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(i.e. fenestrate with slit-like holes, Cercestis mirabilis); heteroblasty frequent, especially in climbing hemiepiphytes, “shingle leaves” sometimes formed (e.g. some Potheae, some Monstereae); seedling leaves usually entire when epigeal, rarely first foliage leaf compound (Gonatopus, Amorphophallus). LEAF VENATION: midrib almost always differentiated, sometimes massive and succulent (e.g. Philodendron crassinervium); primary veins usually arising pinnately from midrib (and then called primary lateral veins), either running into marginal vein (e.g. Philodendron, Dieffenbachia) or joining distally to form a submarginal collective vein on each side (e.g. Caladium, many Anthurium spp.), sometimes primary veins all arising from petiole insertion and running arcuately into leaf apex (e.g. Orontium, Anthurium sect. Digitinervium), rarely strictly parallel (Gymnostachys) or subparallel (Pistia), sometimes not differentiated at all (e.g. Philodendron crassinervium); secondary and tertiary veins either reticulate (e.g. Areae), or parallel-pinnate, i.e. running parallel to primaries (e.g. Philodendron), or arising from primaries at a wide angle and then arching strongly towards leaf margin (e.g. Colocasia), sometimes forming sinuous or zig-zag interprimary veins (e.g. Caladieae); higher order venation reticulated or forming cross connections between lower order veins. INFLORESCENCE: terminal, solitary, or 2 to many in a synflorescence, usually appearing to be axillary to sympodial leaf, consisting of a spadix (spike) of small flowers and subtended by a spathe (bract), usually erect, sometimes pendent (e.g. Anthurium wendlingeri, Stenospermation, Piptospatha), sometimes becoming pendent after anthesis (e.g. Typhonodorum). PEDUNCLE: very short to very long, usually similar to petiole in appearance, coloration, pubescence or armature, normally longer than spadix stipe, sometimes ± suppressed and spadix stipe elongated (Orontium, Lysichiton). SPATHE: nearly always conspicuous (except Gymnostachys, Orontium), very variable in shape and colour, simpler forms (e.g. many Anthurium spp.) often green, reflexed or spreading, more complex forms often showy and highly coloured, erect, usually either boat-shaped or constricted centrally to form a basal tube and an apical blade; tube may enclose the female zone of the spadix or both fertile zones or rarely the entire spadix (e.g. Cryptocoryneae), very occasionally much longer than blade (e.g. many Cryptocoryne spp.), tube margins usually convolute, sometimes connate (e.g. Sauromatum, Stylochaeton, Arisarum); blade usually erect and gaping, sometimes widely spreading, twisted, reflexed or rarely margins ± closed forming slit-like opening (e.g. most Lagenandra spp.); spathe constriction may lie between or above male and female zones or occur in two places (e.g. some Remusatia spp.); spathe entirely deciduous soon after anthesis (e.g. most Monstereae), or tube persistent to fruiting and blade marcescent to deciduous after anthesis (Caladieae, Colocasieae, Schismatoglottideae), or spathe entirely persistent until fruiting (e.g.

Philodendron, Homalomena) or whole spathe gradually withering and rotting (most Areae). SPADIX: usually erect, often fleshy and relatively thick, sessile or shortly stipitate, rarely very long-stipitate (e.g. Lysichiton, Orontium, some Anthurium spp.), usually free, sometimes adnate basally (e.g. Hapaline, Dieffenbachia) or entirely (Spathicarpa) to spathe, either ± uniform in appearance (flowers bisexual or monoclinous), or divided into distinct floral zones (flowers unisexual or diclinous), fertile zones contiguous or separated by sterile zones, female (pistillate) zone always basal and male (staminate) zone either apical or intermediate in position (except Spathicarpa), rarely bisexual flowers occur between male and female zones (e.g. Arophyteae); sterile zones may be basal, intermediate or apical or any combination of these, apical sterile zone usually known as a terminal appendix; rarely a single plant produces inflorescences bearing male flowers only, followed in later years by inflorescences bearing female flowers only, and vice versa (paradioecy, known only in Arisaema). FLOWERS: 2- to 3-merous (-mery often hard to detect in unisexual flowers), bisexual (monoclinous, hermaphrodite) or unisexual (diclinous), very small, protogynous, lacking floral bracts, usually numerous (except e.g. Pistia, Ambrosina), sessile (except Pedicellarum), usually densely arranged, sometimes laxly so (e.g. some Pothos spp., male flowers of Arisarum, female flowers of Dieffenbachieae); bisexual flowers with or without (Calloideae, Monstereae) a perigone (perianth), unisexual flowers usually without a perigone (present in Zamioculcadeae, Stylochaeton) but sometimes including rudimentary organs representing modified sexual parts of the other sex (e.g. staminodes of female flowers in Dieffenbachia and Spathicarpeae, cup-like synandrodium in Arophyteae, pistillodes present in male flowers of e.g. Stylochaeton, Furtadoa, some Spathicarpeae spp., central stigmatoid body of the synandrium present in Taccarum and some Gorgonidium spp.). PERIGONE: composed of free (e.g. Anthurium) or partially connate (some Pothos spp.) tepals, or consisting of a single cup-like structure (e.g. Spathiphyllum cannifolium); when free, tepals 4 to 6 (–8) and imbricate in 2 whorls, membranaceous (e.g. Anadendrum) or more commonly thickened at least apically, truncate (Zamioculcadeae) to cucullate (Lasioideae). STAMENS (bisexual perigoniate, bisexual non-perigoniate, unisexual perigoniate flowers):

usually free (filaments connate in Gonatopus and often in Lasimorpha), equal in number and opposite to tepals (when present), rarely more (e.g. some Dracontium spp.); filaments distinct, often ± oblong and flattened (e.g. Anthurium), rarely filiform (Stylochaeton), usually rapidly elongating to push anthers above perigone or gynoecium at anthesis; anthers usually terminal, basifixed, extrorse (introrse in Zamioculcas, latrorse in Pedicellarum), always composed of 2 thecae each with 2 microsporangia; connective usually slender, inconspicuous, often over-

FA M I L Y D E S C R I P T I O N

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topped by thecae, thecae dehiscing by single longitudinal slit or apical stomial pore, with all intermediate degrees occurring. MALE FLOWER (unisexual nonperigoniate flowers): 1–8 androus (rarely more, e.g. Alocasia brisbanensis), floral grouping of stamens sometimes obvious in mature inflorescence (e.g. many Philodendron and Homalomena spp.), often obscured during ontogeny; stamens free or partially to completely connate to form a synandrium. FREE STAMENS (unisexual non-perigoniate flowers): usually sessile to subsessile, filament sometimes distinct (e.g. Schismatoglottis), connective sometimes ± slender (e.g. Areae) but often strongly thickened, apically broad, fleshy and probably osmophoric (e.g. Philodendron), thecae lying opposite or adjacent on one side of stamen, dehiscing by single longitudinal slit or apical stomial pore, with all intermediate degrees occurring, rarely both microsporangia dehiscing independently by separate stomial pores (some Amorphophallus spp.), rarely theca prolonged apically into a horn dehiscing by single pore (Cryptocoryneae, some Schismatoglottideae). SYNANDRIUM (unisexual non-perigoniate flowers):

usually ± sessile, sometimes formed by fusion of filaments only (e.g. Arisaema, Arisarum, Carlephyton sect. Pseudocolletogyne, Gorgonidium), more commonly composed of completely connate stamens and then usually apically truncate and ± prismatic in apical crosssection (e.g. Caladieae), sometimes mushroom-shaped (Asterostigma) or cylindric (Taccarum), very rarely the synandria themselves connate (Ariopsis); common connective usually broad, fleshy and probably osmophoric (e.g. Caladieae, Alocasia); thecae either lateral, apical or marginal depending on the degree of elongation of the thecae and the extent to which they are overtopped by the common connective, dehiscing by single longitudinal slit or apical stomial pore, with all intermediate degrees occurring. POLLEN: shed in monads, rarely shed in tetrads (Xanthosoma, Chlorospatha), aperturate in most bisexual-flowered genera, inaperturate in most unisexual-flowered genera, exine various (see chapter 10). STERILE ORGANS (pistillodes, staminodes, synandrodes): often forming zones between fertile zones, sometimes present below female zone (Schismatoglottideae), or on base of terminal appendix, very variable in shape, most often ± truncate and prismatic (e.g. Philodendron), more rarely filiform, subulate, bristle-like or elongate-clavate (Areae), spathulate (Bucephalandra), cylindric (Aridarum) or enlarged and pearl-like (Amorphophallus margaritifer). TERMINAL APPENDIX: present only in some genera (e.g. Thomsonieae, Areae), probably always osmophoric, partly or completely (e.g. Pseudodracontium) covered with staminodes, rugose or corrugated or entirely smooth (e.g. most Areae), intermediate conditions also occur (e.g. Ulearum). FEMALE FLOWER (unisexual, non-perigoniate flowers): Gynoecium sometimes surrounded by a whorl of

variously shaped staminodes (e.g. Dieffenbachia, Spathicarpeae), or sometimes ± regularly associated

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with a single clavate staminode (e.g. Homalomeneae, Schismatoglottis). GYNOECIUM (bisexual and unisexual flowers): ovary usually 1–3 locular, rarely more (e.g. Philodendron, most Spathicarpeae), 1-locular ovaries probably always pseudomonomerous; ovules 1–many per locule, orthotropous, hemiorthotropous, campylotropous, amphitropous, hemianatropous or anatropous; placenta 1–several, axile, parietal, apical, basal, or basal and apical; stylar region (tissue lying between ovary and stigmatic epidermis) usually welldeveloped, usually at least as broad as ovary, sometimes attenuate and elongate (e.g. many Amorphophallus, Arisaema, Biarum spp., Dracontium, some Spathiphyllum spp.) or massive and truncate (most Monstereae spp.), rarely dilated and connate with those of neighbouring gynoecia (Xanthosoma); stigma hemispheric, capitate, discoid, umbonate, more-or-less strongly lobed (e.g. some Amorphophallus, Dieffenbachia spp.), rarely stellately lobed (Asterostigma), sometimes brightly coloured (e.g. some Alocasia, Amorphophallus spp.), always wet from copious stigmatic secretion during female anthesis, sometimes producing conspicuous nectar droplet (e.g. Anthurium). FRUIT: normally a juicy berry, rarely mesocarp leathery; berries normally free, rarely connate (Syngonium) or connate and dehiscent as a syncarp (Cryptocoryne), usually red, orange or purplish red, sometimes white (e.g. some Philodendr on, Stenosper mation), yellow (Typhonodorum), green (Arophyton buchetii, Lysichiton, Orontium, Peltandra virginica), very rarely blue (Amorphophallus kerrii, Gymnostachys), or brownish (Jasarum); infructescence densely packed, cylindric to globose, exposed by withering, basal abscission (e.g. Philodendron) or splitting (e.g. Alocasia, Dieffenbachia) of spathe, rarely berries dehiscent, either basally (Lagenandra) or apically with the seeds exposed by ± simultaneous sloughing of stylar regions of all berries (e.g. Monstereae). SEED: 1–many per berry; testa thick to thin, smooth, roughened, verrucose or striate-costate, papery in seeds with highly developed embryos (Gonatopus), sometimes decaying at maturity (Orontium), or lacking altogether (Gymnostachys, Nephthytis), sometimes arillate with a conspicuous strophiole (e.g. most Areae, Ambrosina), rarely operculate (e.g. Pistia); embryo usually straight, sometimes curved (e.g. Cyrtosperma, Epipremnum), usually undifferentiated, rarely with highly developed plumule (e.g. Cryptocoryne ciliata, Orontium, Typhonodorum) and then endosperm lacking and outer cell layers of embryo chlorophyllous; endosperm copious or absent, with all intermediate states occurring. 105 genera, over 3300 spp.; distribution subcosmopolitan, most abundant and diverse in tropical latitudes.

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1. Plants free-floating aquatics; leaves rosulate, hairy; flowers unisexual, naked; inflorescence with a single female flower and a few male flowers · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 105. Pistia 1. Plants terrestrial or helophytes, climbing hemiepiphytes, epiphytes or lithophytes or other, but never floating 2. Leaves not differentiated into petiole and blade, primary venation strictly parallel; inflorescence borne on a culm-like axis 3. Leaves ensiform, unifacial; spadix solitary, pseudolateral and overtopped by a single, erect, leaf-like spathe; flowers 3-merous, tepals 6 · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · Acorus (Acoraceae) 3. Leaves dorsiventrally flattened, bifacial; flowering shoot with long culm-like axis, bearing numerous spadices distally, these borne in axillary clusters subtended by elongate bracts; flowers 2-merous, tepals 4 · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 1. Gymnostachys 2. Leaves with distinct petiole and expanded blade, primary venation never strictly parallel 4. Flowers with obvious perigone of free or fused tepals (except Pycnospatha which lacks perigone, but has dracontioid leaf, tuberous stem and boat-shaped, fornicate spathe – see lead 22) 5. Flowers bisexual, spadix uniform in appearance with flowers of only one type 6. Higher order leaf venation parallel-pinnate; tissues with abundant trichosclereids 7. Spathe persistent; tepals free or connate; ovary 2–4-locular; ovules 2–8 per locule, placenta axile · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 9. Spathiphyllum 7. Spathe deliquescent; tepals connate; ovary 1-locular; ovules several, placenta basal · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 10. Holochlamys 6. Higher order leaf venation clearly reticulated; tissues without trichosclereids or trichosclereids very few 8. Stem aerial, not tuberous or rhizomatous, never aculeate; plant usually a climbing hemiepiphyte or epiphyte, less often lithophyte or terrestrial, only very rarely helophytic (some spp. of Anthurium) 9. Neotropical plants; seeds with copious endosperm; pollen usually forate, never monosulcate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 8. Anthurium 9. Palaeotropical plants; seeds without endosperm; pollen monosulcate or inaperturate 10. Stigma transversely oblong; stamens always 4 per flower; pollen inaperturate; perigone consisting of a single cup-like structure · · · · · · · · · · · · · · · · · · · 11. Anadendrum 10. Stigma hemispheric to discoid; stamens usually 6 per flower; pollen monosulcate; perigone usually consisting of free tepals or when connate and cup-like the flowers are borne on short pedicels 11. Ovary 3-locular; locules 1-ovulate; flowering shoot with inflorescences always axillary 12. Flowers sessile; tepals free, very rarely basally united · · · · · · · · · 5. Pothos 12. Flowers pedicellate; tepals connate · · · · · · · · · · · · · · · · · 6. Pedicellarum 11. Ovary 1-locular; flowering shoots terminating in a branching system of spadices · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 7. Pothoidium 8. Stem typically subterranean, tuberous or rhizomatous, sometimes aerial and creeping or scrambling but then aculeate; plant frequently a helophyte 13. Plants of temperate regions (N. America, NE. Asia); leaf blade always entire, ovate to elliptic 14. Ovary 2-locular; ovules 2 per locule, placenta axile · · · · · · · · · · · · · 3. Lysichiton 14. Ovary 1-locular; ovule 1, placenta apical or basal 15. Placenta basal; spathe inconspicuous; spadix cylindric, stipe very long · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 2. Orontium 15. Placenta apical; spathe thick, ventricose, enclosing spadix; spadix subglobose, stipe short · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 4. Symplocarpus 13. Plants of tropical and subtropical regions; leaf blade sagittate, pinnatifid, pinnatisect or dracontioid

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16. Leaf deeply sagittate, anterior division not pinnatifid or pinnatisect 17. Ovary many- to 2-ovulate, rarely 1-ovulate; seeds with endosperm 18. Plants without stolons; petiole spines dispersed; stamen filaments free; tropical Asia to Oceania · · · · · · · · · · · · · · · · · · · · · · · · 26. Cyrtosperma 18. Plants with long stolons; petiole spines in ridges; stamen filaments free or connate; tropical West Africa · · · · · · · · · · · · · · · · · · · · · 27. Lasimorpha 17. Ovary 1-ovulate, rarely 2-ovulate; seeds without endosperm or rarely with a little endosperm 19. Petiole aculeate, with obvious spines; Malay Archipelago · · 28. Podolasia 19. Petiole smooth to scabrid-verrucose, never aculeate; tropical America 20. Leaf blade never fenestrate; spathe lanceolate, very long-acuminate and usually spirally twisted; ovary locules with (1–)2 to several ovules; neotropics · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 30. Urospatha 20. Leaf blade often perforated with a few perforations of irregular size between primary lateral veins; spathe fornicate; endemic to Brazil (coastal Bahia and Espirito Santo) · · · · · · · · · · · · · · · · · · · 22. Dracontioides 16. Leaf blade pinnatifid, pinnatisect, dracontioid or sometimes ± pedatifid; anterior division always pinnately divided, either pinnatifid, pinnatisect or yet more highly divided 21. Stem aculeate, aerial and scrambling to prostrate, internodes distinct, green · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29. Lasia 21. Stem not aculeate, subterranean, internodes very abbreviated, not green 22. Leaf blade dracontioid, anterior division bipinnatifid or yet more highly divided; stem a depressed-globose tuber; spathe fornicate 23. Tropical America; flowers with perigone of 4–8 free tepals; berries smooth, red · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 21. Dracontium 23. Tropical southeast Asia; flowers without perigone; berries aculeate, dark green · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 24. Pycnospatha 22. Leaf blade pinnatifid, pinnatisect, or sometimes ± pedatifid, anterior division pinnatifid to pinnatisect; stem a vertical or horizontal rhizome; spathe erect, not fornicate, blade often spirally twisted apically 24. Tropical America; testa thick, verrucose; embryo curved · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 23. Anaphyllopsis 24. Southern India; testa membranous, smooth; embryo straight · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 25. Anaphyllum 5. Flowers unisexual, spadix clearly divided into basal female zone and apical male zone; tropical Africa 25. Leaf pinnatisect to tri- or quadripinnatifid; tepals free; spathe margins free 26. Leaf blade pinnatisect; stamens free · · · · · · · · · · · · · · · · · · · · · · · · · 32. Zamioculcas 26. Leaf blade bipinnatifid to quadripinnatifid, at least in lowest pinnae; stamen filaments connate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 33. Gonatopus 25. Leaf entire, linear to cordate, sagittate or hastate; tepals connate into cup; spathe margins connate basally · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 34. Stylochaeton 4. Flowers without perigon of free or fused tepals 27. Flowers bisexual; spadix uniform in appearance with flowers of only one type (sometimes with sterile flowers at spadix base) 28. Helophytes from temperate regions of northern hemisphere; petiole sheath with long apical ligule · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 31. Calla 28. Climbing hemiepiphytes or sometimes epiphytes or very rarely rheophytes (few Rhaphidophora) from tropical regions; petiole sheath non-ligulate or ligule only short 29. Petiole usually very short with non-annular insertion; trichosclereids not present in tissues, leaf never perforated or lobed; primary lateral veins forming distinct submarginal vein · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 8. Heteropsis 29. Petiole well-developed with annular insertion and usually conspicuous sheath; trichosclereids present in tissues, or if absent (or nearly so) then leaf with conspicuously reticulate higher order venation and often perforated or lobed (Amydrium); primary lateral veins usually not forming distinct submarginal vein

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30. Trichosclereids rare or nearly absent; higher order leaf venation completely reticulate; ovary 1-locular, placenta 1, intrusive-parietal, ovules 2 · · · · · · · · · 13. Amydrium 30. Trichosclereids abundant; higher order leaf venation parallel to primary lateral veins, or only finest venation reticulate 31. Ovary 1-locular or incompletely 2-locular 32. Ovules anatropous, more than one 33. Ovules numerous, superposed on 2 (rarely 3) parietal placentas; seeds fusiform, straight, 1.3–3.2 mm long, 0.6–1.0 mm wide · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 14. Rhaphidophora 33. Ovules 2–4 (–6) at base of a single intrusive placenta; seeds curved, 3–7 mm long, 1.5–4.0 mm wide · · · · · · · · · · · · · · · · · · 15. Epipremnum 32. Ovules amphitropous to anatropous, solitary, basal 34. Adult leaf blade entire; palaeotropics · · · · · · · · · · · · · 16. Scindapsus 34. Adult leaf blade pinnatifid; neotropics (Amazonia) 18. Alloschemone 31. Ovary 2–5 locular 35. Seeds fusiform, claviform or lenticular, less than 3 mm long, endosperm present; ovules (2–)3-many per locule; leaf blade entire 36. Placenta basal; seeds fusiform to claviform; leaf blades thickly coriaceous · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 20. Stenospermation 36. Placenta axile; seeds lenticular and flattened, strongly curved; leaf blades mostly membranous · · · · · · · · · · · · · · · · · · · · · · · 19. Rhodospatha 35. Seeds globose to oblong, 6–22 mm long, the raphe S-shaped; endosperm absent; ovules 2 per locule; leaf blade variously shaped, often perforated or pinnatifid or both · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 17. Monstera 27. Flowers unisexual; spadix clearly divided into basal female zone and apical or intermediate male zone, flowers very rarely in longitudinal rows (Spathicarpa) 37. Spadix fused laterally on both sides to spathe and entirely enclosed by it, forming a septum dividing the spathe into two chambers, with a single gynoecium on one side and the male flowers arranged in 2 rows on the other; very small, seasonally dormant plants endemic to western Mediterranean · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 87. Ambrosina 37. Spadix free or fused to spathe in various degrees but never fused laterally on both sides to spathe to form two internal chambers with a single gynoecium on one side and the male flowers on the other 38. Stamens of each male flower free or only the filaments connate 39. Spadix never entirely enclosed by spathe in a basal “ kettle “ formed of connate spathe margins (if spathe margins basally connate then plant never aquatic) 40. Higher order leaf venation parallel-pinnate 41. Upper part of spathe persisting as long as lower part; petiole sheath lacking ligule; ovary 1–many locular; thecae dehiscing by subapical pores or longitudinal slits; connective usually conspicuously thickened 42. Spathe variously shaped, never campanulate; plants tropical American or tropical Asian; peduncle usually short, if long then female flowers in single whorl (Aglaodorum) 43. Plant always terrestrial, rarely aquatic, never climbing or epiphytic; inflorescences not secreting resin at anthesis; endothecium with cell wall thickenings; ovary 1 locular or incompletely 2–5 locular; most tropical Asian (except Homalomena sect. Curmeria) 44. Seed without endosperm, embryo large; ovule 1, placenta basal or parietal 45. Inflorescence with short peduncle; female flowers in spirals; stem erect to repent; placenta basal; forest plants · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 72. Aglaonema 45. Inflorescence with long peduncle; female flowers in a single whorl; stem repent; placenta parietal; on tidal mudflats · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 73. Aglaodorum 44. Seed with copious endosperm, embryo relatively small; ovules several to many, placenta basal, parietal or axile

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46. Male flower consisting of solitary stamen overtopped by flask-shaped pistillode; ovary 1-locular, placenta basal · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46. Furtadoa 46. Male flower consisting of 2–6 stamens, pistillodes absent; ovary incompletely 2–5 locular, placentas parietal and axile · · · · · · · · · · · · · · · · · · · · · 47. Homalomena 43. Plant usually climbing or epiphytic; inflorescences secreting resin from spathe or spadix at anthesis; endothecium nearly always lacking cell wall thickenings; ovary completely 2–many locular, placenta axile to basal; tropical America · · · · · · · · 45. Philodendron 42. Spathe obconic to campanulate; plants from Southern Africa (naturalized in America and Asia); peduncle long, sometimes longer than leaves · · · · · · · · · · · · · · · · · · · · 77. Zantedeschia 41. Upper part of spathe marcescent or caducous at anthesis, lower part long-persistent; petiole sheath with long, marcescent ligule (except most Schismatoglottis spp.); ovary 1-locular; thecae dehiscing by apical pores, connective not conspicuously thickened 47. Placentas parietal; thecae truncate. 48. Spathe constricted; ovules anatropous to hemianatropous; petiole sheath usually not ligulate; upper part of spadix usually sterile. · · · · · · · · · · · · · · · 49. Schismatoglottis 48. Spathe not constricted; ovules hemiorthotropous to orthotropous; petiole sheath with long, marcescent ligule; spadix fertile almost to apex · · · · · · · · · · · · · · · · 50. Piptospatha 47. Placenta basal or basal and apical; thecae truncate or horned 49. Thecae truncate; placenta basal · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 51. Hottarum 49. Thecae horned; placenta basal or basal and apical 50. Stigma smaller than ovary; upper part of spadix sterile with a distinct appendix of hornless sterile flowers; spathe constricted or not; stamens never excavated apically. 51. Spathe not constricted; male flowers smooth or verrucose; sterile flowers between male and female flowers flattened · · · · · · · · · · · · · · · · · · 52. Bucephalandra 51. Spathe constricted; male flowers densely tuberculate; sterile flowers between male and female flowers subcylindric · · · · · · · · · · · · · · · · · 53. Phymatarum 50. Stigma as broad as ovary; upper part of spadix mostly fertile to apex and without a distinct appendix; spathe not conspicuously constricted; stamens all or mostly excavated apically 52. Stamens all excavated; placenta basal · · · · · · · · · · · · · · · · · · · · 54. Aridarum 52. Two lateral stamens of each male flower excavated and thecae horned, central stamen truncate and thecae hornless; placentas basal (fertile ovules) and apical (apparently sterile) · · · · · · · · · · · · · · · · · · · · · · · · · · · · 55. Heteroaridarum 40. Higher order leaf venation reticulate 53. Leaf blade dracontioid, leaf solitary in each growth period 54. Petiole usually aculeate; at least some of the ultimate leaf lobes trapezoid, truncate or shallowly bifid, veins not forming regular submarginal collective vein on each side 55. Peduncle long; ovary 1-locular · · · · · · · · · · · · · · · · · · · · · · · · · · · · 70. Anchomanes 55. Peduncle very short; ovary 2-locular · · · · · · · · · · · · · · · · · · · · 71. Pseudohydrosme 54. Petiole usually smooth, sometimes rugose but never aculeate; ultimate leaf lobes usually oblong-elliptic, acuminate, with primary lateral veins forming regular submarginal collective veins on each side 56. Ovary 1–4-locular; terminal appendix smooth, rugose, rarely verrucose or staminodial (appendix absent in Amorphophallus margaritifer and A. coudercii) · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 79. Amorphophallus 56. Ovary always 1-locular; terminal appendix staminodial, separated from male zone by naked axial region · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 80. Pseudodracontium 53. Leaf blade shape of various types but never dracontioid; usually several leaves present 57. Spadix fertile to apex, terminal appendix absent 58. Helophytes with robust, erect stems and an apical crown of sagittate to hastate (rarely trisect) leaves; tropical America · · · · · · · · · · · · · · · · · · · · · · · · · · · · 76. Montrichardia 58. Terrestrial, hemiepiphytic or epiphytic plants, leaf blade variously shaped; tropical Africa. 59. Leaf blade usually with pellucid resin canals (lines or points); plants mostly climbing hemiepiphytes; spathe boat-shaped, convolute basally; anthers lacking endothecial thickenings 60. Laticifers absent; flagelliform shoots absent; leaf blade always simple, acute to rounded at base; ovary 1–3-locular; spadix stipitate or sessile · · · 74. Culcasia

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60. Laticifers present; flagelliform shoots present; leaf blade oblong-lanceolate to cordate, sagittate, hastate, trifid or laciniate to pinnatifid; ovary 1-locular; spadix sessile · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 75. Cercestis 59. Leaf blade lacking pellucid resin glands; plants terrestrial; spathe ± fully expanded, not convolute; anthers with endothecial thickenings 61. Leaf cordate-sagittate or subtriangular, deeply sagittate or trifid, glabrous; spathe green; spadix entirely free of spathe · · · · · · · · · · · · · · · · · · · 69. Nephthytis 61. Leaf cordate-ovate, minutely hispid abaxially; spathe pure white; female zone of spadix adnate to spathe · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 78. Callopsis 57. Spadix with ± smooth terminal appendix 62. Laticifers anastomosing; tropical South America 63. Ovary 6- to 9-ovulate; leaf blade trisect to pedatisect; stem a subglobose tuber · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 58. Zomicarpa 63. Ovary 1- to 6-ovulate; leaf blade cordate-sagittate; stem a subglobose tuber or rhizome 64. Appendix slender 65. Stamen connective much prolonged, thread-like; stem tuberous; ovary 1ovulate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 61. Filarum 65. Stamen connective not at all prolonged; ovary 1-6-ovulate; stem a creeping rhizome · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 59. Zomicarpella 64. Appendix relatively thick and subcylindric; stem a creeping rhizome; ovary 1ovulate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 60. Ulearum 62. Laticifers simple; temperate Eurasia and palaeotropics 66. Spadix with zone of sterile flowers between male and female zones, rarely with a naked axis between female and male zones of spadix (Arum pictum) or with fertile zones contiguous (Dracunculus) 67. Placenta parietal to subbasal; leaf blade sagittate or hastate · · · · · · · 88. Arum 67. Placenta apical and/or basal; leaf blade variously shaped 68. Placentas basal and apical 69. Male zone of spadix contiguous with female zone; leaf blade pedatifid but lobes not spirally twisted upwards · · · · · · · · · · · · · · 90. Dracunculus 69. Male zone of spadix separated from female zone by subulate to filiform sterile organs; leaf blade variously shaped 70. Appendix covered with subulate to setiform sterile flowers; leaf blade pedatifid, lobes twisting upwards on each side · · 91. Helicodiceros 70. Appendix smooth; leaf blade oblong-lanceolate or sagittate-hastate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 92. Theriophonum 68. Placenta basal 71. Lower spathe margins free (except Typhonium hirsutum) 72. Infructescence borne above ground level, berries dark red to purple, pericarp juicy; sterile zone between male and female zones of spadix relatively long, often partially naked; tropical and subtropical to warm temperate Asia to Australia · · · · · · · · · · · · · · · · · 93. Typhonium 72. Infructescence borne at or below ground level, berries white to pale lilac, pericarp firm, not juicy; sterile zone between male and female zones relatively short and covered entirely with subulate sterile flowers; Turkey, eastern North Africa, Near East, central Asia · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 89. Eminium 71. Lower spathe margins connate for an appreciable distance (entirely free in Biarum aleppicum) 73. Leaf usually solitary, blade deeply pedatifid to pedatisect; ovary 2–several-ovulate · · · · · · · · · · · · · · · · · · · · · · · 94. Sauromatum 73. Leaves several; blade linear to ovate, elliptic or obovate; ovary 1ovulate 74. Leaf blade broadly elliptic, spathe tube septate · · 95. Lazarum 74. Leaf blade linear, ovate, elliptic-oblong or obovate; spathe tube not septate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 96. Biarum 66. Spadix usually without sterile flowers (sometimes present in Arisaema)

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75. Ovary several-ovulate; female zone of spadix free from spathe; spathe without a transverse septum separating male and female zones. 76. Flowers of both sexes always present in a single inflorescence; male flowers 1-androus; lower spathe margins connate; leaf blade ovate or sagittate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 86. Arisarum 76. Flowers of both sexes sometimes present in a single inflorescence, but more often with male and female flowers appearing in separate inflorescences; male flowers 2–5-androus; lower spathe margins convolute; leaf blade normally trisect, pedatisect or radiatisect, rarely simple and ovate · · · · · 98. Arisaema 75. Ovary 1-ovulate; female zone of spadix adnate to spathe; spathe usually with transverse septum between male and female zones · · · · · · · · · · · 97. Pinellia 39. Spadix entirely enclosed by spathe in a basal “kettle” formed of connate spathe margins, plants always helophytic or aquatic 77. Female flowers spirally arranged (pseudo-whorl in Lagenandra nairii, whorled in L. gomezii) and free; spathe tube “kettle” with connate margins occupying entire spathe tube; spathe blade usually opening only slightly by a straight ot twisted slit; berries free, opening from base; leaf ptyxis involute · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 56. Lagenandra 77. Female flowers in a single whorl, connate; spathe tube kettle occupying only lower part of spathe tube, remainder also with connate margins (except Cryptocoryne spiralis), blade spreading or twisted; berries connate into a syncarp which opens from the apex; leaf ptyxis convolute · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57. Cryptocoryne 38. Stamens of each male flower entirely connate into a distinct synandrium, synandrium rarely reduced to single stamen (Colletogyne endemic to Madagascar) or stamens free and basally connate with remote globose thecae (Gorgonidium endemic to Andean South America), or only filaments connate and then stigma stellate and 5–8-lobed (Spathantheum) 78. Laticifers simple 79. Synandria connate, thecae of adjacent synandria encircling pits in the spadix, each pit with a somewhat prominent upper margin; leaf peltate; Burma to India · · · · · · · · 99. Ariopsis 79. Synandria free; leaf not peltate; Africa, Madagascar or Americas 80. Higher order leaf venation parallel-pinnate or if reticulate then stem a creeping rhizome and plant from Amazonia (Bognera) 81. Ovules anatropous; primary lateral veins of leaf forming a single marginal vein, no submarginal collective vein present; plant from tropical America or continental tropical Africa. 82. Female zone of spadix free; plant from tropical west and central Africa · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 48. Anubias 82. Female zone of spadix entirely adnate to spathe; plant from tropical America. 83. Female flowers each with whorl of several staminodes; higher order leaf venation strictly parallel-pinnate · · · · · · · · · · · · · · · · · 35. Dieffenbachia 83. Female flowers without staminodes; higher order leaf venation reticulate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 36. Bognera 81. Ovules orthotropous to hemi-orthotropous; primary lateral veins of leaf forming submarginal collective vein and 1–2 marginal veins; plants from temperate eastern North America or Madagascar. 84. Giant herbs (to 4m) with massive pseudostem of petiole sheaths; staminodes of female flower free; Madagascar (also naturalized in Pemba, Zanzibar and Mascarene Is.) · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 85. Typhonodorum 84. Relatively small herbs (less than 1m) without pseudostem; staminodes of female flower connate into a cup-like synandrodium; eastern North America · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 84. Peltandra 80. Higher order leaf venation reticulate; stem usually a subglobose tuber, if rhizomatous then plant Madagascan 85. Madagascan plants; seed lacking endosperm; ovary 1-locular; leaf venation with primary lateral veins forming submarginal collective vein and 1-2 marginal veins on each side of blade 86. Stamens either completely connate with marginal thecae or only partially connate by filaments; bisexual flowers often present between male and female zones of the spadix · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 82. Carlephyton

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86. Stamens completely connate into truncate synandria or synandria reduced to one stamen 87. Synandria reduced to one stamen, thecae apical on conical filament; spadix fertile to apex; leaf blade always cordate · · · · · · · · · · · · 83. Colletogyne 87. Thecae apical on a truncate synandrium; spadix appendix present or not; leaf blade cordate, hastate, trifid, trisect or pedatifid · · · · · · · · 81. Arophyton 85. South American plants; seed with abundant endosperm; ovary with more than 1 locule (except Spathicarpa); leaf venation with primary lateral veins usually forming single marginal vein on each side, submarginal collective veins usually absent 88. Spadix free or only female zone adnate to spathe 89. Ovules anatropous 90. Ovules 2 per locule; leaf blade entire, linear to subsagittate; spadix, with terminal appendix of synandrodes · · · · · · · · · · · · · · · · 37. Mangonia 90. Ovules 1 per locule; leaf blade entire, pinnatifid to subdracontioid; spadix fertile to apex 91. Leaf blade pinnatifid to bipinnatifid or subdracontioid; synandria elongate; stigma capitate or lobed; staminodes of female flowers free (connate in Taccarum caudatum) · · · · · · · · · · · · · 38. Taccarum 91. Leaf blade usually pinnatifid, rarely entire; synandria short and domed; stigma deeply lobed; staminodes of female flowers free or connate · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39. Asterostigma 89. Ovules orthotropous 92. Styles and synandria elongate; leaf blade pinnatifid or -sect or bipinnatifid or entire and ± cordate 93. Staminodes in female flowers filiform to subclavate; synandria with free filament apices or not; leaf blade pinnatifid, pinnatisect or bipinnatifid · · · · · · · · · · · · · · · · · · · · · · · · · · · 40. Gorgonidium 93. Staminodes in female flowers elongate-triangular; synandria entirely connate; leaf blade entire, ± cordate · · · · · · 41. Synandrospadix 92. Styles and synandria short and squat or synandria very flat; staminodes in female flowers obovate or trapezoid; leaf blade pedatisect or subpalmatifid · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42. Gearum 88. Spadix usually entirely adnate to spathe (male zone free in Spathantheum intermedium) 94. Ovary 6–8-locular; female flowers below, male above; leaf blade entire or pinnately lobed · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43. Spathantheum 94. Ovary 1-locular; female and male flowers intermixed (2 central rows of male flowers, 2 outer rows of female flowers); leaf blade entire · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 44. Spathicarpa 78. Laticifers anastomosing 95. Plants climbing hemiepiphytes, sometimes creeping on ground in submature growth; internodes long; berries connate into a syncarp · · · · · · · · · · · · · · · · · · · · · · · 67. Syngonium 95. Plants terrestrial or geophytic, rarely aquatic, not climbing; internodes very short; berries free from each other 96. Spadix without an appendix (present in Hapaline appendiculata, included here, occasionally absent in Colocasia esculenta, excluded here) 97. Ovary completely to incompletely 2- to several-locular with deeply intrusive parietal placentas (1-locular with basal placenta in Jasarum, Scaphispatha and a few species of Caladium and Xanthosoma); neotropical plants 98. Helophytes or terrestrial; leaf blade ovate, sagittate to hastate or pedatifid 99. Pollen shed in tetrads; style usually laterally thickened or expanded into a diaphanous mantle; leaf blade rarely peltate, sometimes trifid or -sect, or pedatifid or -sect 100. Spathe tube subglobose, inflated; female zone of spadix free; styles normally discoid (laterally swollen) and coherent (except Xanthosoma plowmanii); synandrodes (sterile flowers) between male and female flowers well-developed, ± prismatic · · · · · · · · · · · 65. Xanthosoma

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100. Spathe tube narrow, elongate; female zone of spadix mostly adnate to spathe; stylar region thin, spreading, diaphanous, mantle-like; synandrodes (sterile flowers) betweeen male and female flowers usually irregular or fungiform, not prismatic · · · · · · · · · · 66. Chlorospatha 99. Pollen shed in monads; stylar region not laterally expanded; leaf blade usually peltate, rarely trisect, never pedatifid or -sect 101. Spathe tube always convolute; stylar region as broad as ovary (Caladium paradoxum has discoid, coherent stylar regions); synandrodes (sterile flowers) between male and female flowers well-developed, prismatic; placentas 1–2(–3), parietal; seeds several (rarely 1–2) · · · 63. Caladium 101. Spathe tube gaping widely at anthesis; style much narrower than ovary; synandrodes (sterile flowers) lacking, male and female zones contiguous; placenta 1, basal; seed solitary · · · · · · · · · · · 62. Scaphispatha 98. Submerged aquatics; leaf blade linear · · · · · · · · · · · · · · · · · · · · · 64. Jasarum 97. Ovary clearly 1-locular, placentas not intrusive; palaeotropical plants 102. Ovules more than 1; leaf blade peltate 103. Female flowers with staminodes; spathe not constricted; stem trunk-like or creeping · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 101. Steudnera 103. Female flowers without staminodes; spathe with 1 or 2 constrictions; stem tuberous, producing erect or spreading stolons bearing small tubercles covered in hooked scales · · · · · · · · · · · · · · · · · · · · · · 102. Remusatia 102. Ovule solitary; leaf blade not peltate · · · · · · · · · · · · · · · · · · · · 68. Hapaline 96. Spadix with an appendix (occasionally absent in Colocasia esculenta); palaeotropical plants 104. Leaf blade pedatisect to radiatisect; female flowers each with several large staminodes · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 100. Protarum 104. Leaf blade entire or pinnatifid; female flowers without staminodes (except single small ones in Colocasia esculenta) 105. Placentas parietal; ovules many; leaf blade always entire · · · 103. Colocasia 105. Placenta basal; ovules few; leaf blade entire or pinnatifid · · · 104. Alocasia

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26 D E S C R I P T I O N S O F T H E T R I B E S A N D G E N E R A O F A R A C E A E

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I. Subfamily Gymnostachydoideae Subfamily Gymnostachydoideae Bogner & Nicolson in Willdenowia 21: 37 (1991). Laticifers absent; stem a short rhizome; leaves distichous, bifacial (dorsiventrally flattened), linear, not differentiated into blade and petiole, primary veins parallel; flowering shoot a complex synflorescence consisting of 3–6(7), short, perennating floral sympodia, borne on an erect peduncular axis, each sympodium axillary to a bract and composed of numerous inflorescences; spathe inconspicuous; spadix long-stipitate; flowers bisexual, perigoniate, 2-merous; tepals 4, fornicate, stamens 4, free, thecae dehiscing by longitudinal slit, pollen monosulcate; ovary 1-locular, ovule 1, orthotropous, placenta apical, stigma small; berries deep blue, long-exserted beyond tepals; seed obovoid, testa absent at maturity, embryo axile, endosperm copious.

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1. Gymnostachys Gymnostachys R. Brown, Prodr. 337 (1810). TYPE: G. anceps R. Brown HABIT: acaulescent herbs, stem a short, thick subterranean rhizome. LEAVES: distichous, bifacial (dorsiventrally flattened), somewhat plicate, linear, not differentiated into blade and petiole, margins erose-serrate; midrib not differentiated, primary lateral veins parallel, somewhat prominent, higher order venation parallel. INFLORESCENCE: borne on a long scape, terminating in 3–6(–7), short, perennating floral sympodia separated from each other by a distinct peduncular axis, each sympodium composed of several spadices and borne axillary to a leaf-like bract. SCAPE: subequal to leaves, alate on each side, margins erose-serrate. SPATHE: a simple, short, keeled, inconspicuous bract, ± alternating with the short, linear-lanceolate, 2-keeled prophylls of each article of

the floral sympodium. SPADIX: short- to long-stipitate, cylindric, many-flowered, erect in flower, pendent in fruit. FLOWERS: bisexual, perigoniate, often somewhat distant; tepals 4, ± as long as wide, fornicate, imbricate. STAMENS: 4, free, filaments somewhat flattened, anthers short, connective slender, thecae ellipsoid, dehiscing by a longitudinal slit, connective inconspicuous. POLLEN: monosulcate, ellipsoid, medium-sized (mean 31 µm., range 30–33 µm.), exine foveolate to slightly fossulate, apertural exine fossulate-verrucate. GYNOECIUM: ovary oblong, 1-locular, ovule 1, orthotropous, funicle very short, placenta apical, stylar region shortly attenuate, stigma small, subhemispheric. BERRY: ellipsoid, deep blue, apiculate, 1-seeded, strongly projecting beyond tepals. SEED: ellipsoid to obovoid, testa absent at maturity, embryo axile, ± elongate, endosperm copious. See Plates 1, 107A. CHROMOSOMES: 2n = 48. DISTRIBUTION: 1 sp.; Australia (Queensland, New South Wales). ECOLOGY: southern temperate to subtropical or rarely tropical rainforest; understorey plant in moist hardwood forests, most common on cool southerly slopes and in cool moist gullies. ETYMOLOGY: Greek gymnos (naked) and stachys (spike). TAXONOMIC ACCOUNTS: Engler (1905), Shelton (1980), Elliot & Jones (1990), Hnatiuk (1990), Hay (1993c).

II. Subfamily Orontioideae

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Subfamily Orontioideae Mayo, Bogner & P.C. Boyce, Genera of Araceae p. 346 (1997). Laticifers absent (except Orontium); helophytes, stem a stout, erect rhizome, continuation shoot in axil of last leaf preceding spathe; petiole not geniculate apically, sheath long; leaf blade entire, oblong-elliptic, higher order venation reticulate; flowers bisexual, perigoniate; tepals free, fornicate, stamen with distinct filament, anther terminal, connective slender, thecae dehiscing by longitudinal slit, pollen monosulcate; ovary ± immersed in spadix axis; seed testa smooth, thin or absent at maturity, embryo large, endosperm very sparse to absent.

2. Orontium

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Orontium L., Sp. Pl. 324 (1753). TYPE: O. aquaticum L. SYNONYMS: Amidena Adanson, Fam. 2: 470 (1763); Aronia J. Mitchell, Diss. Gen. Pl. 28 (1769), nom. rej.

1. Gymnostachys

Laticifers present, simple, articulated. HABIT: seasonally dormant, aquatic herbs, continuation shoot of stem sympodium arising in axil of last foliage leaf of each article and beginning with a foliage leaf rather than a cataphyll, rhizome short, erect, buried deep in soil. LEAVES: several. PETIOLE:

ORONTIOIDEAE : ORONTIUM

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Plate 1. Gymnostachys. A, habit × 1/6; B, base of plant × 2/3; C, detail of leaf venation showing erose-serrate midrib and margin x 5; D, cross-section of leaf × 5; E upper part of inflorescence × 1/3; F, detail of peduncle showing erose margin × 5; G, spadix × 1; H, detail of spadix × 10; J, tepal, side view × 20; K, flower with perigone removed × 20; L, gynoecium, longitudinal section × 20; M, berry × 2. Gymnostachys anceps: A, Dransfield s.n. (Kew slide collection); B, Cunningham 337 (K); C–D, Forster 4851 (K); E, Brown s.n. (K); F–M, Covery 10432 (Kew spirit collection 29047.718).

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Plate 2. Orontium. A, habit × 1/5; B, habit × 2/3; C, detail of leaf venation × 5; D, detail of spadix × 5; E, flower × 10; F, stamen, adaxial view × 10; G, gynoecium, longitudinal section × 10; H, fruit × 4; J, fruit, longitudinal section × 4. Orontium aquaticum: A, Boyce s.n. (Kew slide collection); B–C, Curtis s.n. (K); D–G, Mayo s.n. (Kew spirit collection 46576) (K); H, Hooker s.n. (K); J, Bogner 1896 (Kew spirit collection 56428).

ORONTIOIDEAE : ORONTIUM

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ETYMOLOGY: ancient Greek orontion referred to an unknown plant used for treating jaundice; according to some authors named after the River Orontes in Syria. TAXONOMIC ACCOUNTS: Huttleston (1953), Klotz (1991, 1993).

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3. Lysichiton Lysichiton Schott in Oesterr. bot. Wochenbl. 7: 62 (1857). TYPE: L. camtschatcensis (L.) Schott (“camtschatcense”; Dracontium camtschatcense L.) SYNONYMS: Arctiodracon A. Gray in Mem. Amer. Acad. Arts, ser. 2, 6: 408 (1859); [Lysichitum Schott in Oesterr. bot. Wochenbl. 7: 62 (1857), orth. var.]

2. Orontium

terete, sheath long. BLADE: oblong-elliptic, held above water surface or floating on it; primary lateral veins arising at base of blade, arcuately ascending and running into apex, secondary laterals inconspicuous and parallel to primaries, higher order venation transverse-reticulate. INFLORESCENCE: 1–2 in each floral sympodium. PEDUNCLE: absent, spadix supported on elongated stipe. SPATHE: a short, inconspicuous, simple bract, inserted at point of attachment of inflorescence to rhizome, enclosing young spadix, later separated from it and disintegrating. SPADIX: held above water level, conical, slender, flowering acropetally, golden yellow, peduncle-like stipe very long, often partly submerged, greenish to red-brown with white, swollen apical part. FLOWERS: bisexual except for some male flowers at spadix apex, perigoniate, tepals 6, sometimes 4, short, about as long as wide, fornicate, subtruncate, irregularly imbricate. STAMENS: 6, sometimes 4, free, filaments flattened, connective slender, thecae ellipsoid, dehiscing by broad apical slit. POLLEN: monosulcate, ellipsoid, large (mean 64 µm., range 55–73 µm.), exine densely foveolate-fossulate. GYNOECIUM: ovary depressed-globose, broader than long, 1-locular, ovule 1, hemianatropous, held horizontally, funicle short, placenta basal, stylar region ± absent, stigma small, subsessile, discoid-hemispheric. BERRY: depressed-globose, green, 1-seeded, partially immersed in spadix axis. SEED: ± globose, testa thin, smooth, ± transparent, decaying at maturity, raphe conspicuous in furrow-like depression, hilum reddish, embryo large, almost globose to depressed-globose, with a small internal cavity situated below the hilum, outer cell layers green, plumule well-developed with 1(–2) filiform leaf primordia, endosperm absent. See Plates 2, 107B. CHROMOSOMES: 2n = 26. DISTRIBUTION: 1 sp.; temperate E. North America:– USA (Alabama, Carolinas, Connecticut, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Massachusetts, Mississippi, New Jersey, New York, Pennsylvania, Rhode Is., Tennessee, Virginia, West Virginia). ECOLOGY: temperate wetlands, occurring from sea level to 900m alt.; partly submersed aquatic plant growing in open or partially shaded swamps, marshes, lakes and ponds, in dense stands or scattered colonies.

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HABIT: large herbs with thick, hypogeal rhizome. LEAVES: numerous. PETIOLE: thick, flattened to deeply sulcate, sheath long. BLADE: becoming large, ovate-elliptic to ovate-oblong; primary lateral veins pinnate, running into inconspicuous marginal vein, higher order venation ± regularly transversereticulate between primaries. INFLORESCENCE: solitary, appearing just before or with the leaves in early spring. PEDUNCLE: absent, spadix supported on elongated stipe. SPATHE: marcescent, inserted at the point of attachment of spadix stipe to rhizome, cucullate to boat-shaped, lower part narrowly convolute and clasping spadix stipe, blade expanded, ovate-elliptic, white or yellow. SPADIX: subcylindric, subacute at apex, stipe extremely elongated. FLOWERS: bisexual, perigoniate; tepals 4, cucullate, imbricate. STAMENS: 4, free, filaments oblong, flattened, connective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 40 µm., range 38–43 µm.), exine reticulate. GYNOECIUM: elongate-ovoid, attenuate apically, ovary immersed in spadix axis, 2–locular or incompletely 2-locular, ovules 1–2 per locule, orthotropous, broad, conoid, funicle very short and thick, placenta axile, thick, stylar region attenuate, longer than tepals, stigma discoid-hemispheric. BERRY: ellipsoid, mostly 2-seeded, green, decaying or stylar portion breaking off with perigone at maturity to expose seeds embedded in spadix axis. SEED: ± ellipsoid, somewhat compressed, testa smooth, embryo large, ellipsoid, endosperm nearly absent, present only as a single cell layer. See Plates 3, 107C.

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Plate 3. Lysichiton. A, flowering habit × 1/3; B, leaf × 1/3; C, detail of junction of spathe, stipe and rhizome, longitudinal section × 2; D, peduncle surrounded by spathe base, transverse section × 2; E, flowering habit × 1/10; F, vegetative and fruiting habit × 1/8; G, detail of spadix × 3; H, flower × 6; J, flower, longitudinal section × 6; K, seed, adaxial view × 3; L, seed, side view × 3; M, seed, abaxial view × 3. Lysichiton americanus: A-D, Cult. Kew 1969–18002; E, Ballard 364 (Kew slide collection); F, Townsend 87/142 (Kew slide collection); G–J, Cult. Jodrell (Kew spirit collection 29047.78 & 29047.764); K–M, Townsend 87/127 (Kew spirit collection 51789).

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CHROMOSOMES: 2n = 28. DISTRIBUTION: 2 spp.; temperate E. Asia and W. North America:– Canada (British Columbia), Japan, Russian Far East (Kamchatka, Kuril Is., Okhotsk, Sakhalin, Udsk), USA (Alaska, California, Idaho, Montana, Oregon, Washington). ECOLOGY: temperate wetlands; helophytes, in swamps and wet woodlands. ETYMOLOGY: ancient Greek lysis (free) and chiton (tunic). TAXONOMIC ACCOUNTS: Krause (1908), Hultén & St. John (1931,1956), Huttleston (1953, 1955), Nicolson (1981).

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4. Symplocarpus Symplocarpus R.A. Salisbury ex Nuttall in W. Barton, Gen. 1: 105 (1817), (see Taxon 29: 601, Phytologia 72: 80–92) nom. cons. TYPE: S. foetidus (L.) Nuttall (Dracontium foetidum L.) SYNONYMS: Ictodes Bigelow in Amer. Med. Bot. 2: 41 (1818); Spathyema Rafinesque, Fl. Tell. 4: 13 (1838, “1836”); [Symplocarpos J.A. Schultes & J.H. Schultes, Mant. 3: 16 (1827), orth. var.] HABIT: fairly large, seasonally dormant herbs, rhizome thick, erect with thick roots. LEAVES: few. PETIOLE: fairly broad, sulcate, sheath short. BLADE: subcordate- to cordate-ovate; primary lateral veins pinnate, arching towards apex, running into inconspicuous marginal vein, secondary laterals and higher order venation reticulate to transverse-reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing before or with leaves. PEDUNCLE: only shortly exserted above ground. SPATHE: thick, boat-shaped or conchiform, lower part convolute, upper part somewhat to widely gaping, apex 2-keeled, rostrate, curving forwards. SPADIX: stipitate, subglobose, hidden within spathe. FLOWERS: bisexual, perigoniate; tepals 4, fornicate, imbricate. STAMENS: 4, free, filaments flattened, connective slender, thecae oblong, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 33 µm.), exine reticulate, apertural exine

4. Symplocarpus

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coarsely verrucate. GYNOECIUM: ovary somewhat immersed in spadix axis, 1-locular, ovule 1, orthotropous, funicle very short, placenta apical-parietal, stylar region long-attenuate, stigma punctate-discoid. BERRY: tepals and style persistent to ripe fruiting stage, seed and base of berry immersed in spongy spadix axis, infructescence ± globose. SEED: globose, testa thin, smooth, embryo globose, endosperm very sparse, only a single cell layer thick. See Plates 4, 107D. CHROMOSOMES: 2n = 30, 60 (28). DISTRIBUTION: 3 spp.; temperate E. Asia and E. North America:– Canada (New Brunswick, Nova Scotia, Quebec, Ontario), China, Japan, Korea N. & S., Russian Far East, USA (Connecticut, Delaware, Indiana, Illinois, Iowa, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Is., Tennessee, Virginia, West Virginia, Wisconsin). ECOLOGY: temperate damp woodlands, rarely open wetlands, from near sea level to ca. 900m. alt.; usually in shaded sites, frequent near water courses; S. foetidus and S. renifolius flower in early spring before the leaves appear and fruit in the summer of the same year; S. nipponicus flowers after the leaves appear and the fruits ripen in the following spring. ETYMOLOGY: Greek syn- (together), -plo (folded) and karpos fruit); refers to immersion of gynoecia and fruits in spadix axis. TAXONOMIC ACCOUNTS: Huttleston (1953), Lee (1985).

III. Subfamily Pothoideae Subfamily Pothoideae Engler in Nova Acta Acad. Leopold.Carol. 39: 140 (1876). Laticifers absent; stem usually aerial; petiole geniculate apically; higher order leaf venation reticulate; spathe simple, spreading to reflexed, not enclosing spadix; flowers bisexual, perigoniate; stamen filament distinct, anther terminal, thecae dehiscing by longitudinal slit, connective inconspicuous.

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Plate 4. Symplocarpus. A, habit showing vertical rhizome, longitudinally sectioned × 2/3; B, flowering habit × 1/3; C, leaf × 2/3; D, detail of leaf venation × 5; E, inflorescence, nearside half of spathe removed × 1; F, detail of spadix × 3; G, flower × 10; H, flower, nearside tepals removed × 10; J, gynoecium, longitudinal section × 10; K, infructescence × 1; L, seed, side view × 2; M, habit × 1/3. Symplocarpus foetidus: A, Herb Gray 7/82 (K) & Cult. Kew 1969–18003 (Kew spirit collection 51367); B, Cult. Kew 1969–18003 (Kew slide collection); C–D, W.D. s.n. (K); E–L, Cult. Kew 1969–18003 (Kew spirit collection 51367, 51616 & 55681); S. renifolius: M, Furuse 8737 (K).

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Tribe Potheae

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Tribe Potheae Engler in Nova Acta Acad. Leopold.-Carol. 39: 140 (1876, Pothoeae). Laticifers absent; shrubby climbing herbs with tough woody stems, main shoot monopodial, flowering shoots axillary or infra-axillary; leaves distichous; petiole sheath long and often broad, flattened and apically auriculate (reduced in Pedicellarum and Pothos series Goniuri); flowers bisexual, perigoniate, (2–)3-merous; tepals (4–)6, stamens 6, free, pollen monosulcate; ovules 1 per locule, anatropous, stigma sessile, usually umbonate; embryo large, endosperm absent.

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5. Pothos Pothos L., Sp. Pl. 968 (1753). TYPE: P. scandens L. SYNONYMS: Tapanava Adanson, Fam. 2: 470 (1763); Batis Blanco, Fl. Filip. 791 (1837); Goniurus Presl, Epim. Bot. 244 (1851, “1849”); [Potha O. Kuntze, Rev. Gen. 2: 742 (1891), orth.var.] Trichosclereids occasionally present. HABIT: climbing herbs, stems rather woody, lower branches rooting, upper ones free and hanging, nodes rarely bearing short, clustered spines (P. armatus), buds of lateral shoots sometimes perforating the leaf sheath or ± infra-axillary. LEAVES: distichous, juvenile plants of some species with shingle form. PETIOLE: geniculate (articulate) apically, either broad, completely flattened and usually auriculate apically, or morphology normal with a long sheath, sometimes sheath reduced to a pair of hyaline ridges (series Goniuri). BLADE: linear-lanceolate to ovate or elliptic, sometimes oblique; primary lateral veins either mostly arising near base of blade, long arcuate, and running into marginal vein near apex, or primary lateral veins pinnate, weakly differentiated, forming submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate in all types. INFLORESCENCE: axillary or infra-axillary, solitary or forming short branching systems of several inflo-

5. Pothos

98

THE GENERA OF ARACEAE

rescences, bearing 4–6 (sometimes more, e.g. P. insignis) rigid, coriaceous cataphylls at the base. PEDUNCLE: short to long, sometimes reflexed. SPATHE: ovate to linear, rarely very long (P. mirabilis). SPADIX: globose, ovoid, cylindric, ellipsoid or obovoid, sessile to long-stipitate, densely or laxly flowered. FLOWERS: bisexual, perigoniate; tepals 4–6, usually fornicate, free or partially to completely connate (e.g. P. rumphii). STAMENS: 4–6, free, filaments oblong, flattened, connective slender, thecae ellipsoid, dehiscing by slit. POLLEN: monosulcate, ellipsoid-oblong, small (mean 21 µm., range 16–25 µm.), exine foveolate to reticulate or subrugulate, muri psilate or minutely tuberculate. GYNOECIUM: ovary ovoid-oblong or depressed, (2?–)3-locular; ovules 1 per locule, anatropous, funicle short, placenta axile at base of septum, stylar region sometimes as broad as ovary, stigma discoid-hemispheric to umbonate. BERRY: ellipsoid to ovoid, 1–3-seeded, red. SEED: ellipsoid, testa smooth, embryo large, endosperm absent. See Plates 5, 108A. CHROMOSOMES: 2n = 24, 36. DISTRIBUTION: ca. 70 spp.; south and southeast Asia, Australasia, Malagasy region, Malay Archipelago:– Australia (New South Wales, Queensland), Bangladesh, Brunei, Burma, Cambodia, China (Guandong, Guangxi, Guizhou, Hainan, Hunan, Sichuan, Taiwan, Yunnan), Comores Is., India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Japan (Ryukyu Is.), Laos, Madagascar, Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, Philippines, Solomon Is., Sri Lanka, Thailand, Vanuatu, Vietnam. ECOLOGY: tropical humid forest; usually climbing hemiepiphytes often in regrowth forest, rarely on rocks. NOTES: Engler (1905) recognized 2 sections:– sect. Pothos (with 4 series), sect. Allopothos (with 3 series). ETYMOLOGY: modified spelling of a Sinhalese vernacular name “potha”; P. scandens is still known as “pota-wel” in Sri Lanka. TAXONOMIC ACCOUNTS: Engler (1905), Sivadasan (1982), Nicolson (1988a), Sivadasan, Mohanan & Rajkumar (1989), Boyce & Poulsen (1994), Boyce & Nguyen (1995).

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B

C

D E A

F

M

J S H P R

G

N

K

L

Q

Plate 5. Pothos. A, habit × 2/3; B, detail of leaf venation × 5; C, detail of spadix × 10; D, gynoecium, transverse section × 10; E, juvenile habit × 2/3; F, detail of spadix × 6; G, portion of flowering stem × 2/3; H, detail of petiole venation × 5; J, detail of leaf venation × 5; K, flowering shoot showing developing flagelliform shoot × 2/3; L, infructescence × 1; M, flower, top view × 10; N, flowering shoot × 2/3; P, detail of leaf venation × 5; Q, infructescence × 1; R, flower, top view × 10; S, gynoecium, longitudinal section × 10. Pothos beccarianus: A–D, Sumbing Jimpin SAN 110325 (K); P. motleyanus: E, Burbidge s.n. (K); P. barberianus: F, UNESCO (Kostermans) 192 (K & Kew spirit collection 58012); P. scandens: G–J, Cult. Kew (Kew spirit collection 19158); Soedarsono 280 (K); P. junghuhnii: K–M, Warnham s.n. Cult. Kew (K); P. rumphii: N–S, Forman 240 (K); Sands s.n., Cult Kew 1970–02267 (Kew spirit collection 29047.171).

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6. Pedicellarum Pedicellarum M. Hotta in Acta Phytotax. Geobot. 27 (34): 61 (1976). TYPE: P. paiei M. Hotta HABIT: climbing, shrubby herb, branches distichously leaved, branching from below the nodes, juvenile shoots with shingle form, flowering branches producing flagelliform shoots. LEAVES: several. PETIOLE: sometimes pubescent in young leaves, geniculate at apex, sheathed for the most part. BLADE: lanceolate to narrowly elliptic (nearly orbicular in shingle form), acuminate; primary lateral veins pinnate, forming submarginal collective vein, 1–2 distinct marginal veins also present, higher order venation reticulate. INFLORESCENCE: arising below the nodes, bearing several short cataphylls at base of peduncle. PEDUNCLE: shorter than spadix, subequal to or longer than petiole, very slender. SPATHE: small, fully expanded, ovate-cordate, membranaceous. SPADIX: sparsely and very laxly flowered, long-stipitate, axis flexuose, minutely hispid-papillose. FLOWERS: bisexual, perigoniate, on short pedicels, receptacle large and conspicuous; tepals connate, forming cup-like structure. STAMENS: 6, free, filaments broad, flattened, connective slender, thecae ellipsoid, latrorse, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid-oblong, small (mean 17 µm.), exine reticulate, muri minutely tuberculate. GYNOECIUM: obpyramidal, excavated at apex, ovary 3-locular, ovules 1 per locule, anatropous, placenta axile at base of septum, stigma sessile, umbonate. BERRY: obovoid, 1–3-seeded, red. SEED: compressed-ellipsoid, testa smooth, thin, embryo large, endosperm absent. See Plates 6, 108B. CHROMOSOMES: unknown. DISTRIBUTION: 1 sp.; Indonesia (Borneo), Malaysia (Borneo). ECOLOGY: tropical humid forest; climbing hemiepiphyte. NOTES: Appears closely related to Pothos series Goniuri, in which the spadix also has a slender axis and the flowers are distant. ETYMOLOGY: Latin pes, pedis (foot), -ella (diminutive) and Arum; refers to the unique character of pedicellate flowers. TAXONOMIC ACCOUNTS: Nicolson (1984b).

7. Pothoidium

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6. Pedicellarum

7. Pothoidium Pothoidium Schott, Aroideae 6: 26, t. 57 (1857) & in Oesterr. bot. Wochenbl. 7: 70 (1857). TYPE: P. lobbianum Schott HABIT: climbing herb, stems somewhat woody, flowering branches free and hanging. LEAVES: distichous, many. PETIOLE: oblong, entirely flattened, resembling blade, venation parallel, joined to blade by constricted articulation. BLADE: much shorter than petiole, triangular-lanceolate; midrib absent, no primary veins differentiated, veins parallel, running into apex. INFLORESCENCE: several to many, borne in a terminal branching system, lower inflorescences axillary to a foliage leaf, upper ones either subtended by cataphyll or without subtending leaf. PEDUNCLE: peduncular axis slender, composed of one to several internodes, sometimes subtended by a prophyll, sometimes also bearing a cataphyll ± halfway up. SPATHE: occurrence irregular, often absent, linear-lanceolate, widely spreading, margins usually revolute. SPADIX: apparently often functionally unisexual, cylindric, sessile to long-stipitate when subtended

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D

E

B

A

C

J

H

G

F

Plate 6. Pedicellarum. A, habit × 2/3; B, petiole × 3; C, detail of petiole base × 5; D, detail of leaf venation × 5; E, juvenile shingle habit × 1; F, inflorescence × 2; G, flower × 10; H, gynoecium, transverse section × 10; J, infructescence × 2. Pedicellarum paiei: A–D, F, Paie SAN 16354 (K, L); E, Boyce 782 (K & Kew slide collection); G, Lassan SAN 107216 (K); H, Church et al. 303 (A, Kew spirit collection 58609); J, Lee S 54080 (L).

P OT H O I D E A E : P E D I C E L L A R U M

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C D

B

E G

F

A

Plate 7. Pothoidium. A, habit × 2/4; B, detail of leaf blade and petiole venation × 2; C, infructescence × 1; D, detail of basal portion of spadix × 5; E, flower, top view × 10; F, flower, nearside tepal removed × 10; G, gynoecium, longitudinal section. Pothoidium lobbianum: A–B, de Vogel 3866 (K); C, Loher 7047 (K); D, Merrill 2293 (K & Kew spirit collection 58025); E–G, Herb. Lugd. Bat. (K).

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by spathe. FLOWERS: apparently usually unisexual, sometimes bisexual, perigoniate, male flowers with well developed anthers and apparently sterile ovary, female flowers with large fertile ovary and lacking stamens; tepals 6, fornicate, membranaceous. STAMENS: 3–6, free, number often varying on single spadix, filaments elongated and overtopping perigone at anthesis, oblong-triangular, flattened, connective slender, thecae short, ellipsoid, dehiscing by broad slit. POLLEN: monosulcate, ellipsoid-oblong, medium-sized (mean 26 µm.), exine reticulate with psilate muri, apertural exine shallowly fossulate or verrucate. GYNOECIUM: broadly ovoid to subglobose or obovoid, ovary 1-locular, ovule 1, anatropous, funicle short, placenta subbasal, stylar region attenuate, stigma discoid-hemispheric. BERRY: ellipsoid to ovoid, apiculate (stigma remnant), prominently exserted when mature, red. SEED: ellipsoid, testa smooth, embryo large, endosperm absent. See Plate 7. CHROMOSOMES: 2n = 24. DISTRIBUTION: 1 sp.; southeast Asia, Malay Archipelago:– China (Taiwan), Indonesia (Moluccas, Sulawesi, Sumatra), Philippines. ECOLOGY: tropical humid forest; climbing hemiepiphyte. ETYMOLOGY: Pothos and Greek -idion (special); implies a plant distinct from Pothos. TAXONOMIC ACCOUNTS: Engler (1905).

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Tribe Anthurieae Tribe Anthurieae Engler in Nova Acta Acad. Leopold.Carol. 39: 140 (1876).

rotting to a fibrous mass (net-fibrous), sometimes completely disappearing. PETIOLE: geniculate apically (geniculum rarely well below blade, e.g. A. oerstedianum), variously shaped in cross-section, sheath long in juvenile (monopodial) leaves, very short in sympodial leaves. BLADE: small to very large (exceeding 2m), usually coriaceous, more rarely membranaceous or stiff and brittle, extraordinarily variable in shape, linear to orbicular in outline, rarely peltate, entire to trifid or trisect, or pedatifid or subpalmatifid, or pedatisect to radiatisect, rarely the lobes or segments themselves pinnately lobed, blade base cuneate to cordate, sagittate or hastate; primary lateral veins pinnate or more rarely all arising at the base, usually forming one or more submarginal collective veins, basal ribs often present in cordate leaves, higher order venation reticulate. FLOWERING BRANCHES: sympodial units usually comprising one 2-keeled prophyll, one 1-keeled cataphyll, one foliage leaf and terminal inflorescence. INFLORESCENCE: always solitary. PEDUNCLE: usually rather elongated, rarely short. SPATHE: usually persistent, sometimes marcescent or deciduous, usually linear to linear-lanceolate, more rarely elliptic to ovate, broadly cordate to suborbicular, erect, spreading or reflexed. SPADIX: sessile to long-stipitate, usually cylindric to conic, more rarely clavate, rarely globose, very short to very long (over 1m). FLOWERS: bisexual, perigoniate; tepals 4, fornicate, in 2 decussate whorls. STAMENS: 4, free, filaments somewhat flattened, usually equalling tepals at anthesis, sometimes exceeding them, anthers short, connective slender, thecae ovate to oblongovate, dehiscing by longitudinal slit. POLLEN: forate (most often 3–4 pores), more rarely diporate, rarely inaperturate (sect. Polyphyllium), spherical to subspheroidal, small (mean 22 µm., range 14–29 µm.), exine foveolate to retic-

Laticifers absent; climbers, hemiepiphytes, epiphytes, lithophytes or terrestrial herbs, sympodial units of stem each normally composed of prophyll, cataphyll, foliage leaf, inflorescence (often aborted); petiole of sympodial leaves with very short sheath, geniculate apically; primary lateral veins pinnate (rarely all arising at petiole insertion) in blade of entire leaves and in lobes of compound leaves, usually forming one or more submarginal collective veins; spathe usually simple, spreading, reflexed or erect, linear to elliptic or ovate; flowers bisexual, perigoniate; tepals 4, fornicate, stamens 4, free, pollen usually forate (periporate) with 3–4 pores, more rarely diporate, rarely inaperturate (sect. Polyphyllium); ovary 2-locular, ovules 1–2 (rarely more) per locule, anatropous to campylotropous, placenta axile-subapical; mature berries exserted and usually dangling; seed often sticky, endosperm copious.

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8. Anthurium Anthurium Schott, in Wiener Z. Kunst 1829 (3): 828 (1829). LECTOTYPE: A. acaule (Jacquin) Schott (Pothos acaulis Jacquin, see Britton & Wilson, Sci. Surv. Porto Rico 5: 128. 1923). SYNONYMS: Podospadix Rafinesque, Fl. Tell. 4: 821 (1838, “1836”); Strepsanthera Rafinesque, Fl. Tell. 4: 13 (1838, “1836”). HABIT: evergreen herbs, stem erect, creeping, or short- to long-climbing, rarely rhizomatous, internodes very short (plant rosulate) to elongated. LEAVES: prophylls and cataphylls usually ± persistent, entire (membranaceous) or

8. Anthurium

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C

D

F E

B

A

H

J

G

Plate 8 (i). Anthurium. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2; H, leaf × 1/2; J, leaf × 1/2. Anthurium friedrichsthalii: A, Lehmann s.n. (K); A. vallense: B, Nee & Hale 9628 (K); A. carnosum: C, Valerio 181 (K); A. melastomatis: D, McPherson 7678 (K); A. smithii: E, Hahn & Grifo 3323 (K); A. antrophyoides: F, Lehmann 787 (K); A. peltigerum: G, Madison et al. 4599 (K); A. clidemioides: H, Stevens 24526 (K) & Bown 131/37 (Kew slide collection); A. puberulinervium: J, Croat 55033 (K).

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C

B

A

D

E

Plate 8 (ii). Anthurium. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2. Anthurium watermaliense: A, Croat 36713 (K); A. rimbachii: B, Hepper 6445 (K); A. trisectum: C, Bogner 1088 (K); A. longissimum: D, Kalbreyer 853 (K); A. polyschistum: E, Paterson 121 (K).

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A

B

C

D

Plate 8 (iii). Anthurium. A, habit × 1/16; B, habit × 1/8; C, habit × 1/2; D, habit × 1/2. Anthurium salviniae: A, Cult. Kew 1961–66404; A. oerstedianum: B, Cult. Kew 1981–3725; A. interruptum: C, Davidse & Herrera 31343 (K); A. melastomatis: D, McPherson 7678 (K).

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A

B

C

D E

Plate 8 (iv). Anthurium. A, habit × 1/3; B, habit × 1/8; C, habit × 1/16; D, habit × 1/3; E, habit × 1/6. Anthurium radicans: A, Cult. Kew. 1977–5366; A. andraeanum: B, Cult. Kew 1963–49801; A. warocqueanum: C, Cult. Kew. 1986–6028; A. affine: D, Harley et al. 19429 (Kew slide collection); A. wendlingeri: E, Cuadros et al. 3950 (K) & Nee et al. 8735 (K).

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J

K

F A

H

G M

P L

B

C

D

E

N

Plate 8 (v). Anthurium. A, inflorescence × 2; B, inflorescence, spathe mostly removed × 2; C, flower × 15; D, stamen, three quarter view × 15; E, gynoecium, longitudinal section × 15; F, infructescence × 1; G, inflorescence, spathe mostly removed × 2; H, flower × 15; J, stamen × 15; K, gynoecium, longitudinal section × 15; L, infructescence × 2; M, developing berry surrounded by persistent perigone × 8; N, inflorescence × 1; P, flower × 10. Anthurium globosum: A, Croat 67966 (K & Kew spirit collection 58085); A. regale: B–E, Cult. Kew 1962–67111 (Kew spirit collection 51391); A. scandens subsp. scandens: F, Cult. Kew 1984–8010 (Kew spirit collection 52038); A. polyschistum: G–K, Cult. Kew 1976–1533 (Kew spirit collection 51386); A. sp.: L–M, (Kew spirit 59068); A. radicans: N–P, Cult. Kew. 1977–5366 & (Kew spirit collection 58028).

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ulate or subrugulate, rarely tuberculate, muri ± psilate or spinulose, apertural exine mostly psilate, rarely spinulose. GYNOECIUM: ovary ovoid to oblong or obovoid, 2-locular, ovules 1–2 per locule, rarely more, anatropous, hemianatropous or subcampylotropous, funicle short, placenta axile near apex of septum, stylar region inconspicuous to attenuated; stigma small, subcapitate, secreting conspicuous nectar droplet at anthesis. BERRY: variously shaped from globose to elongate-fusiform, when mature exserted from tepals and usually held dangling by tiny strips of inner tepal epidermis, sometimes simply falling out of spadix, 2–4seeded (more in sect. Tetraspermium), variously coloured, from con-spicuous reds and oranges to dull purplish green, white or blueish. SEED: ± oblong to ellipsoid or subglobose, sometimes curved, testa usually smooth or somewhat verrucose, thin, usually with sticky gelatinous mass adhering to raphe, small strophiole sometimes present, embryo axile, subcylindric to conoid, sometimes curved, endosperm copious. See Plates 8i–v, 108C. CHROMOSOMES: 2n = 30, 60, 90 (20, 24, 28, 40, 48, 56, 84) DISTRIBUTION: over 800 spp.; tropical America, West Indies:– Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad & Tobago, Uruguay, Venezuela. ECOLOGY: tropical humid forest, especially diverse in cloud forests; climbing hemiepiphytes, terrestrial on forest floor, epiphytes, lithophytes, rarely helophytes or rheophytes. NOTES: 19 sections are recognized by Croat & Sheffer (1983):– Tetraspermium, Gymnopodium, Porphyrochitonium, Pachyneurium, Polyphyllium, Leptanthurium, Oxycarpium, Xialophyllium, Polyneurium, Anthurium (syn. Urospadix), Episeiostenium, Digitinervium, Cardiolonchium, Chamaerepium, Calomystrium, Belolonchium, Semaeophyllium, Schizoplacium, Dactylophyllium. ETYMOLOGY: Greek anthos (flower), oura (tail) and -ion (diminutive). TAXONOMIC ACCOUNTS: Engler (1905), Madison (1978c), Croat (1980), Mayo (1982), Croat & Sheffer (1983), Croat (1984, 1986), Rodriguez (1987, 1989), Croat (1991), Sakuragui (1994).

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IV. Subfamily Monsteroideae Subfamily Monsteroideae Engler in Nova Acta Acad. Leopold.-Carol. 39: 142 (1876). Laticifers absent; trichosclereids (H- or T- shaped) abundant (except Anadendreae, Heteropsideae; in Amydrium present only in certain parts); terrestrial, climbing hemiepiphytes or more rarely epiphytes; petiole geniculate apically, sheath usually long; leaf blade never sagittate, base narrowed to subcordate, sometimes pinnatifid or perforated, outline always ± oblong-ovate to -elliptic or narrower, often oblique; spathe expanded or boat-shaped, not constricted centrally; spadix fertile to apex; flowers bisexual, perigone present or absent, 2-merous (2–3-merous in Spathiphylleae).

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Tribe Spathiphylleae Tribe Spathiphylleae Engler in Engler & Prantl, Nat. Pflanzenfam. II (3): 112, 121 (1887). Laticifers absent; tissues with small trichosclereids occurring in bundles; leaves sometimes distichous, petiole sheath long; blade oblong, cuspidate-acuminate, higher order venation parallel-pinnate; inflorescence solitary; spathe persistent or marcescent; flowers bisexual, perigoniate, 2–3-merous; tepals free or connate, thecae dehiscing by longitudinal slit, pollen inaperturate, exine striate; ovules anatropous to hemianatropous; seed ± oblong, narrowed towards micropyle, endosperm copious.

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9. Spathiphyllum Spathiphyllum Schott in Schott & Endlicher, Melet. Bot. 22 (1832). TYPE: S. lanceifolium (Jacquin) Schott (“lancaefolium”; Dracontium lanceaefolium Jacquin) SYNONYMS: Hydnostachyon Liebmann in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1849: 23 (1849); Massowia K. Koch in Bot. Zeitung (Berlin) 10: 277 (1852); Spathiphyllopsis J.E. Teysmann & S. Binnendijk, Natuurk.

9. Spathiphyllum

MONSTEROIDEAE : SPATHIPHYLLUM

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Tijdschr. Ned.-Indië 25: 400 (1863); Amomophyllum Engler in Gard. Chron., ser. 2, 7: 139 (1877, non Watelet 1866); [Massovia Bentham & J.D. Hooker, Gen. Pl. 3: 998 (1883), orth. var.]. Trichosclereids present. HABIT: evergreen herbs usually with short, erect to creeping stem, appearing acaulescent, sometimes stoloniferous, occasionally erect and climbing (S. solomonense). LEAVES: several. PETIOLE: geniculate apically, sheath long. BLADE: oblong to elliptic or narrowly elliptic, cuspidate-acuminate; primary lateral veins pinnate, running into marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation transverse-reticulate. INFLORESCENCE: solitary. PEDUNCLE: subequal to or longer than petiole. SPATHE: oblong, elliptic, ovate or obovate, cuspidate-acuminate, ± decurrent at insertion, membranaceous to subcoriaceous, fully expanded, rarely fornicate or clasping, persistent, with distinct midrib and pinnate primary lateral veins, usually white, rarely green, turning green in fruit. SPADIX: usually stipitate, rarely sessile, stipe often partially adnate to spathe, spadix cylindric, erect, shorter than spathe. FLOWERS: bisexual, perigoniate; tepals 4–6, free, fornicate and almost truncate at apex, or partly or completely connate into a truncate cup. STAMENS: 4–6, free, filaments short, oblong, flattened, connective slender, thecae oblong-ellipsoid to ovoid, dehiscing by longitudinal slit. POLLEN: inaperturate, ellipsoid to ellipsoid-oblong, medium-sized (mean 32 µm., range 27–41 µm.), exine striate. GYNOECIUM: ovoid, subcylindric, obovoid or flask-shaped, ovary 3-locular, more rarely 2- or 4-locular, ovules 2, 4, 6 or 8 per locule, anatropous to hemianatropous, placenta axile, stylar region usually long, conic and longexserted beyond perigone, sometimes shortly attenuate, sometimes ± truncate and not exserted and inconspicuous, stigma 2–3-lobed or subcapitate to punctiform. BERRY: rounded, ovoid to obovoid, or conically attenuate apically, 1–8-seeded, greenish. SEED: oblong, ellipsoid to ovoid or slightly curved and ± reniform, pale yellow to brown, funicle short, testa sparsely foveolate, otherwise smooth or verrucose, embryo axile, elongate, slightly curved, endosperm copious. See Plates 9, 108D. CHROMOSOMES: 2n = 30, 60. DISTRIBUTION: 41 spp.; tropical America, West Indies, eastern Malay Archipelago, Melanesia:– Belize, Brazil (Amazonia, Central-West, Northeast), Colombia, Costa Rica (incl. Isla del Coco), Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Indonesia (Irian Jaya, Moluccas, Palau Is., Sulawesi), Mexico, Nicaragua, Panama, Papua New Guinea

10. Holochlamys

(incl. New Britain, New Ireland), Peru, Philippines, Solomon Is., Surinam, Trinidad, Venezuela. ECOLOGY: tropical humid forest, rarely cloud forest; forest floor, in wet sites, sometimes on rocks in streams, rarely hemiepiphytic (S. solomonense). NOTE: Bunting (1960a) recognized 4 sections:– sect. Spathiphyllum, sect. Dysspathiphyllum, sect. Amomophyllum, sect. Massowia; S. solomonense has recently been placed in a separate section, sect. Chlaenophyllum (Nicolson 1994). ETYMOLOGY: Greek spathe (spathe) and phyllon (leaf). TAXONOMIC ACCOUNTS: Engler & Krause (1908), Bunting (1960a), Nicolson (1968b, 1994), Williams & Dressler (1976).

10. Holochlamys Holochlamys Engler in Beccari, Malesia 1: 265 (1883). TYPE: H. beccarii (Engler) Engler (Spathiphyllum beccarii Engler). Trichosclereids present. HABIT: evergreen herbs, stem short, upright. LEAVES: several. PETIOLE: geniculate apically. BLADE: oblong-elliptic, or ovate to lanceolate, often oblique, apex cuspidate to acuminate, base attenuate to rounded; primary lateral veins pinnate, running into marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation forming transverse cross connections, often obscured. INFLORESCENCE: solitary. PEDUNCLE: shorter than petiole. SPATHE: white, with distinct midrib and pinnate primary lateral veins, tightly clasping the spadix, marcescent after anthesis, gradually decomposing. SPADIX: sessile to shortly stipitate, cylindric, fertile to apex. FLOWERS: bisexual, perigoniate; tepals 4, fornicate apically and ± truncate, connate into a truncate cup. STAMENS: 4, free, filaments short, oblong, subequal to anthers, connective slender, thecae oblong, dehiscing by longitudinal slit. POLLEN: inaperturate, ellipsoid, mediumsized (mean 33 µm., range 32–34 µm.), exine striate. GYNOECIUM: subcylindric to ovoid, ovary 1-locular, ovules many, anatropous, funicle long, placenta basal, stylar region cylindric, ± as broad as ovary, stigma oblong or 3–4-lobed. BERRY: 1–few-seeded. SEED: irregularly oblongellipsoid, narrowed towards micropyle, testa minutely verrucose or smooth, embryo elongate, endosperm copious. See Plate 10. CHROMOSOMES: 2n = 60. DISTRIBUTION: 1 sp.; Indonesia (Irian Jaya), Papua New Guinea (incl. New Britain). ECOLOGY: tropical humid forest; on forest floor, particularly along small streams, on river banks or on rocks. ETYMOLOGY: Greek holos (whole) and chlamys (short mantle, cloak); refers to the connate tepals of the flowers. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Hay (1990a).

Tribe Anadendreae Tribe Anadendreae Bogner & French in Taxon 33(4): 689 (1984). Laticifers and trichosclereids absent; climbing hemiepiphytes; leaves distichous; petiole sheathed almost to apex; finer venation reticulate; peduncle relatively long; spathe boat-shaped to reflexed, longer than spadix, marcescent or deciduous soon after anthesis; flowers bisexual, perigone membrana-

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Plate 9. Spathiphyllum. A, habit × 1/5; B, leaf × 2/3; C, detail of leaf venation × 5; D, inflorescence × 2/3; E, detail of spadix × 3; F, flower × 6; G, flower, perigone removed × 6; H, gynoecium, longitudinal section × 6; J, inflorescence × 2/3; K, detail of spadix × 3; L, flower with developing fruit, longitudinal section × 6; M, fruit, transverse section × 6; N, inflorescence × 2/3; P, detail of spadix × 3; Q, flower × 6; R, flower, perigone removed × 6; S, gynoecium, longitudinal section × 6. Spathiphyllum ‘Clevelandii’ : A, Cult Kew 1962–43301 (Kew slide collection); S. laeve: B–D, Dressler 4465 (K); S. cannifolium: E–H, Jermy 2872, Cult. Kew 1976–56863 (Kew spirit collection 37366); S. humboldtii: J–K, Cremers 7813 (K); L–M, Jonker et al. 5845 (K); S. cochlearispathum: N–S, Cult. Kew 1972–68267 (Kew spirit collection 29047.83).

MONSTEROIDEAE : SPATHIPHYLLUM

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Plate 10. Holochlamys. A, habit × 1/6; B, leaf × 2/3; C, detail of leaf venation × 5; D, inflorescence, lower portion of peduncle removed × 1; E, detail of spadix × 5; F, flower × 10; G, flower, perigone removed × 10; H, gynoecium, longitudinal section × 15. Holochlamys beccarii: A–C, Cult. Kew 1970–1474; D, Sands 889 (Kew spirit collection 34304); E–H, Sands s.n. (Kew spirit collection 56122).

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ceous, urceolate, shorter than gynoecium; stamens 4, thecae dehiscing by longitudinal slit, pollen inaperturate; gynoecium truncate apically, ovary 1-locular, ovule 1, hemianatropous, placenta basal, stigma transversely oblong or subspheroid; embryo large, endosperm absent.

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11. Anadendrum Anadendrum Schott in Bonplandia 5: 45 (1857). LECTOTYPE: A. montanum Schott (see Engler in De Candolle, Monogr. Phan. 2: 97, 250 (1879)). SYNONYMS: [Anadendron Schott in Oesterr. bot. Wochenbl. 7: 118 (1857), orth. var.]. Trichosclereids absent. HABIT: climbing herbs. LEAVES: distichous. PETIOLE: geniculate apically, sheathed nearly to apex, sheath persistent or marcescent. BLADE: obliquely ovateoblong, entire; primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–3 in each floral sympodium. PEDUNCLE: relatively long. SPATHE: oblong-ovate, boat-shaped to reflexed, greenish white, rostrate apically and overtopping the spadix, deciduous after anthesis. SPADIX: stipitate, cylindric. FLOWERS: bisexual, perigoniate; perigone membranaceous, a single cup-like structure, truncate, equalling or shorter than gynoecium. STAMENS: 4, free, filaments relatively short, broad, spathulate, connective slender, thecae linear-elliptic, dehiscing by longitudinal slit. POLLEN: inaperturate, subspheroidal, small (mean 22 µm.), exine psilate or subretipilate, pilae spinulose tipped and solitary, or united into groups of 2–4 or more. GYNOECIUM: ovary obconic or obpyramidal, subquadrangular, 1-locular, ovule 1, ana-tropous, funicle short, placenta basal, stylar region as broad as ovary, stigma transversely oblong. BERRIES: distinctly truncate apically, subglobose, orange red (A. microstachyum). SEED: rounded, subglobose, testa smooth, glossy, embryo large, endosperm absent. See Plate 11. CHROMOSOMES: 2n = 60.

DISTRIBUTION: ca. 7 spp.; southeast Asia, Malay Archipelago:Brunei, Cambodia, China (Guandong, Hainan, Yunnan), Indonesia (Borneo, Java, Sulawesi, Sumatra), Laos, Malaysia (Borneo, Peninsula), Philippines, Thailand, Vietnam. ECOLOGY: tropical humid forest; climbing hemiepiphytes, sometimes on rocks. ETYMOLOGY: Greek ana (up) and dendron (tree), a tree climber. TAXONOMIC ACCOUNTS: Engler (1905), Bogner & French (1984).

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Tribe Heteropsideae Tribe Heteropsideae Engler in Engler, Pflanzenreich 21 (IV.23B): 20 (1905). Laticifers and trichosclereids absent; climbing hemiepiphytes, main shoot monopodial, flowering articles usually short, axillary; leaves distichous; petiole usually almost entirely adnate to succeeding internode, leaving only apical geniculum free; blade oblong to lanceolate, primary and secondary lateral veins parallel-pinnate, forming submarginal collective vein, higher order venation reticulate; spathe boat-shaped, marcescent or deciduous soon after anthesis; flowers bisexual, perigone absent; stamens 4, thecae dehiscing by apical slit, pollen zonate or dicolpate; gynoecium truncate, ovary incompletely 2-locular, ovules 2 per locule, anatropous, placenta axile at base of partial septum, stigma oblong or rounded; endosperm absent.

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12. Heteropsis Heteropsis Kunth, Enum. Pl. 3: 59 (1841). LECTOTYPE: H. salicifolia Kunth (see Nicolson in Taxon 24: 468. 1975). Trichosclereids absent. HABIT: evergreen climbing herbs with woody-fibrous roots. LEAVES: numerous. PETIOLE: usually very short, entirely geniculate, concave and somewhat flat-

11. Anadendrum

MONSTEROIDEAE : HETEROPSIS

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Plate 11. Anadendrum. A, habit × 2/3, B, detail of leaf venation × 5; C, juvenile habit × 5; D, detail of spadix × 5; E, flower × 6; F, flower, perigone removed × 6; G, gynoecium, longitudinal section × 6. Anadendrum microstachyum: A, Cult. Kew 1982–4984, Scortechini 82 (K); B, de Wilde 14630 (K); C, ‘Native collector’ 2398 (K); D–F, Cult. Kew 1982–4984; G, Burkill 3231 (K).

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Plate 12. Heteropsis. A, habit × 2/3; B, detail of leaf venation × 5; C, fruiting habit × 2/3; D, juvenile habit × 2/3; E, spadix × 2; F, detail of spadix × 5; G, flower × 10; H, gynoecium, longitudinal section × 10; J, berry × 2; K, berry, longitudinal section × 2; L, berry, transverse section × 2. Heteropsis spruceana: A–B, Traill 1135 (K); H. oblongifolia: C, E–L, Bell et al. 88–178 (K); Gentry & Young 31969 (K); Pinheiro & Santos 2266 (K) & Santos 1276 (K); H. cf. jenmanii: D, Harley et al. 17848 (K).

MONSTEROIDEAE : HETEROPSIS

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ETYMOLOGY: Greek heteros (different) and opsis (appearance); this aroid is different from all others in that the petiole sheaths are adnate to the internodes in most species, leaving only a very short free portion of the petiole. TAXONOMIC ACCOUNT: Engler (1905).

Tribe Monstereae

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Tribe Monstereae Engler in Nova Acta Acad. Leopold.Carol. 39: 143 (1876).

12. Heteropsis

tened, sheath adnate to subtended internode, rarely petiole free with long sheath (e.g. H. melinonii). BLADE: oblong, elliptic or lanceolate, ± long-cuspidate, subcoriaceous; primary lateral veins pinnate, forming a submarginal collective vein very close to margin, 1(-2) marginal veins also present, secondary laterals ± parallel to primaries, higher order venation reticulate. INFLORESCENCE: solitary, subtended by several small cataphylls, borne terminally on free, axillary branches. PEDUNCLE: very short. SPATHE: ovate-elliptic to ovate-oblong, cuspidate, convolute, opening at anthesis, then caducous. SPADIX: erect, free, stipitate, shorter than spathe, cylindric or ellipsoid. FLOWERS: bisexual, or lowermost flowers female by abortion of stamens, perigone absent. STAMENS: 4 or fewer by abortion, free, filaments short, flattened, rather broad, connective slender, thecae ovate-ellipsoid, overtopping connective a little, dehiscing by apical slit. POLLEN: zonate or dicolpate, ellipsoid-oblong or hamburger-shaped, medium-sized (mean 40 µm., range 37–42 µm.), exine foveolate or foveolate-fossulate, apertural exine psilate. GYNOECIUM: obpyramidal- prismatic, truncate, ovary incompletely 2-locular with conspicuous septal aperture, ovules 2 per locule, anatropous, collateral, placenta axile at base of partial septum, stylar region dense and thickened, broader than ovary, stigma very small, ellipsoid, oblong or subhemispheric. BERRY: shortly obovoid or obpyramidal, ± prismatic, stylar region forming a ± broad, flattened scar-like structure, 1-4 seeded, orange or greenish-white with brown apex. SEED: obovoid to ellipsoid, testa thin, smooth, black, shiny, embryo large, endosperm absent. See Plate 12. CHROMOSOMES: 2n = 28. DISTRIBUTION: ca. 13 spp.; tropical America:- Bolivia, Brazil (Amazonia, Atlantic region), Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Nicaragua, Panama, Peru, Surinam, Venezuela. ECOLOGY: tropical moist and humid forest; climbing hemiepiphytes, sometimes on rocks. NOTES: Heteropsis melinonii differs from other species of the genus in having free petiole sheaths, but agrees in ovary structure (incompletely 2-locular ovary with 2 ovules per locule), leaf venation and lack of trichosclereids.

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Laticifers absent; trichosclereids abundantly present (sparse in Amydrium), large, solitary and scattered in tissues; climbing hemiepiphytes and epiphytes; leaves distichous; petiole sheath usually long, almost equalling petiole (except Amydrium, Alloschemone); blade normally oblique; peduncle relatively short (except Stenospermation), spathe usually erect and boat-shaped (except Amydrium), marcescent or deciduous soon after anthesis; spadix equalling or shorter than spathe; flowers bisexual, perigone absent; stamens 4, thecae dehiscing by longitudinal slit not reaching base, pollen usually fully zonate, dicolpate; style well developed, rather massive, usually prismatic with ± truncate apex, as wide or wider than ovary, containing abundant trichosclereids, stigma ± hemispherical to linear, appearing sessile; stylar region of berry deciduous at maturity.

13. Amydrium Amydrium Schott in Ann. Mus. Bot. Lugduno-Batavum 1: 127 (1863). TYPE: A. humile Schott SYNONYMS: Epipremnopsis Engler, Pflanzenreich 37 (IV.23B): 1–3 (1908). Trichosclereids sparsely present in vegetative parts (petiole and sheath only, Seubert 1996b), more abundant in style (Carvell 1989). HABIT: evergreen herbs, often very robust, stem climbing or prostrate, usually producing long flagelliform shoots with reduced cataphylls. LEAVES: many, often remote from one another. PETIOLE: geniculate apically, sheath usually less than half as long as petiole. BLADE: ovatecordate or pandurate-trilobed or pinnatifid to pinnatisect, sometimes with ± numerous round to oval perforations near midrib; primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–several in each floral sympodium. PEDUNCLE: erect, subequal or half as long as petiole. SPATHE: conchiform to ovate, apiculate, sometimes reflexed at anthesis and then deciduous. SPADIX: sessile to long-stipitate, sometimes very short. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments short, broadly linear, anthers equalling or shorter than filaments, thecae ovoid, extrorse, dehiscing by longitudinal slit. POLLEN: fully zonate, hamburger-shaped, medium-sized (mean 39 µm., range 38–41 µm.), exine either densely and minutely punctate in one half and virtually psilate in other, or uniformly foveolate-fossulate, apertural exine psilate or obscurely verrucate. GYNOECIUM: obpyramidal or obconoid, tetragonal, ovary 1-locular, ovules 2, anatropous, funicle short, placenta near base of deeply intrusive septum, stylar region broader than ovary, slightly prominent centrally below stigma, otherwise ± truncate, stigma small, hemispheric. BERRY: subglobose, truncate to domed at apex, white (A. medium, A. humile) or orange-red (A. zippelianum)

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Plate 13. Amydrium. A, partial habit × 2/3; B, leaf × 1/2; C, tip of flagelliform shoot × 2/3; D, detail of spadix × 3; E, flower × 6; F, gynoecium, longitudinal section × 6; G, infructescence × 1; H, fruit, transverse section × 4; J, leaf × 1/2; K, detail of leaf venation × 5; L, seed, side view × 2; M, leaf × 1/2. Amydrium medium: A, Synge 35 (K); B, Foxworthy 4605 (K); C, Boyce 463, Cult. Kew 1989–3217; D–F, Motley 778 (K); G, Boyce 763 (Kew spirit collection 59090); H, Sam & Dewol s.n. (K); A. zippelianum: J–K, Ridley 26 (K); A. humile: L–M, de Wilde 13078 (K).

MONSTEROIDEAE : AMYDRIUM

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13. Amydrium

when ripe. SEED: subglobose to heart-shaped, testa smooth, glossy, embryo curved and partly green, endosperm present (E. Seubert 1993). See Plates 13, 109A. CHROMOSOMES: 2n = 60 DISTRIBUTION: 4–6 spp.; tropical southeast Asia, Malay Archipelago:– Brunei, Burma, China (Guandong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Sichuan, Yunnan), Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Malaysia (Borneo, Peninsula), Papua New Guinea, Philippines, Thailand, Vietnam. ECOLOGY: tropical humid forest; climbing hemiepiphytes on tree trunks or creeping on forest floor (A. humile). ETYMOLOGY: Greek amydron (obscure, faint) and ion (diminutive). TAXONOMIC ACCOUNTS: Nicolson (1968c), Carvell (1989a).

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14. Rhaphidophora Rhaphidophora Hasskarl in Flora 25 (2) Beibl. 1: 11 (1842). TYPE: R. lacera Hasskarl, nom. illeg. (Pothos pertusa Roxburgh, R. pertusa (Roxburgh) Schott). SYNONYMS: Afrorhaphidophora Engler in Engler & Prantl, Nat. Pflanzenfam. Nachtr. 3: 31 (1906); [Raphidophora Hasskarl, Cat. Hort. Bogor. 58 (1844), orth.var.]. Trichosclereids abundant. HABIT: evergreen, usually climbing herbs, more rarely repent, often extremely robust, climbing branches often thick, producing anchor and feeder roots, flagelliform shoots also produced, stem often square in cross section. LEAVES: many, distichous, juvenile shingle plants occur in some species. PETIOLE: geniculate

14. Rhaphidophora

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Plate 14. Rhaphidophora. A, habit × 1/2; B, leaf × 1/2; C, detail of leaf venation × 5; D, leaf × 1/2; E, leaf × 1/2; F, detail of spadix × 5; G, flower × 8; H, gynoecium, longitudinal section × 8; J, gynoecium, transverse section × 8; K, flower × 8; L, gynoecium, longitudinal section × 8; M, seed, side view × 20. Rhaphidophora foraminifera: A, Boyce 235 (Kew slide collection); de Wilde 14592 (K); de Wilde & de Wilde-Duyfjes 18154 (K). R. glauca: B–C, Henry 12728 (K); R. tenuis: D, Haviland & Hose 3605 (K); R. sylvestris: E, Mamit S 33623 (K); F–J, Cult. Kew 1967–460 (Kew spirit collection 35951); R. africana: K–L, Bogner 100 (Kew spirit collection 32065); R. decursiva: M, J. Arn. Arb. 57(4): 185–201, pl. I, 12 (1976).

MONSTEROIDEAE : RHAPHIDOPHORA

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Plate 15. Epipremnum. A, infructescences and associated stem × 1/2; B, leaf × 1/2; C, juvenile habit × 1/2; D, seed, side view × 5; E, inflorescence and subtending leaf × 1/2; F, detail of leaf venation × 5; G, detail of spadix × 3; H, flower × 5; J, gynoecium, longitudinal section × 5; K, gynoecium, transverse section × 5. Epipremnum pinnatum: A, Kostermans 18558 (K); B, Bloembergen 3827 (K); C, Powell & Chey 783 (K); D, J. Arn. Arb. 57(4): 185–201, pl. I, 3 (1976); E. nobile: E–K, Forman 289 (K & Kew spirit collection 6919).

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apically, sheath usually relatively long. BLADE: lanceolate or oblong, ± oblique, entire, perforate or pinnatifid to pinnatisect, often very large, lobes often subfalcately narrowed; primary lateral veins pinnate, running into marginal vein, often not differentiated from secondaries, secondary laterals ± parallel-pinnate, higher order venation reticulate. INFLORESCENCE: usually solitary, rarely more. PEDUNCLE: relatively short. SPATHE: boat-shaped, deciduous. SPADIX: subcylindric, conic, clavate, often extremely thick, sessile to stipitate, shorter than spathe. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments oblong-linear, anthers much shorter than filaments, connective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: dicolpate, extended monosulcate to perhaps fully zonate, ellipsoid or hamburger-shaped, medium-sized (mean 33 µm., range 24–55 µm.), exine foveolate, subreticulate, rugulate, fossulate, scabrate, retiscabrate, verrucate, or psilate. GYNOECIUM: obconic-prismatic to oblong, truncate, ovary 1- to partially 2-locular, ovules few to many, anatropous, funicle long, placentae parietal to basal, sometimes ± subaxile, partial septa variably intrusive, stylar region well developed, usually broader than ovary, usually truncate apically, rarely elongate-conic, stigma broadly elliptic or oblong and then transverse or longitudinal, or punctate-prominent. BERRY: usually many-seeded, stylar region deciduous at maturity, red or yellow. SEED: oblong, testa thin, smooth, embryo axile, straight, endosperm copious. See Plates 14, 109B. CHROMOSOMES: 2n = 60, 120 (42, 54, 56). DISTRIBUTION: ca. 120 spp.; tropical Africa, tropical southeast Asia, Malay Archipelago, Melanesia, Australasia, Pacific:– Australia (Queensland), Bangladesh, Bhutan, Brunei, Burma, Cambodia, Cameroon, Caroline Is., China (Fujin, Guandong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Xizang, Yunnan), Equatorial Guinea (Bioko, Rio Muni), Fiji, Gabon, Ghana, India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Palau Is, Sulawesi, Sumatra), Ivory Coast, Japan (Bonin Is., Ryukyu Is.), Laos, Liberia, Malaysia (Borneo, Peninsula), Nepal, New Caledonia, Nigeria, Papua New Guinea, Philippines, Samoa, Sierra Leone, Singapore, Solomon Is., Sri Lanka, Thailand, Togo, Uganda, Vanuatu, Vietnam. ECOLOGY: subtropical and tropical humid or rain forest or deciduous forest; climbing hemiepiphytes, rarely rheophytic (R. beccarii). ETYMOLOGY: Greek rhaphis, rhaphidos (needle) and pherô (I bear); refers to the macroscopic (to 1cm long), needle-like unicellular trichosclereids present in tissues. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Sivadasan (1982), Nicolson (1988a), Hay (1990a, 1993b).

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15. Epipremnum Epipremnum Schott in Bonplandia 5: 45 (1857). TYPE: E. mirabile Schott (1858). SYNONYM: Anthelia Schott in Ann. Mus. LugdunoBatavum 1: 127 (1863). Trichosclereids abundant. HABIT: evergreen climbing herbs, producing flagelliform shoots. LEAVES: several to many, distichous. PETIOLE: geniculate apically, sheath long, marcescent to deciduous, often decomposing to conspicuous net-fibrous mass. BLADE: entire, often oblique, lanceolate, elliptic, elliptic- oblong, or pinnatipartite to pinnatisect, rarely

15. Epipremnum

minutely perforate (E. pinnatum); primary lateral veins pinnate, running into marginal vein, secondary and often tertiaries parallel-pinnate, tertiary and higher order venation often reticulate. INFLORESCENCE: 1(–2) in each floral sympodium. PEDUNCLE: relatively short. SPATHE: boat-shaped, withering after anthesis, usually deciduous. SPADIX: subcylindric, conic, often quite thick, sessile or stipitate, shorter than spathe. FLOWERS: bisexual, or lowermost ones female, perigone absent. STAMENS: 4, free, filaments linear, somewhat broad, anthers much shorter than filaments, connective slender, thecae oblong-ellipsoid, dehiscing by longitudinal slit. POLLEN: fully zonate, hamburger-shaped, medium-sized (mean 40 µm., range 36–44 µm.), exine foveolate-fossulate, psilate at periphery, apertural exine coarsely verrucate. GYNOECIUM: ovary subtetragonal-prismatic, truncate, 1-locular, ovules usually 2, more rarely 4 or 6–8 (E. amplissimum), anatropous, funicle short, placenta parietal or near base of parietal partial septa, stylar region prismatic, as broad or broader than ovary, stigma umbonate to oblong-linear and longitudinal. BERRY: 1–8-seeded, stylar region deciduous at maturity. SEED: reniform, testa thickish, brittle, smooth, embryo curved, endosperm copious. See Plates 15, 109C. CHROMOSOMES: 2n = 60 (56, 84). DISTRIBUTION: 20 spp.; tropical southeast Asia, Australasia, Pacific:– Andaman Is., Australia (Queensland, Northern Territories), Brunei, Burma, Cambodia, Caroline Is., China (Guandong, Guangxi, Hainan, Taiwan,Yunnan), Fiji, Indonesia (Irian Jaya, Java, Moluccas, Sulawesi, Sumatra, Timor), Japan (Bonin Is., Ryukyu Is.), Malaysia (Borneo, Peninsula), Marshall Is., Papua New Guinea, Philippines, Singapore, Solomon Is., Sri Lanka, Thailand, Vanuatu, Vietnam. ECOLOGY: tropical humid forest; high-climbing hemiepiphytes, on trees and rocks. NOTES: Engler & Krause (1908) kept Epipremnum separate from Rhaphidophora because of the differences in seed structure. ETYMOLOGY: Greek prefix epi- (on) and premnon (bottom of tree trunk). TAXONOMIC ACCOUNTS: Engler & Krause (1908), Hay (1990a).

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Plate 16. Scindapsus. A, habit × 1/3; B, stigma, top view × 6; C, juvenile habit × 1/3; D, habit × 1/3; E, detail of leaf venation × 5; F, seed, side view × 2; G, habit showing fertile shoot and flagelliform shoot × 1/3; H, detail of spadix × 4; J, stigma, top view × 9; K, flower × 6; L, gynoecium, longitudinal section × 6; M, stigma, top view × 6; N, flower with window cut in gynoecium to show ovule × 6. Scindapsus rupestris: A, Chew & Corner 4253 (K); B, Beaman 9984 (K); S. pictus: C, Boyce 225 (K & Kew slide collection); S. officinalis: D–E, Kerr 20536 (K); F, J. Arn. Arb. 57(4): 185–201, pl. II, 31 (1976); S. beccarii: G, Boyce 318 (Kew slide collection); Cult. Kew 1965–47801 (Kew spirit collection 40953); H–L, Cult. Kew 1965–47801 (Kew spirit collection 40953); S. perakensis: M–N, Fedilis & Sumbing 68774 (K).

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16. Scindapsus Scindapsus Schott in Schott & Endlicher, Melet. Bot. 21 (1832). LECTOTYPE: S. officinalis (Roxburgh) Schott (Pothos officinalis Roxburgh; see Schott, Prodr. syst. Aroid. 395–397 (1860)). Trichosclereids abundant. HABIT: evergreen climbing herbs, sometimes very robust, sometimes producing flagelliform shoots, shoots with leaves evenly spaced or forming rosulate flowering zones separated by zones with elongated internodes and smaller leaves, or solitary leaf trapping bole epiphyte. LEAVES: many, juvenile plants often of shingle form. PETIOLE: geniculate apically, sheath usually broad, rarely decomposing to form persistent net-fibrous mass with abundant, stinging sclereids. BLADE: always entire, lanceolate, elliptic or ovate to obovate, acuminate, rarely variegated; primary lateral veins hardly differentiated, pinnate, running into marginal vein, secondaries and also sometimes tertiaries parallel-pinnate, higher order venation inconspicuous, transverse-reticulate. INFLORESCENCE: always solitary. PEDUNCLE: shorter than petiole. SPATHE: boat-shaped, gaping only slightly, caducous to deciduous. SPADIX: sessile to shortly stipitate, cylindric, narrowly ellipsoid or clavate, a little shorter than spathe. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments oblong, flattened, broadish, connective slender, thecae oblong-ellipsoid, dehiscing by apical slit. POLLEN: fully zonate, hamburger-shaped, medium-sized (mean 38 µm., range 33–45 µm.), exine shallowly and sparsely punctate, scabrate or nearly psilate. GYNOECIUM: ovary sometimes short, compressed ± cylindric, 1-locular, ovules 1(–2), anatropous, funicle short, placenta basal, stylar region well-developed, prismatic, truncate or with shortly conic central projection supporting stigma, stigma globose, elongate-globose, elliptic, linear, or punctiform. BERRY: stylar region deciduous when mature, red. SEED: rounded, subreniform, compressed, testa thickish, sparsely verruculose or smooth, embryo curved, endosperm present (Seubert 1993). See Plates 16, 109D. CHROMOSOMES: 2n = 60 (42, 56, 58, 64, 70, 112). DISTRIBUTION: ca. 36 spp.; tropical Asia, Malay Archipelago, Melanesia, Pacific:– Andaman Is., Australia (Queensland),

16. Scindapsus

Bangladesh, Bhutan, Brunei, Burma, Cambodia, Caroline Is., China, Fiji, India (Assam, Bengal, Sikkim), Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Laos?, Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, Philippines, Samoa, Solomon Is., Sri Lanka, Thailand, Vietnam. ECOLOGY: tropical humid forest or dry, deciduous or evergreen forest; climbing hemiepiphytes, also creeping over rocks, often terrestrial when juvenile, rarely rheophytic (S. rupestris). ETYMOLOGY: Greek skindapsos, once used for an ivy-like plant. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Bogner & Boyce (1994).

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17. Monstera Monstera Adanson, Fam. Pl. 2: 470 (1763), nom. cons. TYPE: M. adansonii Schott (Dracontium pertusum), typ. cons. SYNONYMS: Tornelia Gutierrez ex Schott, Gen. Aroid. t. 74 (1858); Serangium W. Wood ex R.A. Salisbury, Gen. Pl. Fragm. 5 (1866). Trichosclereids abundant. HABIT: evergreen climbing herbs. LEAVES: distichous, juvenile leaves sometimes of shingle plant form, rarely variegated. PETIOLE: geniculate apically, sheath usually long, persistent or decomposing to fibrous or membranous mass or entirely deciduous. BLADE: entire, oblique, oblong to ovate-elliptic, often conspicuously and elaborately perforated, more rarely deeply pinnatifid; primary lateral veins pinnate, running into marginal vein, rarely forming an irregular submarginal collective vein (M. obliqua), secondary laterals often parallel-pinnate, sometimes reticulated (e.g. M. dubia), higher order venation reticulate. INFLORESCENCE: 1–several in each floral sympodium. PEDUNCLE: shorter than petiole. SPATHE: ovate or oblong-ovate, cuspidate, boat-shaped and somewhat convolute basally, white to rose-coloured within, remaining open after anthesis, caducous. SPADIX: sessile, subcylindric, somewhat shorter than spathe. FLOWERS: bisexual, perigone absent, lowermost flowers usually sterile. STAMENS: 4, free, filaments flattened, connective slender, thecae oblongellipsoid, dehiscing by longitudinal slit. POLLEN: fully zonate,

17. Monstera

MONSTEROIDEAE : MONSTERA

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Plate 17. Monstera. A, leaf × 1/3; B, detail of leaf venation × 5; C, leaf × 1/3; D, gynoecium × 4; E, gynoecium, longitudinal section × 4; F, leaf × 1/3; G, infructescence × 1/3; H, inflorescence × 1/3; J, detail of spadix showing emerging stamens × 2; K, flower × 4; L, gynoecium, longitudinal section × 4; M, seed, side view × 3; N, flowering shoot × 1/3; P, juvenile shingle habit × 1/3. Monstera oreophila: A–B, Grayum et al. 6395 (K); M. subpinnata: C, Santos & Souza 1667 (K); D–E, Vasquez 1853 (K); M. adansonii var. laniata: F, Gomez 19565 (K); G, Whitmore 749 (Kew spirit collection 25790); M. lechleriana: H–L, Cult. Mason (Kew spirit collection 29047.113); M, J. Arn. Arb. 57(4): 185–201, pl. III, 40 (1976); M. tuberculata var. tuberculata: N, Mayo & Madison 341 (Kew spirit collection 29047.346) & Contr. Gray Herb. 207: 3–100, f. 63 (1977); P, Mayo & Madison 341 (K).

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hamburger-shaped, medium-sized (mean 48 µm, range 40–52 µm), exine densely foveolate to subreticulate, sparsely and shallowly foveolate or psilate, apertural exine verrucate or rugulate. STERILE FLOWERS: with 4 minute, conic staminodia, pistillode 2-locular, prismatic, lacking ovules. GYNOECIUM: obovoid to ellipsoid, prismatic, ovary 2-locular, ovules 2 per locule, anatropous, funicle short, placenta axile at base of septum; stylar region often massive, broader than ovary, apex truncate to shortly attenuate, stigma oblong-elliptic to linear and longitudinal or round. BERRY: 1–3-seeded, shedding prismatic stylar region at maturity, pulpy within. SEED: obovoid to ellipsoid, compressed, testa smooth, embryo large, endosperm absent. See Plates 17, 110A. CHROMOSOMES: 2n = 60 (24, 48, 56, 58, 70). DISTRIBUTION: ca. 40 spp. (T. Croat, pers. comm.); tropical America, West Indies:– Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Lesser Antilles, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad & Tobago, Venezuela. ECOLOGY: tropical moist and humid forest, cloud forest; climbing hemiepiphytes, usually on tree trunks, also on rocks or ground (M. deliciosa). NOTES: Madison (1977a) recognized four sections:– sect. Monstera, sect. Marcgraviopsis, sect. Echinospadix and sect. Tornelia. ETYMOLOGY: Latin monstrum (monster), refers to the peculiar perforations of the leaves of many species. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Madison (1977a).

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18. Alloschemone Alloschemone Schott, Gen. Aroid. 99 (1858). TYPE: A. poeppigiana Schott, nom. illeg. (Scindapsus occidentalis Poeppig, A.. occidentalis (Poeppig) Engler & Krause). Trichosclereids abundant. HABIT: evergreen climbing herbs, stem epidermis becoming distinctly corky and longitudinally furrowed with age. LEAVES: large, juvenile leaves entire, ovate to ovate-elliptic. PETIOLE: geniculate apically, thickened at base, sheath short. BLADE: pinnatifid, subcordate, lobes acute, falcate, 4–6 per side, primary lateral veins 1 per lobe, secondary laterals ± parallel-pinnate, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: shorter than petiole. SPATHE: ovate-cymbiform, deciduous. SPADIX: stipitate, cylindric, apex obtuse. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments flattened, free or connate, shorter than ovary, thecae oblong, dehiscing laterally by oblique, apical, pore-like slit. POLLEN: not fully zonate, ellipsoid, medium-sized (mean 46 µm), exine shallowly foveolate. GYNOECIUM: ovary prismatic, 1-locular, with abundant locular mucilage, ovule 1, amphitropous, funicle with trichomes, placenta basal, stylar region densely packed with trichosclereids, stigma sessile, elliptic. BERRY: unknown. SEED: unknown. See Plate 18. CHROMOSOMES: 2n = 84. DISTRIBUTION: 2 spp.; Brazil (Amazonas, Pará, Rondonia). ECOLOGY: tropical humid forest; climbing hemiepiphytes. NOTES: An incompletely known genus, which has been attributed sometimes to Asiatic Scindapsus. ETYMOLOGY: Greek allos (other), schema, schêmatos (form) and ône (being). TAXONOMIC ACCOUNTS: Engler & Krause (1908), Madison (1976a), Boyce & Bogner (in prep.).

18. Alloschemone

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19. Rhodospatha Rhodospatha Poeppig in Poeppig & Endlicher, Nov. Gen. Sp. 3: 91 (1845). LECTOTYPE: R. latifolia Poeppig (see Nicolson in Taxon 16: 518. 1967). SYNONYMS: Anepsias Schott, Gen. Aroid. t. 73 (1858); Atimeta Schott, Gen. Aroid. t. 71 (1858). Trichosclereids abundant. HABIT: evergreen, usually climbing herbs, producing flagelliform shoots. LEAVES: many, distichously arranged. PETIOLE: geniculate apically, sheath long, persistent to marcescent. BLADE: oblong-elliptic, ± oblique, always entire; primary lateral veins pinnate, numerous, running into ± distinct marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation transverse-reticulate. INFLORESCENCE: usually solitary. PEDUNCLE: shorter to longer than petiole. SPATHE: broadly ovate or oblong-ovate, abruptly cuspidate, yellowish white, cream, purplish or pink within, caducous after anthesis. SPADIX: long-stipitate to sessile, cylindric-conic, basal flowers sometimes sterile or female and scattered. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments linear-oblong, flattened, connective slender, thecae ovoid to ellipsoid, dehiscing by longitudinal slit. POLLEN: extruded in strands, fully zonate or inaperturate, hamburger-shaped or ellipsoid to oblong, medium-sized (mean 47 µm., range 34–57 µm.), exine densely to sparsely foveolate and nearly psilate to obscurely fossulate or verrucate. GYNOECIUM: compressed obconic to cylindric, ovary 2-locular, ovules usually numerous per locule, rarely few (R. venosa), anatropous to hemianatropous, funicle fairly long, placenta axile, rarely subbasal, stylar region well-developed, broader than ovary, prismatic, truncate to convex apically, stigma elliptic to linear, usually longitudinal. BERRY: cylindric-prismatic,

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Plate 18. Alloschemone. A, leaf × 1/3; B, detail of leaf venation × 5; C, leaf × 1/3; D, detail of leaf venation × 5; E, mature main stem showing corky epidermis × 1; F, inflorescence, part of spathe removed, remainder flattened out × 1/2; G, detail of spadix × 1; H, flower × 5; J, gynoecium, longitudinal section × 5. Alloschemone sp.: A–B, E, Madison et al. 6310 (K); A. occidentalis: C–D, F, H–J, Plowman et al. 12207 (NY); G, Krukoff 7162 (NY).

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Plate 19. Rhodospatha. A, habit × 1/2; B, flower × 8; C, gynoecium, longitudinal section × 8; D, seed × 20; E, habit × 1/2; F, detail of leaf venation × 5; G, detail of spadix × 5; H, flower × 8; J, gynoecium, longitudinal section × 8; K, gynoecium, transverse section × 16; L, flagelliform shoot × 1. Rhodospatha oblongata: A–C, Harley et al. 18234 (K); D, Harley et al. 18193 (K); R. rubropunctata: E, J. Arn. Arb. 57(4): 185–201, pl. I, 4 (1976); R. venosa: F–L, Montfort 155 (K).

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19. Rhodospatha

truncate, many- to few-seeded. SEEDS: rounded-reniform, flattened, testa brittle, very hard, smooth or with verrucose crest, embryo rather large, strongly curved, endosperm present but sparse. See Plates 19, 110B. CHROMOSOMES: 2n = 28, 56. DISTRIBUTION: ca. 75 spp. (T. Croat pers. comm.); tropical America:– Belize, Bolivia, Brazil (Amazonia, Atlantic region), Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad, Venezuela. ECOLOGY: tropical humid forest; climbing hemiepiphytes, true epiphytes (?) or sometimes on rocks, juvenile plants often on forest floor. ETYMOLOGY: Greek rhodon (rose, roseate) and spathê (spathe); refers to the spathe colour of some species. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Croat (1978).

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20. Stenospermation Stenospermation Schott, Gen. Aroid. t. 70 (1858). LECTOTYPE: S. mathewsii Schott (see Nicolson 1967). SYNONYM: [Stenospermatium Schott in Oesterr. bot. Zeitschr. 9: 39 (1859), orth. var.] Trichosclereids abundant. HABIT: epiphytic, climbing hemiepiphytic or terrestrial evergreen herbs, stem rather densely leaved, erect, often elongated. LEAVES: many. PETIOLE: geniculate apically, sheath long. BLADE: oblong-elliptic or lanceolate, oblique, often rather thick, venation usually obscured; midrib narrowly sulcate above, primary lateral veins weakly or not differentiated, pinnate,

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20. Stenospermation

running into margin, secondary laterals parallel-pinnate, higher order venation inconspicuous. INFLORESCENCE: solitary, often nodding. PEDUNCLE: relatively long. SPATHE: convolute, gaping at anthesis, boat-shaped or opening widely, white, caducous. SPADIX: usually stipitate, rarely sessile, cylindric. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments oblong, flattened, connective slender, thecae ovoid-ellipsoid, dehiscing by longitudinal slit. POLLEN: extruded in strands, fully zonate or inaperturate, hamburger-shaped or subspheroidal, medium-sized (mean 42 µm., range 30–58 µm.), exine psilate to shallowly and sparsely foveolate, fossulate-foveolate, verrucate or baculate. GYNOECIUM: compressed obconic to cylindric, ovary (1–)2-locular, ovules 4–many per locule, anatropous, arranged in 2 rows, funicles long, placenta basal, stylar region well-developed, usually broader than ovary and truncate, stigma elliptic to punctiform. BERRY: obovoid, truncate apically, locules 3–many-seeded, white, orange (S. ulei) to reddish orange, or yellow. SEED: clavate to ellipsoid, raphe prominent, testa thickish, smooth, embryo axile, elongate, endosperm copious. See Plates 20, 110C. CHROMOSOMES: 2n = 28. DISTRIBUTION: ca. 36 spp.; tropical America:– Bolivia, Brazil (Amazonia, Atlantic region), Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Nicaragua, Panama, Peru, Surinam, Venezuela. ECOLOGY: tropical humid forest, especially cloud forests; epiphytes or terrestrial on forest floor. NOTES: Vestigial tepals occasionally present (S. ulei). ETYMOLOGY: Greek stenos (narrow), sperma, spermatos (seed) and -ion (diminutive); refers to the slender seeds. TAXONOMIC ACCOUNT: Engler & Krause (1908), Pérez de Gómez (1983).

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Plate 20. Stenospermation. A, habit, inflorescence removed × 1/2; B, inflorescence × 1/2; C, gynoecium, longitudinal section; D, habit × 1/2; E, detail of leaf venation × 5; F, detail of spadix × 5; G, flower × 10; H, gynoecium, longitudinal section × 10; J, infructescence × 1; K, habit × 1/2. Stenospermation rusbyi: A–B, Rusby 2609 (K); S. ulei: C, E–H, Cult. Kew 1990–2738; D, Ule 8490 (K); S. angustifolium: J–K, Gómez et al. 20533 (K).

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V. Subfamily Lasioideae Subfamily Lasioideae Engler in Nova Acta Acad. Leopold.Carol. 39: 144 (1876); Hay (1992). Laticifers and trichosclereids absent; terrestrial or rooted aquatics, stem tuberous or rhizomatous, usually geophytic (except Lasia, Podolasia); petiole usually aculeate or warty or with striking coloration, long, usually ± geniculate apically; primary lateral veins of leaf blade divisions pinnate to arcuate-parallel, higher order venation reticulate; spadix flowering and fruiting in basipetal sequence; flowers bisexual, perigoniate (except Pycnospatha); tepals fornicate, ± truncate, free, in 2 or more whorls, stamens free (except Lasimorpha) with distinct filaments, anthers terminal, connective slender, pollen monosulcate; stigma capitate to subcapitate; embryo large, endosperm present (except Anaphyllum) and forming a thin but distinct layer.

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21. Dracontium Dracontium L., Sp. Pl. 967 (1753). LECTOTYPE: D. polyphyllum L. (Britton & Wilson 1923, p.130). SYNONYMS: Eutereia Rafinesque, Fl. Tell. 4: 12 (1838, “1836”); Echidnium Schott in Oesterr. bot. Wochenbl. 7: 62 (1857); Ophione Schott in Oesterr. bot. Wochenbl. 7: 101 (1857); Chersydrium Schott in Oesterr. bot. Zeitschr. 15: 72 (1865); Godwinia Seemann in J. Bot. 7: 314 (1869). HABIT: usually seasonally dormant, often robust, sometimes gigantic herbs, tuber hypogeal, depressed-subglobose, often bearing few to very numerous tubercles usually on upper surface. LEAVES: usually solitary. PETIOLE: long, often verrucose-asperate, sometimes bearing few to many longer, prickle-like processes, covered with striking, transversely banded variegation, usually weakly geniculate at apex, sheathed only at very base. BLADE: dracontioid, i.e trisect, sometimes perforate, primary anterior division usually trisect at middle and primary posterior divisions bisect below the middle, secondary and tertiary divisions further subdivided, ultimate lobes usually elliptic-acuminate, rarely linear (D. margaretae); primary lateral veins of ultimate lobes pinnate, forming arching submarginal collective veins, higher order venation reticulate. INFLORESCENCE: 1(–2), preceded by cataphylls, appearing before, with or after the leaf. PEDUNCLE: very short to long, epidermis similar to petiole. SPATHE: oblong, usually ± boat-shaped, erect, acute, cuspidate to acuminate, often fornicate, convolute basally, gaping above, sometimes pubescent within, marcescent to persistent, usually dark brown-purple outside, more reddish-purple within and often white at the base within. SPADIX: sessile to stipitate, shortly cylindric to ellipsoid, much shorter than spathe, apical flowers sometimes sterile or with enlarged, irregularly shaped, projecting structures. FLOWERS: bisexual, perigoniate; tepals 4–8, dilated apically, fornicate. STAMENS: usually 4–6, free, sometimes up to 19, usually longer than tepals at anthesis, filaments somewhat dilated, ± compressed, anthers longer than connective, connective slender, thecae oblong-ellipsoid, dehiscing by apical slit. POLLEN: extruded in a conglutinate mass, monosulcate, ellipsoid to oblong, medium-sized (mean 38 µm., range 29–48 µm.), exine foveolate-fossulate to subreticulate, apertural exine rugulate or verruculate. GYNOECIUM: ovary ovoid, incompletely 1–6locular, ovules 1 per locule, anatropous or

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21. Dracontium

subcampylotropous, funicle short, placenta axile to subbasal, stylar region as long or much longer than ovary, projecting beyond perigone, stigma small, subcapitate or slightly 3— lobed. BERRY: usually obpyramidal, bearing style rudiment, greenish to brown. SEED: rounded-reniform, a little compressed, testa verrucose to almost smooth, rather thick, embryo curved, endosperm present. See Plates 21, 110D. CHROMOSOMES: 2n = 26 DISTRIBUTION: 23 spp.: tropical America, West Indies:– Bolivia, Brazil (Amazonia, Central West), Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad, Venezuela. ECOLOGY: tropical moist and humid forest, rarely in savannas (D. margaretae); geophytes on forest floor. NOTES: Engler (1911) recognized 2 sections:– Sect. Dracontium, Sect. Godwinia; Sect. Echidnium is recognized by modern workers (G.H. Zhu pers. comm.). ETYMOLOGY: ancient name, Latin draco, dracontis (dragon, snake) and suffix -ium (diminutive), referring to the similarities of the petiole markings to a snake; also perhaps from the Greek drakontion. TAXONOMIC ACCOUNTS: Engler (1911), Bogner (1986a), Bunting (1986), Hay (1992a), Zhu (1996, 1997).

22. Dracontioides Dracontioides Engler in Pflanzenreich 48 (IV.23C): 36 (1911). TYPE: D. desciscens (Schott) Engler (Urospatha desciscens Schott). SYNONYM: Dracontium sect. Urospathopsis Engler in Bot. Jahrb. 5: 178 (1884). HABIT: unarmed herbs, sometimes robust (to 2m), rhizome subterranean, erect, sparsely branched, sometimes bearing a few subterranean tubercles. LEAVES: several, ± erect. PETIOLE:

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V

Plate 21. Dracontium. A, habit × 1/10; B, detail of petiole × 2/4; C, rachis/petiole insertion × 2/3; D, tubercules at soil–level junction of petiole and tuber × 2/3; E, leaf segment × 1/2; F, leaf segment × 1/2; G, seedling × 1; H, partially buried inflorescence × 2/3; J, flower, longitudinal section × 6; K, inflorescence × 1/5; L, inflorescence × 2/3; M, spadix × 1; N, detail of spadix × 3; P, flower, longitudinal section × 6; Q, spadix × 1; R, flower, longitudinal section × 6; S, leaf segment × 1/2; T, leaflet, transverse section × 1; U, infructescence × 1; V, berry, side view × 3; W, seed, side view × 4. Dracontium asperum: A–D, Cult. Kew 1977–5355; D. gigas: E, Cult. Bull 1878 (K) & Cult. Bull 1881 (K); D. spruceanum: F, Plowman 4490 (K); G, Plowman 4490 (Kew illustration collection); D. changuango: H, Boyce s.n.; J, Aristeguieta 12734 (Kew spirit collection 53986); D. prancei : K, Cult. Kew 1977–5372 (Kew slide collection); D. asperum: L, Cult. Kew 1979–3887 (Kew spirit collection 46575); M–P, Cult. Kew 1979–3887 (Kew spirit collection 29047.455); D. soconuscum: Q–R, Cult. Kew 1980–1628 (Kew spirit collection 51365); D. margaretae: S, Emmerich 4053 (K illustration collection); T–W, Emmerich 4053 (Kew spirit collection 29047.358 & 43972).

LASIOIDEAE : DRACONTIUM

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F

G

K

L

H

J

D

C

E

A

B

Plate 22. Dracontioides. A, habit × 1/8; B, seedling × 1/3; C, perforate leaf × 2/3; D, eperforate leaf × 2/3; E, base of plant showing erect rhizome and leaf–axillary tubercles × 2/3; F, inflorescence × 2/3; G, inflorescence, nearside half of spathe removed × 1; H, detail of spadix × 5; J, flower, longitudinal section × 8; K, infructescence × 2/3; L, seed, side view × 3. Dracontioides desciscens: A, Harley et al. 18009 (Kew slide collection); B, Cult. Kew 1977–640 (Kew spirit collection 39121); C, Harley et al. 18009 (K); D, Lewis & de Carvalho 1060 (K); E, Harley et al. 18241A (Kew spirit collection 47695) & Lewis & de Carvalho 1060 (K); F–L, Harley et al. 18009 (Kew spirit collection 46561, 46564 & 47699).

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23. Anaphyllopsis Anaphyllopsis A. Hay in Aroideana 11 (1): 25–31 (1989, “1988”). TYPE: A. americana (Engler) A. Hay

22. Dracontioides

long, smooth to roughened-verruculate, often transversely variegated, geniculate at apex, sheath less than half petiole length. BLADE: deeply sagittate to subtripartite, often with a few perforations of irregular size between primary lateral veins; basal ribs very well-developed, primary lateral veins mostly arising near petiole insertion, very long-arcuate towards apex of each division, running into margin, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: shorter than petiole. SPATHE: marcescent, tube with convolute margins, longitudinally white-striped, blade strongly fornicate, usually obscuring mouth of tube, brown-purple. SPADIX: sessile to shortly stipitate, cylindric, obtuse, shorter than spathe tube, flowering sequence basipetal. FLOWERS: bisexual, perigoniate; tepals 4, fornicate, subtruncate. STAMENS: 4, free, strongly exserted from flower at anthesis, anthers longer than connective, connective slender, thecae ovate-ellipsoid, dehiscing by pore-like apical slit. POLLEN: monosulcate, ellipsoid-oblong, small (mean 23 µm.), exine subreticulate. GYNOECIUM: ovate-conoid, ovary 2-locular, ovules 1 per locule, anatropous, placenta axile, stylar region attenuate, longer than tepals, stigma small, button-like. BERRY: obovoid, somewhat furrowed, 1–2-seeded, dark purplish-red. SEED: reniform, attenuate towards micropyle, testa strongly dentate-cristate, thick, hard, brown, embryo curved, endosperm present. See Plates 22, 111A. CHROMOSOMES: 2n = 26. DISTRIBUTION: 1 sp.; Brazil (Atlantic region); the record from Tijuca, Rio de Janeiro (Peyritsch 1879) is almost certainly a mistake. ECOLOGY: tropical humid forest; helophytes in swamp forest, marshes, stream margins, open or shaded sites, in peat or sand. ETYMOLOGY: Dracontium and Greek suffix -oides (resemblance), i.e. similar to Dracontium. TAXONOMIC ACCOUNTS: Mayo (1978), Hay (1992a).

HABIT: seasonally dormant (A. americana), solitary herbs, rhizome hypogeal, erect. LEAVES: solitary, rarely 2. PETIOLE: unarmed, smooth to tubercular, geniculate apically. BLADE: pinnatifid and perforate or pinnatisect, juvenile leaves entire; basal ribs well-developed, primary lateral veins of ultimate lobes pinnate, higher order venation reticulate. INFLORESCENCE: solitary, rarely 2. PEDUNCLE: long, similar to petiole in colour and texture. SPATHE: membranaceous, convolute basally, spirally twisted apically, marcescent, papery when dry. SPADIX: stipitate, stipe mostly adnate to spathe, flowering sequence basipetal. FLOWERS: bisexual, perigoniate; tepals 4, fornicate. STAMENS: 4, free, filaments short, linear, connective slender, thecae dehiscing by short, apical pore-like slit. POLLEN: monosulcate, ellipsoid, medium-sized (31 µm.), exine foveolate, apertural exine psilate. GYNOECIUM: ovary 1–locular, ovules 1–2, anatropous, placenta basal, stylar region ± attenuate, stigma subcapitate. BERRY: ovoid to obpyramidal, reddish. SEED: campylotropous, testa thick, verruculose to channelled, embryo curved, endosperm present. See Plates 23, 111B. CHROMOSOMES: 2n = 26. DISTRIBUTION: 3 spp., tropical South America:– Brazil (Amazonia), French Guiana, Surinam, Venezuela. ECOLOGY: tropical humid swamp forest; helophytes in sandy, partially flooded places along streams, swamp forest. ETYMOLOGY: Anaphyllum and Greek opsis (appearance); i.e. “like Anaphyllum”. TAXONOMIC ACCOUNTS: Hay (1989, 1992a).

23. Anaphyllopsis

LASIOIDEAE : ANAPHYLLOPSIS

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B F

E D A

C

K

H

L

J

G

Plate 23. Anaphyllopsis. A, leaf × 1/4; B, gynoecium, longitudinal section × 5; C, leaf × 1/4; D, infructescence × 2/3; E, immature fruit, longitudinal section × 6; F, seed, side view × 4; G, leaf × 1/4; H, base of plant showing erect rhizome × 1/2; J, inflorescence, nearside half of spathe removed × 2/3; K, detail of spadix × 4; L, flower, nearside tepal removed × 6. Anaphyllopsis pinnata: A–B, Wessels Boer 2387 (U); A. cururuana: C–F, Anderson 10627 (NY); A. americana: G, J, Leprieur 152 (P); H, Aroideana 11(1) 30, fig. 3 (1988) (as A. cururuana); K–L, Mélinon 52 (P).

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C

24. Pycnospatha

C

25. Anaphyllum

Pycnospatha Thorel ex Gagnepain in Bull. Soc. Bot. France 88: 511 (1941). TYPE: P. palmata Thorel ex Gagnepain

Anaphyllum Schott in Bonplandia 5: 126 (1857). TYPE: A. wightii Schott

HABIT: seasonally dormant herbs with subglobose tuber. LEAVES: 1–2. PETIOLE: smooth, rough or aculeate, mottled, sheath very short, inconspicuous. BLADE: dracontioid, i.e. trisect, anterior division trifid, segments simple to pinnatifid, posterior divisions bifid to pedatifid, segments then simple to pinnatifid, ultimate lobes decurrent, ovate-elliptic to triangular, acute to acuminate; primary lateral veins pinnate, running into distinct marginal vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing before or with leaf. PEDUNCLE: much shorter than petiole, similar in appearance and texture to petiole. SPATHE: margins not overlapping, strongly fornicate, thick, ± purple, marcescent. SPADIX: much shorter than spathe, conic to ovoid-conoid, stipitate, fertile to apex, flowering sequence basipetal. FLOWERS: bisexual, perigone absent. STAMENS: ca. 6 or more per flower, free, crowded densely together with those of neighbouring flowers, filaments oblong, flattened, connective slender, thecae oblong-ellipsoid, dehiscing by apical, pore-like slit. POLLEN: monosulcate, ellipsoid to oblong, medium-sized (mean 34 µm.), exine subreticulate to rugulate. GYNOECIUM: elongated-flask-shaped, ovary 1-locular, ovule 1, anatropous to hemianatropous, funicle short, placenta basal to subparietal, stylar region greatly elongated, straight or somewhat curved, projecting well beyond stamens, stigma small, scarcely or no broader than style. BERRY: globose, pericarp densely covered with conic prickles, with conspicuous, persistent style remnant, dark green. SEED: reniform, black or very dark brown, testa hard, thick, verrucose, containing white druses, embryo large, ellipsoid to slightly curved, endosperm very sparse, only a few cell layers thick. See Plates 24, 111C. CHROMOSOMES: 2n = 26. DISTRIBUTION: 2 spp.; tropical southeast Asia:– Laos, Thailand, Vietnam. ECOLOGY: tropical humid forest; geophytes on forest floor in sandy loam. ETYMOLOGY: Greek pyknos (compact, thick) and spathê (spathe). TAXONOMIC ACCOUNTS: Bogner (1973c), Hay (1992a), Boyce (1993b).

HABIT: evergreen herbs, clump- or colony-forming, rhizome creeping. LEAVES: solitary to few. PETIOLE: smooth to tuberculate, geniculate apically, sheath short. BLADE: sagittate-hastate to pedatifid when juvenile, trisect at maturity, anterior division remotely pinnatisect with ± oblong-lanceolate, acute lobes, the upper ones decurrent, posterior divisions either ± oblong-lanceolate or deeply divided into 3 coherent segments, rachis geniculate at insertion of anterior and posterior divisions; basal ribs well-developed, primary lateral veins of ultimate lobes pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: very long and slender, similar in colour and texture to petiole. SPATHE: membranous to coriaceous, marcescent, either convolute basally and becoming spirally twisted and long-acuminate apically, or oblong-ovate, ± flat and fully expanded. SPADIX: cylindric, much shorter than spathe, stipitate or sessile, flowering sequence basipetal. FLOWERS: bisexual, perigoniate, tepals 3–4, fornicate. STAMENS: 3–5, free, filaments fairly wide, connective slender, thecae ellipsoid, dehiscing by short, apical, pore-like slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 29 µm., range 25–33 µm.), exine foveolate to reticulate, sometimes with elevated psilate regions, apertural exine psilate. GYNOECIUM: ovary ovoid, 1-locular, ovule 1, hemianatropous, funicle short, placenta parietal on single intrusive septum, stylar region thick, attenuate, stigma subcapitate, exuding droplet at anthesis. BERRY: ovoid, smooth, red. SEED: ovoid, funicle slender, testa membranaceous, smooth, embryo stout, straight, endosperm absent. See Plates 25, 111D. CHROMOSOMES: 2n = 26. DISTRIBUTION: 2 spp.; southern India (Kerala, Tamil Nadu). ECOLOGY: tropical evergreen forest in leaf litter and in swamp forest undergrowth; geophytes, rare. ETYMOLOGY: Greek ana- (up, erect) and phyllon (leaf). TAXONOMIC ACCOUNTS: Engler (1911), Sivadasan (1982), Hay (1992a).

24. Pycnospatha

25. Anaphyllum

LASIOIDEAE : ANAPHYLLUM

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A

C B

Q

P D F

H G

L

N

M

E

J K

Plate 24. Pycnospatha. A–E, leaf developmental sequence from first leaf (A) to flowering size (E) × 1/3; F, detail of petiole × 1; G, habit × 1/8; H, inflorescence × 2/3; J, spathe sectioned to show spadix × 1; K, detail of spadix × 3; L, flower × 6; M, gynoecium, longitudinal section × 6; N, gynoecium, transverse section × 6; P, infructescence × 2/3; Q, seed, side view × 5. Pycnospatha arietina: A–F, Bogner 395 (K); G, Cult. Kew 1971–1039 (Kew slide collection 8021); H–J, Bogner 395 (Kew spirit collection 34429); K–Q, Bogner 395 (Kew spirit collection 29047.184).

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H

A

G

J

D

B

E

C

F

Plate 25. Anaphyllum. A, habit × 1/8; B, leaf × 1/3; C, base of plant × 2/3; D, inflorescence × 2/3; E, detail of spadix × 5; F, flower, longitudinal section × 6; G, seed, hilum view × 3; H, leaf × 1/3; J, inflorescence × 2/3. Anaphyllum wightii: A–B, Cult. Kew 1984–04519; Fliegner 245 (Kew spirit collection 49762 & 51853); C, Sivadasan CU 9060 (K); D–F, Cult. Kew 1983–02560; Hay s.n. (Kew spirit collection 49796); G, Cult. Kew 1984–04519; Fliegner 245 (Kew spirit collection 51853); A. beddomei: H, Anon. s.n. (K); J, Blasco s.n. (K).

LASIOIDEAE : ANAPHYLLUM

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C

26. Cyrtosperma Cyrtosperma Griffith, Notul. Pl. Asiat. (Posthum. Pap.) 3: 149 (1851); Icon. pl. Asiat. 3, t.169 (1851). TYPE: C. lasioides Griffith (“lacioides”), [= C. merkusii (Hassk.) Schott]. SYNONYM: Arisacontis Schott in Bonplandia 5: 129 (1857). HABIT: slender to gigantic evergreen herbs, usually solitary, sometimes clump-forming, rhizome thick, condensed, creeping. LEAVES: several. PETIOLE: sometimes very long, aculeate, geniculate apically, sheath short. BLADE: deeply sagittate, hastate-sagittate or ± tripartite (posterior divisions usually larger than anterior), veins sometimes aculeate on lower surface; basal ribs very well-developed, primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing with the leaves. PEDUNCLE: long, similar to petioles. SPATHE: marcescent, erect, blackish purple to white, convolute or not in lower part, upper part rarely somewhat fornicate, long-acuminate and twisted in some species. SPADIX: sessile or stipitate. FLOWERS: bisexual, perigoniate; tepals 4–6, somewhat thickened at apex, fornicate. STAMENS: 4–6, free, filaments free, flat and broad, connective slender, thecae oblong-ovate, dehiscing by apical slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 29 µm., range 28–30 µm.), exine foveolate, apertural exine psilate. GYNOECIUM: 1-locular, ovules 1–many, campylotropous to subamphitropous, placenta basal to parietal, stylar region short or inconspicuous, stigma subhemispheric, exuding droplet at anthesis. BERRY: obovoid, 1–7-seeded, usually red when mature, bearing remnant of stigma (in C. cuspidispathum ripe berries are extruded and dangle on strips of tepal epidermis; A. Hay pers. comm.). SEED: reniform to orbicular to helically twisted, cristate, warty or smooth, embryo curved, endosperm present. See Plates 26, 112A. CHROMOSOMES: 2n = 26. DISTRIBUTION: 11–12 spp.; tropical southeast Asia, Malay Archipelago, Melanesia, Pacific:– Brunei, Caroline Is., China (cultivated?), Cook Is., Fiji, Gilbert & Ellice Is., Guam, Indonesia

26. Cyrtosperma

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THE GENERA OF ARACEAE

(Borneo, Irian Jaya, Java, Palau Is., Sumatra), Malaysia (Borneo, Peninsula), Mariana Is., Marquesas Is., Marshall Is., Papua New Guinea, Philippines, Samoa, Singapore, Society Is., Solomon Is., Tahiti, Vanuatu, Vietnam (cultivated?). ECOLOGY: tropical humid forest, swamp forest, open swamps and cultivated areas; helophytes in streams, ponds and other wet places. NOTES: Hay (1988) recognized 4 informal groups:– “Merkusii” group, “Cuspidispathum” group, “Carrii” group, “Macrotum” group. ETYMOLOGY: Greek kyrtos (curved) and sperma (seed); the seeds are strongly curved. TAXONOMIC ACCOUNTS: Engler (1911), Thompson (1982), Hay (1988, 1992a).

27. Lasimorpha Lasimorpha Schott in Bonplandia 5: 127 (1857). TYPE: L. senegalensis Schott (syn. Cyrtosperma senegalense (Schott) Engler). SYNONYM: [Lasiomorpha Engler in Engler, Pflanzenreich 48 (IV.23C): 14 (1911), orth. var.]. HABIT: robust to gigantic evergreen herbs forming large colonies, rhizome short, thick, hypogeal, vigorously stoloniferous. LEAVES: several. PETIOLE: very long, 4–6 angled, aculeate along angled ridges, weakly geniculate at apex, sheath short. BLADE: deeply sagittate, sometimes weakly hastate, to over 1m long, coriaceous, ± erect; basal ribs well-developed, primary lateral veins pinnate, long arcuate towards apex of each leaf division and running into marginal vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: subequal to petiole and similar in appearance. SPATHE: erect, ovate, ± convolute in lower third, gaping at anthesis, striped purple and yellow within, persistent. SPADIX: sessile to stipitate, cylindric, obtuse, shorter than spathe, flowering sequence basipetal. FLOWERS: bisexual, perigo-

27. Lasimorpha

C

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G

B

A

K

N H

D

M

J

E

C

F

L

Plate 26. Cyrtosperma. A, inflorescence × 2.3; B, flower, longitudinal section × 8; C, infructescence × 2/3; D, seed, side view × 6; E, leaf × 2/3; F, base of plant × 2/3; G, habit × 1/8; H, leaf × 2/3; J, base of petiole × 2/3; K, inflorescence × 2/3; L, detail of spadix × 4; M, flower, longitudinal section × 8; N, infructescence × 2/3. Cyrtosperma cuspidispathum: A–B, NGF 10241 (K & Kew spirit collection 18946); C–D, Hay s.n. (Kew spirit collection 45943); C. beccarianum: E, Reksodihardjo 412 (K); F, van Royen 4792 (K); C. macrotum: G, Aroids of Papua New Guinea pl. x, 3 (1990); C. merkusii: H, Lobb s.n. (K); J, Nicolson 1217 (K); L–M, Bogner 1510 (Kew spirit collection 45187); N, Reksodihardjo 311 (K).

L A S I O I D E A E : C Y RTO S P E R M A

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H

E

F

D

G

B

C

J

A

Plate 27. Lasimorpha. A, base of plant showing stolons × 2/3; B, leaf × 2/3; C, inflorescence × 2/3; D, detail of spadix × 4; E, flower, tepals removed × 8; F, gynoecium, longitudinal section × 8; G, stamens, top view × 8; H, infructescence × 2/3; J, seed, side view × 5. Lasimorpha senegalensis: A, Bogner 691 (K & Kew spirit collection 29047.472); B, Leeuwenberg 1873 (K); C, Bogner 691 (Kew spirit collection 57526); D–J, Meikle 642 (Kew spirit collection 25024).

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niate; tepals 4(–5), free, fornicate. STAMENS: 4(–5), filaments free or partially to completely connate, anthers partially exserted from tepals at anthesis, connective slender, thecae dehiscing by short apical slit. POLLEN: monosulcate, ellipsoid, small (22 µm.), exine subreticulate, apertural exine psilate. GYNOECIUM: ovoid-ellipsoid to oblong, ovary 1-locular, ovules 4–6(–8), campylotropous, placenta single, ± prominent, parietal and basal, stylar region short, somewhat narrowed, stigma discoid-hemispheric. BERRY: irregularly globose, 1–4-seeded, red. SEED: strongly curved, weakly strophiolate, testa hard, brown, warty to spiny, embryo rather large, curved, endosperm present but only as a few cell layers. See Plates 27, 112B. CHROMOSOMES: 2n = 26. DISTRIBUTION: 1 sp.; tropical west and central Africa:– Angola, Benin, Cameroon, Central African Republic, Chad, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Nigeria, Senegal, Sierra Leone, Zaïre. ECOLOGY: tropical swamp forest, open wet areas; helophyte along streams, in ditches and ponds, in forest gaps, often very abundant. ETYMOLOGY: Lasia and Greek morphê (shape). TAXONOMIC ACCOUNTS: Engler (1911, as Cyrtosperma), Knecht (1983), Hay (1988, 1992a).

C

veins mostly arising near petiole insertion, long-arcuate towards division apex and running into margin, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole and similar in appearance. SPATHE: ovatelanceolate, fully expanded, persistent, red brown. SPADIX: shorter or equalling spathe, cylindric, stipitate, stipe basally adnate to spathe, flowering sequence basipetal. FLOWERS: bisexual, perigoniate; tepals 4–6, fornicate. STAMENS: 4–6, free, filaments oblong, flattened, connective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid-oblong, small (mean 18 µm.), exine scabrate or verruculate, minutely foveolate between verruculae. GYNOECIUM: cylindric to obovoid, ovary 1-locular, ovule 1, anatropous, placenta parietal to subbasal, stigma discoidhemispheric. BERRY: ovoid to ellipsoid, smooth, strongly exserted at maturity, very prominent and thicker than spadix, red. SEED: curved, ± spherical, testa thin, hard, smooth, dark brown, embryo curved, endosperm present but only as a few cell layers. See Plates 28, 112C. CHROMOSOMES: 2n = 26. DISTRIBUTION: 1 sp.; Malay Archipelago:– Indonesia (Sumatra), Malaysia (Sarawak, Peninsula). ECOLOGY: tropical humid forest, usually peat swamp forest; helophyte, along streams. ETYMOLOGY: Greek pous, podos (foot) and Lasia; refers to the ± basal placentation of the ovule and the resemblance to Lasia. TAXONOMIC ACCOUNTS: Engler (1911), Hay (1988, 1992a).

28. Podolasia Podolasia N.E. Brown Gard. Chron., ser.2, 18: 70 (1882). TYPE: P. stipitata N.E. Brown HABIT: evergreen herbs, solitary or forming small clumps, rhizomatous, aerial stem erect to decumbent, internodes distinct, unarmed. LEAVES: several. PETIOLE: long, geniculate apically, aculeate, spines either patent or pointing downwards, sheath short. BLADE: unarmed, sagittate to hastate or almost tripartite, coriaceous, posterior divisions ± equalling anterior, sometimes longer; basal ribs well-developed, primary lateral

28. Podolasia

C

29. Lasia Lasia Loureiro, Fl. Cochinch. 64, 81 (1790). TYPE: L. aculeata Lour. (= L. spinosa (L.) Thwaites). SYNONYM: [Lasius Hasskarl, Cat. Bogor. 59 (1844), orth.var.] HABIT: clump- and colony-forming evergreen herbs, stoloniferous, stem thick, aculeate or unarmed (L. concinna), erect to decumbent, green, epigeal or submersed, internodes

29. Lasia

LASIOIDEAE : LASIA

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C

D

E A

G

F

B

Plate 28. Podolasia. A, habit × 2/3; B, detail of spadix × 5; C, flower × 10; D, flower, nearside tepals removed × 10; E, gynoecium, longitudinal section × 10; F, infructescence, × 2/3; G, seed, side view × 3. Podolasia stipitata: A, Corner 30898 (K); Kunstler (Dr King’s collector) 5499 (K); Parris 10979 (K); B, Corner 30898 (K); C–G, Parris 10979 (K).

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K

J

F

B

H

A

D

G

E

C

Plate 29. Lasia. A, leaf × 2/3; B, leaf × 2/3; C, leaf and base of plant with adventitious shoots, large portion of petiole removed × 2/3; D, habit × 1/15; E, inflorescence × 2/3; F, tranverse section of spiral portion of spathe × 2/3; G, detail of spadix × 5; H, flower, longitudinal section × 7; J, infructescence × 2/3; K, seed, side view × 5. Lasia spinosa: A, Cult. Kew (K); B, Hooker & Thomson s.n. (K); C, Clarke 7934 (K); Bogner 428 (Kew spirit collection 46984); D, Forman & Blewett 975 (Kew slide collection); E–H, Mayo 133 (Kew spirit collection 47735); J–K, Hooker & Thomson s.n. (K); Boyce 340 (Kew slide collection).

LASIOIDEAE : LASIA

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relatively long or short. LEAVES: several. PETIOLE: long, aculeate, weakly geniculate apically, sheath relatively short. BLADE: sagittate to hastate-sagittate when juvenile, adult blade deeply pinnatifid in anterior division, posterior divisions pedatifid, sometimes simple, adult blade rarely entire, or bipinnatifid (L. concinna), major veins aculeate on lower surface; primary lateral veins pinnate in anterior division, pedate in posterior divisions, primary lateral veins of each pinna forming submarginal collective vein in pinnatifid leaves, or running into marginal vein in simple leaves, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole, aculeate as petiole. SPATHE: linear, very long and narrow (L. spinosa) or broader (L. concinna), very thick and spongy, spirally twisted, marcescent or deciduous, basal part enclosing spadix, gaping at anthesis. SPADIX: shortly cylindric, obtuse, sessile. FLOWERS: bisexual, perigoniate; tepals usually 4, more rarely 6, fornicate. STAMENS: 4(–6), free, filaments broad, connective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 27 µm.), exine reticulate, psilate along aperture margins. GYNOECIUM: ovoid to ellipsoid, ovary 1-locular, ovule 1, anatropous, funicle very short, placenta apical, stylar region well developed, shortly attenuate to cylindric, stigma discoid-hemispheric. BERRY: borne in cylindric infructescence, crowded, quadrangular, apically densely muricate to spinose or smooth, 1-seeded, green. SEED: large, compressed-obovoid, testa thin, brown, hard, somewhat rugose, embryo large, somewhat curved, endosperm present but only as a single cell layer. See Plates 29, 112D. CHROMOSOMES: 2n = 26. DISTRIBUTION: 2 spp.; tropical southeast Asia, Malay Archipelago:- Bangladesh, Brunei, Burma, Cambodia, China (Guandong, Guangxi, Hainan, Hong Kong, Taiwan, Yunnan), India, Indonesia, Laos, Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, Singapore, Sri Lanka, Thailand, Vietnam. ECOLOGY: tropical humid forest; helophytes, wet places in forest, open swamps, along streams, rice fields, tidal flats. ETYMOLOGY: Greek lasios (shaggy), refers to the densely spiny stems. TAXONOMIC ACCOUNTS: Engler (1911), Sivadasan (1982), Hay (1988, 1990b, 1992a).

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30. Urospatha Urospatha Schott, Aroideae 3 (1853). LECTOTYPE: U. sagittifolia (Rudge) Schott (“sagittaefolia ”, Pothos sagittaefolia Rudge; see Nicolson 1967). SYNONYMS: Urophyllum K. Koch in Berliner Allg. Gartenzeitung 25: 173 (1857, non Jack ex Wallich 1824); Urospathella Bunting in Phytologia 65: 391 (1988). HABIT: robust to slender evergreen herbs, solitary to clumpforming, rhizome subterranean, horizontal or vertical, spongy. LEAVES: few, basal, ± erect. PETIOLE: long, spongy, smooth or scabrid-verrucose, sometimes angled, often variegated, apical geniculum absent or only very weakly developed, sheath long to short. BLADE: deeply sagittate or subtripartite to hastate, posterior divisions usually longer than anterior division, rarely blade lanceolate-linear and lacking posterior divisions (U. wurdackii); basal ribs welldeveloped, primary lateral veins of both anterior and posterior divisions pinnate, long-arcuate towards division

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30. Urospatha

apex and running into marginal vein, higher order venation reticulate. INFLORESCENCE: solitary, rarely 2 in each floral sympodium. PEDUNCLE: equal to or longer than leaf, similar in appearance to petiole. SPATHE: erect, persistent, convolute below, gaping above, apically long-acuminate, usually spirally twisted, sometimes flattened. SPADIX: shortly stipitate or sessile, usually much shorter than spathe, cylindric to subcylindric, obtuse. FLOWERS: bisexual, perigoniate; tepals 4–6, fornicate. STAMENS: 4–6, free, filaments broadish, a little compressed, connective slender, thecae ellipsoid, dehiscing by apical slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 26 µm., range 25–28 µm.), exine subreticulate, subrugulate to reticulate, apertural exine psilate. GYNOECIUM: ovoid, ovary 1–2-locular, ovules (1–)2–4 or more per locule, anatropous, placenta axile in 2-locular ovaries, basal in 1-locular ovaries, stylar region a little narrower than ovary, stigma broad, circular, discoid. BERRY: obovoid, 1–5(–8)-seeded, green to greenish yellow. SEED: curved, testa hard, brown, thickish, strongly warty, or spiny or crested, embryo curved, endosperm present but only as a very thin layer about 2 cells thick. See Plates 30, 113A. CHROMOSOMES: 2n = 52. DISTRIBUTION: ca. 10 spp. (T. Croat pers. comm.); tropical America:– Brazil, Colombia, Costa Rica, ?Ecuador, French Guiana, Guatemala, Guyana, Nicaragua, Panama, Peru, Surinam, Venezuela. ECOLOGY: tropical humid forest and wetlands; helophytes, open aquatic habitats, swamps, along streams, brackish water. NOTE: Urospatha wurdackii (syn. Urospathella wurdackii) is notable for its small stature, linear-lanceolate leaves, and very slender spathe. ETYMOLOGY: Greek “oura” (tail) and “spathê” (spathe). TAXONOMIC ACCOUNTS: Engler (1911), Bunting (1988, 1989a), Hay (1992a).

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N

G

M

L

K

F B

A

D

J

C

H E

Plate 30. Urospatha. A, habit × 1/10; B, leaf × 2/3; C, inflorescence × 2/3; D, detail of spadix × 4; E, flower, longitudinal section × 10; F, infructescence showing persistent spathe × 2/3; G, seed, side view × 5; H, leaf × 2/3; J, leaf × 2/3; K, inflorescence × 2/3; L, diagrammatic longitudinal section of inflorescence to show partial spathe/spadix fusion × 2/3; M, detail of spadix × 4; N, flower, longitudinal section × 10. Urospatha sagittifolia : A, Mayo (Kew slide collection); U. sagittifolia: B, Jenman 5777 (K); C, Bogner s.n. (Kew spirit collection 29047.469); D–G, Bogner 580 (Kew spirit collection 29047.191); U. angustiloba : H, Spruce 3761 (K); U. wurdackii: J, Huber, Tillet & Davidse 3721 (K); K–N, Davidse, Huber & Tillet 17085 (K & Kew spirit collection 57275).

LASIOIDEAE : UROSPATHA

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VI. Subfamily Calloideae

VII. Subfamily Aroideae

Subfamily Calloideae Endlicher, Gen. Pl. 239 (1837, “Callaceae”).

Subfamily Aroideae

Laticifers present, simple, articulated; trichosclereids absent; rooted aquatic, stem rhizomatous; leaves distichous; petiole not geniculate apically, sheath long; blade cordate, venation parallel-pinnate, uniformly fine; spathe fully expanded, elliptic- to ovate-lanceolate, persistent; flowers bisexual, perigone absent, 3-merous; stamens 6 (or more), thecae dehiscing by longitudinal slit, pollen dicolpate; ovary 1-locular, ovules 6–9, anatropous, placenta basal; endosperm copious.

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31. Calla Calla L., Sp. Pl. 968 (1753). TYPE: C. palustris L. SYNONYMS: Provenzalia Adanson, Fam. 2: 469 (1763); Aroides Heister ex Fabricius, Enum., ed. 2, p. 42 (1763); Callaria Rafinesque in Amer. Monthly Mag. & Crit. Rev. 2: 267 (1818). Laticifers present, simple, articulated. HABIT: seasonally dormant herb with repent or submersed, green, rhizomatous stem, rooting at nodes. LEAVES: distichous. PETIOLE: sheath long, with long, free, ligulate apex. BLADE: cordate to broadly cordate, rounded, cuspidate-apiculate; primary lateral veins not differentiated, higher order venation parallel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: erect, as long or longer than petiole. SPATHE: fully expanded, elliptic- or ovate-lanceolate, acuminate, shortly decurrent, persistent, white within at anthesis, green without. SPADIX: stipitate, cylindric, obtuse. FLOWERS: bisexual, often male at spadix apex, perigone absent. STAMENS: ca. 6, free, sometimes more, filaments somewhat flattened, anthers short, connective slender, short, thecae ellipsoid, opposite, dehiscing by longitudinal slit. POLLEN: diaperturate, globose, small (mean 23 µm.), exine foveolate, apertural exine verrucate. GYNOECIUM: shortly ovoid, 1-locular, ovules 6–9, anatropous, oblong, funicles short, placenta basal, stylar region attenuate, stigma small, subhemispheric. BERRY: spheroid-conic, several-seeded, red. SEED: tereteoblong, testa thick, scrobiculate towards chalaza, sulcate-striate towards micropyle, raphe prominent, embryo axile, elongate, endosperm copious. See Plates 31, 113B. CHROMOSOMES: 2n = 36, 54, 72. DISTRIBUTION: 1 sp.; circumboreal:– Austria, Belgium, Canada, China, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, ?Hungary, ?Japan, Korea (N.), Netherlands, Norway, Poland, Romania, Russia, Slovak Republic, Slovenia, Sweden, Switzerland, Turkey, USA. ECOLOGY: in forest swamps, often between forest margins and raised bogs, often with Sphagnum, up to 1270m alt.; creeping helophyte in streams and ponds. ETYMOLOGY: classical name, first used by Pliny for two different kinds of aroids; perhaps from the Greek kallos (beauty). TAXONOMIC ACCOUNTS: Krause (1908), Dudley (1937), Huttleston (1953), Riedl (1977–1979), Topic & Ilijanic (1989).

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Laticifers present (except Pistieae, Stylochaetoneae, Zamioculcadeae), usually simple and articulated, more rarely anastomosing (in Caladieae, Colocasieae, Zomicarpeae); trichosclereids absent; stem most frequently hypogeal, tuberous or rhizomatous, less often aerial, rarely hemiepiphytic climbers or epiphytes (Culcasieae, Philodendreae, Syngonium), very rarely floating aquatics (Pistieae); petiole usually not geniculate apically (except Anubiadeae, Bognera, Culcasieae, Zamioculcadeae, rarely in Philodendreae and Homalomeneae); higher order leaf venation more usually reticulate, less often parallel-pinnate (in Aglaonemateae, Anubiadeae, Colocasieae, Dieffenbachia, Homalomeneae, Peltandreae, Philodendreae, Schismatoglottideae, Zantedeschieae); spathe usually differentiated into lower, convolute tube and upper, gaping blade; flowers unisexual, perigone absent (except Stylochaetoneae, Zamioculcadeae); pollen inaperturate (except Zamioculcadeae and sometimes Stylochaetoneae).

Tribe Zamioculcadeae

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Tribe Zamioculcadeae Engler in Nova Acta Acad. Leopold.Carol. 39: 141 (1876, “Zamioculcaseae ”). Laticifers absent; seasonally dormant or evergreen (Zamioculcas), stem hypogeal; petiole geniculate; leaf blade compound; spathe convolute basally, blade reflexed at anthesis; spadix subequalling spathe, female zone separated from male by short, constricted zone of sterile flowers, male zone cylindric to clavate, fertile to apex; flowers unisexual, perigoniate; tepals 4, free, ± prismatic; stamens 4, surrounding ± clavate pistillode, filaments free, (connate in Gonatopus), anther terminal, connective inconspicuous, pollen extended-monosulcate to fully zonate, extruded in strands; ovary 2-locular, ovules 1 per locule, hemianatropous, placenta axile to basal, style short, distinct, stigma large, hemispheric-discoid; berries and seeds large, testa smooth, endosperm absent.

32. Zamioculcas Zamioculcas Schott, Syn. Aroid. 71 (1856). TYPE: Z. loddigesii Schott, nom. illeg. (Caladium zamiaefolium Loddiges, Z. zamiifolia (Loddiges) Engler). HABIT: seasonally dormant or evergreen herb with short, very thick rhizome. LEAVES: few to many, erect, leaflets deciduous during dormancy leaving persistent petiole. PETIOLE: terete, base greatly thickened and succulent, geniculate at apex, sheath ligulate, free almost to the base, very short, inconspicuous. BLADE: pinnatisect, leaflets oblong-elliptic, thickly coriaceous, capable of rooting at base once shed and forming new plants; primary lateral veins of each leaflet pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, held at ground level. PEDUNCLE: very short. SPATHE: entirely persistent to fruiting stage, slightly constricted between tube and blade, green without, whitish within, tube convolute, blade longer than tube, expanded and horizontally reflexed

C

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B

F C

D

A

E

Plate 31. Calla. A, habit × 2/3; B, spadix × 4; C, flower × 15; D, gynoecium, longitudinal section × 15; E, infructescence × 2/3; F, seed, side view × 10. Calla palustris: A, Bogner s.n. (Kew spirit collection 58031, 58904); B–F, Bogner 2117 (Kew spirit collection 57571).

31. Calla

CALLOIDEAE : CALLA

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C

B

D

A

E

J

H

K

L

G

Q

P

N

F

M Plate 32. Zamioculcas. A, habit × 1/3; B, leaflet × 2/3; C, leaflet with adventitious tuberlet × 2/3; D, first leafy shoot from adventitious tuberlet × 2/3; E, base of leaf showing swollen petiole × 2/3; F, inflorescence × 1; G, spadix × 1; H, male flower, nearside tepal removed × 8; J, detail of male flowers × 5; K, sterile flower, nearside tepal removed × 8; L, detail of sterile flowers × 5; M, gynoecium, longitudinal section × 8; N, detail of female flowers × 5; P, infructescence, longitudinal section × 1; Q, seed, side view × 6. Zamioculcas zamiifolia: A–M, Bogner s.n. Cult. Kew 1967–49401 (Kew spirit collection 49776, 29047.131 & 58712); P–Q, Faden & Faden 77/377 (K).

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SYNONYMS: Heterolobium A. Peter, Nachr. Ges. Wiss. Göttingen, Math.-Phys. Kl. 1929 (3): 211, 221 (1930); Microculcas A. Peter, Nachr. Ges. Wiss. Göttingen, Math.Phys. Kl. 1929 (3): 212, 222 (1930).

32. Zamioculcas

at anthesis. SPADIX: sessile, female zone subcylindric, separated from male zone by short constricted zone bearing sterile flowers, male zone cylindric, ellipsoid to clavate, fertile to apex. FLOWERS: unisexual, perigoniate; tepals 4, in two whorls, decussate. MALE FLOWER: tepals subprismatic, apex thickened, stamens 4, free, shorter than tepals, filaments free, oblong, thick, somewhat flattened, anthers introrse, connective slender, thecae ovate-ellipsoid, dehiscing by apical slit, pistillode clavate, equalling tepals. POLLEN: extruded in strands, extended monosulcate to perhaps fully zonate, ellipsoid, large (mean 60 µm.), exine thick, fossulate-foveolate, apertural exine verrucate. STERILE FLOWERS: each consisting of 4 tepals surrounding a clavate pistillode. FEMALE FLOWER: tepals strongly thickened apically, staminodes lacking, gynoecium equalling tepals, ovary ovoid, 2-locular, ovules 1 per locule, hemianatropous, funicle very short, placenta axile near base of septum, stylar region attenuate, stigma large, discoid-capitate. BERRY: depressed-globose with furrow at septum, 1–2-seeded, surrounded by persistent tepals, white, infructescence ellipsoid. SEED: ellipsoid, testa smooth, brown, raphe conspicuous, embryo large, rich in starch, endosperm nearly absent, present only as a few cell layers at chalazal end. See Plates 32, 113C. CHROMOSOMES: 2n = 34. DISTRIBUTION: 1 sp.; tropical east and subtropical southeast Africa:– Kenya, Malawi, Mozambique, South Africa (Natal), Tanzania (incl. Pemba, Zanzibar), Zimbabwe. ECOLOGY: tropical moist forest, savannas; geophytes on forest floor or in stony ground. ETYMOLOGY: Zamia (genus of cycads) and Greek kolokasia (from Middle Eastern “qolqas”); perhaps in reference to the pinnately compound leaves, in fanciful comparison to Zamia. TAXONOMIC ACCOUNTS: Engler (1905), Obermeyer & Strey (1969), Mayo (1985a).

C

HABIT: Seasonally dormant herbs, stem subterranean, a subglobose tuber or a cylindric, horizontal rhizome. LEAF: solitary, rarely pilose or scabrous, preceded by lanceolate cataphylls. PETIOLE: geniculate basally or centrally. BLADE: usually trisect, rarely not (G. petiolulatus), primary divisions trifid to trisect or pinnatifid, or pinnatisect to quadri-pinnatifid, pinnae geniculate at junction with rachis, ultimate lobes varying from linear to broad-elliptic, often decurrent; primary lateral veins of each lobe pinnate, forming arching submarginal collective vein, higher order venation reticulate. INFLORESCENCE: 1–4 in each floral sympodium, appearing before or with leaves, subtended by several cataphylls. PEDUNCLE: erect, very short to long. SPATHE: constricted between tube and blade, tube convolute, subglobose, cylindric or suburceolate, blade oblong to elliptic, reflexed at anthesis, marcescent. SPADIX: subequal to spathe, female zone subcylindric, separated from male zone by very short, constricted zone of sterile flowers, male zone longer than female, cylindric to clavate, fertile to apex. FLOWERS: unisexual, perigoniate; tepals 4(–6), in 2 decussate whorls, fleshy, truncate to ± cucullate. MALE FLOWER: stamens with connate filaments forming tube around central, cylindric to clavate pistillode, often exserted above tepals at anthesis, connective slender, thecae dehiscing by apical pore. POLLEN: extruded in strands, extended monosulcate or fully zonate, hamburger-shaped, large (mean 76 µm., range 73–79 µm.), exine thick, foveolate, the foveolae scattered or grouped in fossulae, apertural exine psilate to ± verrucate. FEMALE FLOWER: usually lacking staminodes, exceptionally 1 staminode present, ovary 2-locular, ovules 1 per locule, anatropous, funicle very short to almost absent, placenta basal-axile, stylar region thick, somewhat attenuate, stigma large, discoid-hemispheric. BERRY: ovoid-ellipsoid, 1–2seeded, red or orange to yellow, or whitish. SEED: ovoid-ellipsoid, testa thin, smooth, embryo large, plumule lateral, superficial, endosperm absent. See Plates 33, 113D.

33. Gonatopus Gonatopus J.D. Hooker ex Engler in A. & C. De Candolle, Monogr. Phan. 2: 208 (1879). TYPE: G. boivinii (Decaisne) Engler (Zamioculcas boivinii Decaisne).

33. Gonatopus

Z A M I O C U L C A D E A E : G O N ATO P U S

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B D

EE

DD

M

Y

W

Z E

X AA

F

V FF

A

C T

U

K H

GG N

S

P R Q

CC

J

G

L

BB

Plate 33. Gonatopus. A, habit × 1/2; B, stamens, perigone removed × 5; C, detail of male flower × 5; D, gynoecium, longitudinal section × 5; E, infructescence × 2/3; F, seed, front view × 5; G, habit × 1/2; H, stamens, perigone removed × 5; J, detail of female flower × 5; K, gynoecium, longitudinal section × 5; L, habit × 1/2; M, leaflet × 1; N, inflorescence × 1; P, detail of male flowers × 2; Q, detail of female flowers × 2; R, stamens, perigone removed × 5; S, detail of male flower × 5; T, gynoecium, longitudinal section × 5; U, single tepal, three quarter view × 5; V, habit × 1/2; W, stamens, perigone removed × 5; X, male flower × 5; Y, gynoecium, longitudinal section × 5; Z, infructescence × 2/3; AA, seed, side view × 2; BB, habit × 1/2; CC,, inflorescence, lower part of spathe removed to display spadix; DD, stamens, perigone removed × 5; EE, single tepal, three quarter view × 5; FF, male flower × 5; GG, gynoecium, longitudinal section × 5. Gonatopus angustus: A, Nuvunga & Conjo 385 (K); B–D, Cult. Kew 1971–02232 (Kew spirit collection 45233); E, Mogg 30003 (K); F, Bogner s.n. (Kew spirit collection 45229); G. petiolulatus: G, Vollesen 4788 (K); H–K, Bogner 241 (Kew spirit collection 37580); G. clavatus: L–M, Milne-Redhead & Taylor 7670 (K & Kew spirit collection 22034); N–U, Milne-Redhead & Taylor 7670 (Kew spirit collection 22034); G. marattioides: V–AA, Bogner 247 (Kew spirit collection 45222 & 52143); G. boivinii: BB, Bullock 1323 (K); CC–GG, Bogner 127 (Kew spirit collection 29047.481 ).

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CHROMOSOMES: 2n = 34, 68. DISTRIBUTION: 5 spp.; tropical east and subtropical southeast Africa:– Kenya, Malawi, Mozambique, South Africa (Natal, Transvaal), Tanzania (incl. Pemba, Zanzibar), Zaïre, Zambia, Zimbabwe. ECOLOGY: tropical evergreen forest; geophytes, on forest floor, also in rocky crevices with humus deposits. NOTE: The leaf blade of G. petiolulatus, in which the lower pinnae are reduced to irregular, linear rudimentary leaflets, is intermediate in form between Gonatopus and Zamioculcas. In G. boivinii, the fallen leaflets may root and form new plants. ETYMOLOGY: Greek gony, gonatus (knee) and pous, podos (foot); in reference to the petiole geniculum in G. boivinii. TAXONOMIC ACCOUNTS: Engler (1905), Obermeyer & Bogner (1979), Mayo (1985a).

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Tribe Stylochaetoneae Tribe Stylochaetoneae Schott, Syn. Aroid. 132 (1856, “Stylochitoneae”). Laticifers absent; stem hypogeal, rhizomatous, roots often very thick, fleshy; higher order leaf venation reticulate; spathe tube margins connate; spadix mostly hidden within spathe, fertile to apex; flowers unisexual, perigoniate; perigone a single, cup-like (urceolate) structure; male flower usually with central peg-like pistillode, stamen filaments long, slender, filiform, anthers terminal, dehiscing by longitudinal slits, connective slender, pollen sometimes monosulcoidate, otherwise inaperturate; ovary 1–4 locular, ovules 1–many per locule, anatropous, placenta basal, parietal or axile, style thick, exserted from perigone, stigma capitate to thickly discoid; endosperm copious.

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34. Stylochaeton Stylochaeton Leprieur in Ann. Sci. Nat., ser. 2, Bot., 2: 184 (1834). TYPE: S. hypogeum Leprieur SYNONYM: [Stylochiton Schott, Aroideae 10 (1855), orth.var.]. Laticifers absent. HABIT: seasonally dormant or evergreen herbs, rhizome subterranean, horizontal to erect, sometimes stoloniferous, roots thick, spindle-shaped, often very fleshy, almost tuberous. LEAVES: 1 to several, cataphylls often conspicuously mottled and apically auriculate, sometimes persistent as fibrous mass. PETIOLE: sheath short to long, often purplish-spotted or banded. BLADE: lanceolate, ovate, ± rounded, cordate-sagittate, sagittate, hastate-sagittate or hastate; primary lateral veins pinnate or mostly arising basally, long-arcuate and running into marginal vein, lowermost primaries sometimes retrorse and then ascending, higher order venation reticulate. INFLORESCENCE: 1–4 in each floral sympodium, appearing before or with leaves, borne at or partially below ground level. PEDUNCLE: short, much shorter than petiole. SPATHE: erect, marcescent; tube: margins connate, often ventricose at extreme base, sometimes constricted between a lower and an upper inflated zone, rarely entire spathe narrowly cylindric; blade: lanceolate-elliptic, ± gaping or opening only by narrow longitudinal slit, often much thickened. SPADIX: free, shorter than spathe, female zone densely flowered, sometimes contiguous with male zone, often separated by axis bearing a few (to many) sterile or

bisexual flowers, male zone fertile to apex. FLOWERS: unisexual, perigoniate, borne in single basal whorl or in spirals; perigone a single cup-like (urceolate) structure. MALE FLOWER: perigone often with very thick, fleshy margins, stamens 2–7, free, filaments filiform, long, rarely much thickened apically, connective slender, sometimes slightly thickened, thecae oblong, dehiscing by longitudinal slit, pistillode central, cylindric to conoid, sometimes absent. POLLEN: inaperturate, or sometimes vestigially monosulcate, ellipsoid, large (mean 53 µm., range 43–58 µm.), exine foveolate-reticulate or subreticulate. FEMALE FLOWER: perigone usually greatly thickened and sticky-glandular or farinaceous on upper surface, ovary 1–4-locular, ovules 1–many per locule, anatropous, placenta basal, parietal or axile, stylar region thick, ± cylindric, exserted beyond perigone, stigma capitate to broadly discoid and massive. BERRY: borne at or below ground level in globose to cylindric infructescence, often rugose, fleshy, 1–few-seeded. SEED: ovoid to ellipsoid, slightly compressed, testa black, thin, costate, embryo axile, elongate, endosperm copious. See Plates 34, 114A. CHROMOSOMES: 2n = 28, 56. DISTRIBUTION: 17 spp.; tropical and southeast subtropical Africa:– Angola, Benin, Burkina Faso, Cabinda, Cameroon, Central African Republic, ?Chad, Equatorial Guinea (Bioko, Rio Muni), Ethiopia, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Malawi, Mali, Mozambique, Nigeria, Senegal, Sierra Leone, Somalia, South Africa (Transvaal, Natal), Sudan, Swaziland, Tanzania, Togo, Uganda, Zaïre, Zambia, Zimbabwe. ECOLOGY: tropical humid forests (evergreen species, e.g. S. zenkeri), tropical savannas and deciduous forests (seasonally dormant species, e.g. S. natalensis); geophytes. NOTES: Engler (1920a) recognized 2 sections:– sect. Stylochaeton (syn. sect. Cyclogyne) and sect. Spirogyne. ETYMOLOGY: Greek stylos (style) and chitôn (tunic); refers to connate perigone surrounding gynoecium. TAXONOMIC ACCOUNTS: Engler (1920a), Knecht (1983), Bogner (1984c), Mayo (1985a), Ntépé-Nyame (1988), Malaisse & Bamps (1994).

34. Stylochaeton

S T Y L O C H A E TO N E A E : S T Y L O C H A E TO N

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F

D

A

J

E K

G

B

C

H

L

Plate 34. Stylochaeton. A, habit × 1/2; B, flowering habit × 2/3; C, spadix × 2; D, habit × 1/2; E, habit × 1/2; F, infructescence × 1; G, habit × 1/2; H, spadix × 2; J, male flower × 8; K, female flower × 5; L, gynoecium, longitudinal section × 5. Stylochaeton lancifolius: A, Andrews 732 (K); B–C, Meikle 1294 (K & Kew spirit collection 27685); Schweinfurth 199 (K); Synge 220 (K & Kew spirit collection 58038); S. zenkeri: D, Letouzey 14413 (K); S. borumensis: E, Greenway & Kanuri 12751 (K); Verdcourt 3224A (K); S. bogneri: G, Bogner 145 (K); Lucas 253 (K); H–L, Bogner 148 (Kew spirit collection 7707).

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Tribe Dieffenbachieae Tribe Dieffenbachieae Engler in Nova Acta Acad. Leopold.Carol. 39: 148 (1876). SYNONYM: Tribe Bognereae Mayo & Nicolson in Taxon 33 (4): 689 (1984). Laticifers present, simple, articulated; terrestrial to helophytic, stem aerial, internodes distinct; spadix: female zone entirely adnate to spathe, laxly flowered, sterile male flowers (synandrodes) usually present between female and male zones; flowers unisexual, perigone absent; stamens connate into a truncate, prismatic synandrium, fused connectives strongly thickened, thecae lateral, pollen grains large; female flowers distant from one another, ovary 1–3 locular, ovules 1 per locule, anatropous, placenta basal or basal-axile.

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35. Dieffenbachia Dieffenbachia Schott in Wiener Z. Kunst 1829 (3): 803 (1829). TYPE: D. seguine (Jacquin) Schott (“seguinum”; Arum seguine Jacquin). SYNONYMS: Seguinum Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Maguirea A.D. Hawkes in Bull. Torrey Bot. Club 75: 635 (1948). HABIT: evergreen herbs, sometimes robust, stems erect to decumbent, sometimes ± rhizomatous, rooting proximally, erect and unbranched distally, internodes distinct, green, smooth, with conspicuous annular leaf scars. LEAVES: numerous, forming an apical crown. PETIOLE: sheath more than half as long as petiole or reaching blade. BLADE: oblong-ovate, elliptic to oblanceolate, dark to light green or sometimes variegated with white, silver, yellow or various shades of green; midrib thick, sulcate or prominent on upper surface, primary lateral veins pinnate, sometimes only weakly differentiated, running into margin, secondary laterals parallel-pinnate, connected by transverse tertiary veins. INFLORESCENCE: (1–)2-several in each floral sympodium, cataphylls short and usually inconspicuous. PEDUNCLE: shorter than petiole. SPATHE: persistent, slightly or distinctly constricted between tube and blade, green, lower part convolute into a usually rather long, persistent tube which splits longitudinally in fruit, upper part expanded into a short, erect or recurved blade. SPADIX: slightly shorter than spathe, female zone entirely adnate to spathe, enclosed within tube, laxly flowered, separated from male zone by subnaked axis with a few, scattered sterile male flowers with reduced staminodes, rarely fertile zones contiguous (D. humilis), male zone fertile to apex, free, subcylindric, densely flowered, erect. FLOWERS: unisexual, perigone absent. MALE FLOWER: stamens 4–5, connate into a subsessile, rhomboid to hexagonal synandrium, truncate at apex, sulcate laterally, anthers lateral, common connective thick, fleshy, thecae oblong-ellipsoid, dehiscing by short, apical, pore-like slit. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong or nearly spherical, large (mean 79 µm., range 54–99 µm.), exine almost perfectly psilate to obscurely verruculate and/or sparingly punctate-foveolate to densely foveolate, rarely coarsely tuberculate (D. parlatorei). STERILE MALE FLOWERS: composed of a whorl of (3–)4–5(–6), ± flattened, irregularly globose-ellipsoid, sometimes ± connate staminodes.

35. Dieffenbachia

FEMALE FLOWER: staminodes 4–5 in a whorl, white, clavate with rounded apices, spreading to erect, surrounding and longer than gynoecium; ovary stout, subglobose to ovoid, thickwalled, 1–3-locular, locule walls bulging outwards giving ovary distinctly lobed appearance when plurilocular, ovules 1 per locule, anatropous, placenta axile to basal, stylar region inconspicuous, stigma massive, almost as broad to broader than ovary, 2–3-lobed or subhemispheric (when unilocular), usually yellow, saturated with oily secretion at anthesis. BERRY: usually borne in arching infructescence, berries globose to 2–3-furrowed, stigma remnants persistent, 1–3-seeded, scarlet red to orange. SEED: globose to ovoid, testa smooth, green to blackish green, embryo large, endosperm absent. See Plates 35, 114B. CHROMOSOMES: 2n = 34, 68. DISTRIBUTION: 30 spp.; tropical and subtropical America, West Indies: Argentina (Corrientes, Misiones), ?Belize, ?Bolivia, Brazil (Amazonia, Central West, South), Colombia, Costa Rica, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad, Venezuela. ECOLOGY: tropical and subtropical humid forests, helophytes, at edges of stream banks, or terrestrial in forest leaf litter. ETYMOLOGY: named after J. Dieffenbach (1796–1863), head gardener at the Imperial Palace of Schönbrunn, where Schott was Director. TAXONOMIC ACCOUNTS: Engler (1915), Young (1986).

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Plate 35. Dieffenbachia. A, habit × 1/4; B, leaf × 1/3; C, inflorescence, nearside half of spathe removed × 2/3; D, detail of spadix showing sterile zone between male and female zones × 2; E, synandrium × 6; F, female flower × 6; G, gynoecium, transverse section × 6; H, infructescence × 2/3; J, habit, upper part of stem removed × 1/4; K, leaf × 1/2; L, detail of leaf venation × 5. Dieffenbachia seguine cv. Reginae: A, Cult. Kew 1950–48701; D. seguine: B, Bunting 2026A (K); D. seguine var. viridis: C–G, Cult. Kew 1973–13142 (Kew spirit collection 58927); D. seguine: H, Croat 65536 (Kew slide collection); D. elegans: J, Cremers 5730 (K); D. paludicola: K–L, Liesner 4006 (K).

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36. Bognera Bognera Mayo & Nicolson in Taxon 33: 690 (1984). TYPE: B. recondita (Madison) Mayo & Nicolson (Ulearum reconditum Madison). HABIT: evergreen herb with creeping rhizomatous stem, cataphylls marcescent, persistent and entire, drying reddish-brown. LEAVES: erect, several. PETIOLE: shortly-sheathed, pilose, weakly geniculate at apex and base. BLADE: ovate, base rounded, apex acute; primary lateral veins pinnate, ca. 12 per side, glabrous adaxially, minutely pilose abaxially especially along veins, running into marginal vein, secondary laterals arcuate and ± parallel to primaries, finer veins reticulate. INFLORESCENCE: solitary, erect. PEDUNCLE: shorter than petiole, subtended by 3 cataphylls. SPATHE: erect, broadly ovoid, convolute in lower two-thirds, not constricted, apiculate, persistent. SPADIX: subequal to spathe, fertile to apex or most apical flowers sterile, female zone laxly flowered and entirely adnate to spathe, separated from male zone by very sparsely flowered sterile zone, male zone densely flowered. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4-androus, stamens connate into a synandrium, synandria truncate, irregularly rhombic, somewhat domed, usually with central slit, common connective very thick, thecae ellipsoid-oblong, lateral, dehiscing apically by very broad slit. POLLEN: inaperturate, broadly ellipsoid, large (50–60 µm.), exine scabrate to nearly smooth (psilate). STERILE MALE FLOWERS: each usually consisting of 4 staminodes, those of lower flowers ± free and roundish, upper ones nearly completely connate and irregularly formed, sometimes all 4 connate with a central slit. FEMALE FLOWER: gynoecia arranged in somewhat distant spirals, ovary depressed globose, 1-locular, ovule 1, anatropous, funicle short, placenta basal, stigma sessile, discoid, broad. BERRY: unknown. SEED: unknown. See Plates 36, 114C. CHROMOSOMES: 2n = 34 (karyotype resembles that of Dieffenbachia). DISTRIBUTION: 1 sp.; Brazil (Amazonas), ?Peru. ECOLOGY: tropical humid forest (“terra firme”); terrestrial, creeping in leaf litter on sandy soil. NOTES: The mature shoot architecture seems to consist of repeating units of one small cataphyll, one larger cataphyll and one foliage leaf (T. Ray, pers. comm.). ETYMOLOGY: Named after Josef Bogner (born 1939). TAXONOMIC ACCOUNTS: Madison (1980, as Ulearum reconditum).

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Tribe Spathicarpeae Tribe Spathicarpeae Schott, Syn. Aroid. 214 (1856). Laticifers present, simple, articulated; seasonally dormant, stem tuberous, hypogeal; primary lateral veins of leaf or leaf lobes forming single marginal vein (except Spathantheum and Spathicarpa), higher order venation reticulate; spathe unconstricted; spadix fertile to apex (except Mangonia); flowers unisexual, perigone absent; male flower a synandrium of partially or completely connate stamens (sometimes completely free in Gorgonidium); in female flower the gynoecium surrounded by whorl of free staminodes or by cup-like synandrode, ovules 1 per ovary locule (except Mangonia); embryo relatively small, axile, endosperm copious.

36. Bognera

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37. Mangonia Mangonia Schott in Oesterr. bot. Wochenbl. 7: 77 (1857). TYPE: M. tweedieana Schott (“twedieana”). SYNONYMS: Felipponia Hicken in Anales Soc. Ci. Argent. 84: 242 (1917), non Felipponea Brotherus (1912); Felipponiella Hicken in Darwiniana 2: 30 (1928). HABIT: seasonally dormant small herbs, tuber depressed-globose, deeply embedded in soil. LEAVES: few to several. PETIOLE: sheath long. BLADE: linear-elliptic or broadly ovateelliptic to oblong-sagittate; primary lateral veins pinnate, few, running into margin, higher order venation reticulate. INFLORESCENCE: solitary, appearing before the leaves. PEDUNCLE: fairly long, mostly subterranean. SPATHE: erect, not constricted, lower part convolute into subcylindric tube, blade lanceolate to oblong, erect, gaping. SPADIX: subequal to spathe, sessile, female zone short, densely flowered, male zone less densely to laxly flowered especially in basal portion, either contiguous with female or separated by short ± naked axis, apical third to half of spadix more densely flowered and covered with sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–5-androus, stamens with filaments connate, forming stipitate or sessile synandrium, anthers free, connective inconspicuous, thecae ellipsoid to globose, dehiscing by a narrow slit or apical pore. STERILE MALE FLOWERS: synandrode stipitate, consisting of 3–5 subcapitate, basally connate staminodes. POLLEN: inaperturate, ellipsoid, medium-sized (40 × 25 µm.), exine areolate to subrugulate. FEMALE FLOWER: gynoecium surrounded by whorl of 3–4 clavate-spathulate staminodes, ovary globose to ovoid, 2–3-locular, ovules 2 per locule, anatropous, funicle short, placenta axile in upper part of septum, stylar region distinct, narrower than ovary, stigma 3-lobed or discoid, broader than style, concave centrally. BERRY: depressed-globose, in dense, subglobose to shortly cylindric infructescence. SEED: ellipsoid, somewhat compressed, testa smooth, embryo and endosperm unknown. See Plates 37, 114D. CHROMOSOMES: unknown.

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Plate 36. Bognera. A, habit × 2/3; B, detail of leaf venation and trichomes on abaxial surface × 15; C, inflorescence, nearside lower half of spathe removed × 1; D, synandrium, top view × 8; E, synandrium, side view × 8; F, staminode, top view × 8; G, staminode, side view × 8; H, gynoecium × 8; J, gynoecium, longitudinal section × 8. Bognera recondita: A–B, Bogner 1995 (K); C–J, Bogner 1995 (Kew spirit collection 57525).

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R Plate 37. Mangonia, A, habit showing inflorescence and petiole base only × 2/3; B, habit × 2/3; C, spadix × 1; D, synandrium, top view × 15; E, synandrium, side view × 15; F, synandrode × 15; G, gynoecium with associated staminodes × 15; H, gynoecium, longitudinal section × 15; J, infructescence × 1; K, leaf × 2/3; L, inflorescence × 1; M, spadix × 2; N, synandrium, top view × 15; P, synandrium, side view × 15; Q, synandrode × 15; R, gynoecium with associated staminodes × 15; S, gynoecium, longitudinal section × 15. Mangonia uruguaya: A, Darwiniana 18: 73, fig. 1,1; B, Darwiniana 18: 74, fig. 2,2; C–H, Felippone 5772 (SI); Felippone SI 297 (K, Kew spirit collection 58120 & SI); J, Darwiniana 18: 74, fig. 2,3 (1973); M. tweedieana: K, Waechter 2347 (K); L–S, Tweedie s.n. (K, LE); Waechter 2347 (K & Kew spirit collection 58090).

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37. Mangonia

DISTRIBUTION: 2 spp.; warm temperate South America:– Brazil (Rio Grande do Sul), Uruguay. ECOLOGY: subtropical gallery forest; geophytes, stony, well drained soils, flat areas, slopes near water courses. NOTES: The first reliable record of M. tweedieana for Brazil was recently made by Prof. Jorge Waechter (Waechter 2347 [ICN, K!]), who also provided important new information contained in this generic treatment. ETYMOLOGY: Latin mango, mangonis (dealer, hence English monger). TAXONOMIC ACCOUNTS: Engler (1920a), Herter (1943), Bogner (1973a).

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38. Taccarum Taccarum Brongniart ex Schott in Oesterr. bot. Wochenbl. 7: 221 (1857). TYPE: T. weddellianum Brongniart ex Schott SYNONYMS: Lysistigma Schott in Bonplandia 10: 222 (1862); Endera Regel in Gartenflora 21: 226 (1872). HABIT: seasonally dormant herbs, often robust, tuber depressed-globose. LEAF: solitary. PETIOLE: terete, mottled and variegated or not, sheath very short. BLADE: juvenile leaves simply lobed, adult leaves subdracontioid, trifid to trisect, anterior division usually deeply bipinnatifid, often tripinnatifid in lower pinnae, posterior divisions deeply pedatifid with segments themselves pinnatifid, ultimate lobes subtriangular to elliptic or lanceolate, acute to acuminate, broadly decurrent; basal ribs well-developed, primary lateral veins of ultimate lobes pinnate, running into inconspicuous marginal vein, higher order venation reticulate. INFLORESCENCE: 1, rarely 2 in each floral sympodium. PEDUNCLE: usually much shorter than petiole. SPATHE: not constricted, tube convolute, blade gaping to widely spreading, marcescent

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38. Taccarum

and later deciduous (T. weddellianum). SPADIX: free or female zone adnate to spathe, sessile or stipitate, erect, much longer than, subequal or shorter than spathe, male zone usually contiguous with female, rarely with a few bisexual flowers in between, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–8-androus, stamens connate, synandrium very long-stipitate with apical whorl of anthers to short with anthers near base, stigmatoid apex inconspicuous or distinct and 4–6-lobed, or very large and dome-shaped, thecae oblong or broadly ellipsoid, dehiscing by short apical slit or pore. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, large (mean 63 µm., range 49–76 µm.), exine scabrate or verruculate or spinulose-spinose, or smooth (psilate). FEMALE FLOWER: gynoecium surrounded by whorl of 4–6 free, erect, filiform, clavate or oblong, often flattened staminodes, or by urceolate synandrode composed of connate staminodes (T. caudatum), ovary 3–6(–7)-locular, ovules 1 per locule, anatropous, funicle short, placenta axile, style very long and slender or very short to ± absent, always narrower than ovary, stigma thick, capitate or 5–7-lobed, lobes erect or stellate. BERRY: borne in cylindric infructescence, depressed-globose, slightly furrowed, stylar region persistent, 3–5-seeded. SEED: ellipsoid, raphe conspicuous, testa granulate, light brown, embryo straight, elongate, endosperm copious. See Plates 38, 115A. CHROMOSOMES: 2n = 34. DISTRIBUTION: 5 spp.; tropical and subtropical South America:– N. Argentina (Misiones), Bolivia, Brazil, Paraguay, Peru. ECOLOGY: open tropical woodland (cerrado), forests, humid mountain valleys; geophytes, often in stony soils or between rocks. ETYMOLOGY: Malay taka (name for Tacca in the Taccaceae) and Arum; refers to similarity of the leaf to that of Tacca leontopetaloides. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1989a).

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Plate 38. Taccarum. A, leaf × 6; B, flowering habit, leaf removed × 1/8; C, spadix × 1/2; D, synandrium × 3; E, female flower × 3; F, gynoecium, longitudinal section × 3; G, gynoecium, tranverse section × 3; H, infructescence × 1/2; J, habit × 1/5; K, inflorescence, nearside half of spathe removed × 1/3; L, synandrium × 3; M, female flower × 3; N, gynoecium, longitudinal section × 3. Taccarum weddellianum: A, Pirani et al. 1946 (K); B, Cult. Hetterscheid (Kew slide collection); C–G, Bogner 458 (Kew spirit collection 29047.741); H, Harley 20436A (Kew spirit collection 29047.453); T. warmingii: J, Type drawing (Kew illustration collection); K–N, Cult. Kew 9 April 1890 (K & Kew spirit collection 58086).

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39. Asterostigma Asterostigma F.E.L. Fischer & C.A. Meyer in Bull. Cl. Phys.Math. Acad. Imp. Sci. Saint Pétersbourg, ser. 2, 3: 148 (1845). TYPE: A. langsdorffianum F.E.L. Fischer & C.A. Meyer SYNONYMS: Staurostigma Scheidweiler in Allg. Gartenzeitung 16: 129 (1848); Andromycia A. Richard in R. de la Sagra, Hist. Fis. Cuba 11: 282 (1850); Rhopalostigmium Schott in Oesterr. bot. Zeitschr. 9: 39 (1859); [Rhopalostigma B.D. Jackson, Index Kew. 2: 713 (1895), orth. var., non R.A. Philippi (1860)]. HABIT: medium-sized, seasonally dormant herbs, tuber depressed-globose. LEAVES: usually solitary. PETIOLE: smooth, maculate, sheath very short. BLADE: usually pinnatisect, rarely entire, lobes oblong-lanceolate, acute to acuminate; primary lateral veins of ultimate lobes pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–3 in each floral sympodium, appearing with or before leaves, cataphylls often beautifully variegated. PEDUNCLE: rather long, similar to petiole. SPATHE: erect, hardly to not constricted, slender, persistent, lower part convolute into narrowly cylindric tube, blade gaping to widely expanded at anthesis. SPADIX: female zone free or partly adnate to spathe, laxly flowered, contiguous with male zone, male zone longer, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 3–4-androus, surrounding and concrescent with pistillode, subhexagonal to rounded, sometimes very shallow, umbonate at apex, shortly stipitate or sessile, thecae subglobose, distant, almost pendent from upper margin of synandrium, dehiscing by transversely elongated pore. POLLEN: inaperturate, ellipsoid to ellipsoid-oblong, mediumsized (mean 36 µm., range 35–38 µm.), virtually psilate or coarsely verrucate. FEMALE FLOWER: gynoecium surrounded by cup-like synandrode, or by whorl of 3–5 short, somewhat thickened, subcuneate, truncate, often irregularly connate staminodes, ovary pear-shaped to depressed-globose, 3–5-locular, ovules 1 per locule, anatropous, funicle

39. Asterostigma

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short, placenta axile at base of septum, stylar region distinct, stigma usually deeply and stellately 3–5-lobed with each lobe itself lobed, or 2–3-lobed with oblong, entire lobes. BERRY: subglobose to depressed-globose, deeply 4–5-sulcate, pale to whitish green. SEED: oblong to ellipsoid, with swollen arillike funicle, testa nearly smooth, embryo axile, elongate, endosperm copious. See Plates 39, 115B. CHROMOSOMES: 2n = 34. DISTRIBUTION: ca. 7 spp.; tropical and subtropical South America:– Bolivia, Brazil (Central, Northeast, Southeast, South), Ecuador, Peru. ECOLOGY: tropical and subtropical humid forests; geophytes, forest floor. NOTES: Engler (1920a) recognized 2 sections: sect. Asterostigma, sect. Rhopalostigma. ETYMOLOGY: Greek astêr (star) and stigma (a mark), refers to the star-shaped stigmas. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1969), Bogner (in press).

40. Gorgonidium Gorgonidium Schott in Ann. Mus. Bot. Lugduno-Batavum 1: 282 (1864). TYPE: G. mirabile Schott HABIT: seasonally dormant herbs, tuber depressed globose. LEAF: solitary. PETIOLE: terete, sheath short. BLADE: pinnatifid, pinnatisect or bipinnatifid, pinnae (9–)11–14, elliptic, entire or pinnatifid (with all intermediates), acute, upper lobes decurrent, lower ones sessile to shortly stalked; primary lateral veins of ultimate lobes pinnate, running into margin, higher order venation reticulate. INFLORESCENCE: 1(–2) per floral sympodium, appearing before leaf. PEDUNCLE: much shorter than petiole. SPATHE: erect or slightly fornicate, boat-shaped, very shortly convolute at base, not constricted, purple, persistent at least in lower part. SPADIX: sessile or stipitate (G. mirabile), fertile to apex, female zone contiguous with and much shorter than male zone. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–7-androus, stamens free or connate to different degrees into a synandrium, connective slender, sometimes stipe-like between thecae (G. mirabile), otherwise inconspicuous, thecae globular, sometimes remote from one another, dehiscing by apical pore or slit, pistillode present or absent, if present composed of 3–4 stylodia with slightly lobed or discoid-capitate apices (stigmatoids). POLLEN: inaperturate, ellipsoid, medium-sized (mean 34 µm, range 27–42 µm.), exine verrucose (G. mirabile) to retiverruculate, sometimes verrucae irregularly formed and flattened (G. vargasii). FEMALE FLOWERS: gynoecium surrounded by whorl of 6–8 filiform or subclavate staminodes, ovary subglobular to broadly ovoid, (2–)4–5(–7)-locular, ovules 1 per locule, orthotropous, ovoid, funicle half as long as ovule, placenta axile at base of septum, stylar region short to long, stigma 4–5-lobed or subcapitate. BERRY: globular to depressed-globular, blackish-purple to purple, sometimes upper surface somewhat warty (G. vargasii), stigma and style remnants persistent. SEED: ovoid, globular to ellipsoid, testa brownish, smooth to rough, embryo elongate, endosperm copious. See Plates 40, 115C. CHROMOSOMES: 2n = 34. DISTRIBUTION: 3 spp.; Western and Andean South America:– N. Argentina, Bolivia, Peru. ECOLOGY: tropical and subtropical forest or open places, between 900–3000m; geophytes, stony ground.

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Plate 39. Asterostigma. A, leaf × 1/2; B, detail of leaf venation × 5; C, base of plant × 1/2; D, juvenile infructescence enclosed by persistent spathe × 1/2; E, spadix × 2; F, female flower × 10; G, synandrium × 10; H, gynoecium, longitudinal section × 15; J, young plant × 2/3; K, spadix × 2; L, female flower × 10; M, synandrium × 10; N, female flower × 10; P, gynoecium, longitudinal section × 15; Q, infructescence × 1; R, leaf × 1/2; S, base of plant × 1/2. Asterostigma cryptostylum: A–H, Bogner 1237 (K & Kew spirit collection 48202); A. riedelianum: J–P, Harley et al. 18565 (K & Kew spirit collection 39166, 46592, 47693 & Kew slide collection); A. integrifolium: Q–S, Rawlins 204 (K).

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Plate 40. Gorgonidium. A, vegetative habit × 1/5; B, flowering habit, tuber longitudinally split; C, spadix × 2; D, synandrium × 8; E, female flower × 8; F, gynoecium, longitudinal section × 8; G, gynoecium, transverse section × 8; H, leaf × 2/3; J, infructescence × 2/3; K, synandrium × 3; L, female flower × 3; Gorgonidium vermicidum: A, Bot. Jahrb. Syst. 109:550, fig. 23 (1988); B, Araque & Barkley s.n. (K); Aichinger 2 (Kew slide collection); C–G, Bogner 101(Kew spirit collection 45269); G. vargasii : H, Brunel 557 (K); J, Vargas s.n. (Kew spirit collection 49725); G. mirabile : K–L, Nicolson 3390 (K & Kew spirit collection 58918).

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40. Gorgonidium

ETYMOLOGY: Greek Gorgo (the Grim One), -ides (descendant) and -ion (diminutive); the name alludes to the filiform staminodia and stamens in G. mirabile, the appearance of which recalls mythic Gorgo, whose hair was a mass of writhing serpents. TAXONOMIC ACCOUNTS: Bogner & Nicolson (1988).

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41. Synandrospadix Synandrospadix Engler in Bot. Jahrb. 4: 61 (1883). TYPE: S. vermitoxicus (Grisebach) Engler (Asterostigma vermitoxicum Grisebach). SYNONYMS: Lilloa Spegazzini, Pl. Nov. Crit. Argentina 3: 10 (1897); [Synandriospadix Engler in Engler, Pflanzenreich 73 (IV.23F): 49 (1920), orth. var.]. HABIT: seasonally dormant herb, tuber subglobose. LEAVES: several. PETIOLE: fairly long, with darker longitudinal striations, sheath long. BLADE: ovate-cordate to -emarginate, somewhat glaucous; primary lateral veins pinnate, running into inconspicuous marginal vein, higher order venation reticulate. INFLORESCENCE: 1, rarely 2 in each floral sympodium, usually appearing with leaves. PEDUNCLE: short. SPATHE: ovate-lanceolate, boat-shaped, unconstricted, persistent, convolute at base, gaping above, light green outside with dark longitudinal lines, inside purple and roughened. SPADIX: much shorter than spathe, subcylindric, obtuse, female zone partly or completely adnate to spathe, ± laxly flowered, separated from male zone by a few bisexual or sterile flowers, male zone longer, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 4–5-androus, long-stipitate, stipe elongate-conoid, whorl of anthers apical or shortly overtopped by central pistillode, thecae oblong-ellipsoid, dehiscing by longitudinal slit. POLLEN: purple to pinkish, inaperturate, spherical to subspheroidal, medium-sized (mean 49 µm.), exine spinose. BISEXUAL FLOWERS: as for female but with whorl of 4–5 free stamens with flattened elongate-triangular filaments and

41. Synandrospadix

thecae dehiscing as above. FEMALE FLOWER: gynoecium surrounded by 3–5 free, elongate-triangular, flattened, acute staminodes, ovary ovoid-globose, 3–5-locular, ovules 1 per locule, orthotropous, funicle distinct, placenta axile-basal, stylar region attenuate, stigma discoid-subcapitate, slightly 3–5-lobed. BERRY: ± subglobose, 3–5-furrowed, 3–5-seeded, style persistent. SEED: rather large, ovoid to obnapiform, testa ± rough, embryo elongate, straight, endosperm copious. See Plates 41, 115D. CHROMOSOMES: 2n = 34. DISTRIBUTION: 1 sp.; N. Argentina, Bolivia, Paraguay, Peru. ECOLOGY: subtropical dry thorn forest; geophytes in clearings or on stony ground in shade. ETYMOLOGY: Greek syn (together), anêr, andros (man) and spadix (spadix). TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), Croat & Mount (1988).

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42. Gearum Gearum N.E.Brown in J. Bot. 20: 196–197 (1882). TYPE: G. brasiliense N.E.Brown HABIT: seasonally dormant, tuberous herbs with aromatic exudate when cut. LEAF: solitary. PETIOLE: sheath distinct, about half length of petiole. BLADE: subpalmatifid to pedatisect, lobes or segments 5–9; primary lateral veins of each lobe forming submarginal collective vein on each side, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: very short, much shorter than petiole. SPATHE: erect, slightly constricted, purplish, tube subcylindric, convolute, blade erect, gaping, oblong, cuspidate. SPADIX: sessile, shorter than spathe, fertile to apex, female zone shorter than male and separated from it by a sterile zone of synandrodes. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4-androus, synandrium very shallow, subrhombic, truncate, subpeltate or flat and not subpeltate, thecae ± remote, situated near margin of synan-

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A

E

D F

G

H

J

C

K B

Plate 41. Synandrospadix. A, habit × 1/4; B, juvenile plant × 1/2; C, leaf × 2/3; D, inflorescence × 2/3; E, spadix × 1; F, synandrium × 6; G, bisexual flower × 6; H, female flower × 6; J, gynoecium, longitudinal section × 6; K, infructescence × 1/2. Synandrospadix vermitoxicus: A, 184/10 (Kew slide collection); B, Cult. Kew 9 October 1891 (K); C, Araque & Barkley 440 (K); D–F, H–J, Bogner s.n. (Kew spirit collection 56099); G, Burkart et al. 30412 (K); K, Badcock 815 (K).

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42. Gearum

drium, globose to oblong-ellipsoid, dehiscing by apical pore or occasionally by an oblique slit. POLLEN: ellipsoid, inaperturate, large (mean 56µm, range 52–60 µm), exine smooth (psilate) to scabrous-granulate. STERILE MALE FLOWERS: composed of irregularly elongated, shallow, truncate synandrodes. FEMALE FLOWER: densely crowded, gynoecium surrounded by usually 4 staminodes, staminodes laterally compressed, thick, rounded to obovoid or subtrapezoid, ovary subglobose, 3–4-locular, ovules 1 per locule, orthotropous, rather elongate, placenta axile at base of septum, stigma subsessile, ± discoid, slightly (3–)4-lobed. BERRY: unknown. SEED: unknown. See Plate 42. CHROMOSOMES: unknown. DISTRIBUTION: ?2 spp.; Brazil (Central–West). ECOLOGY: tropical open herbaceous vegetation (cerrado , in campo limpo); geophytes in low-lying sites prone to periodic flooding. ETYMOLOGY: Greek gê (earth) and Arum. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner & Nicolson (1988), Mayo, Bogner & Boyce (1994).

H

D F

B

C

G

A

E

Plate 42. Gearum. A, leaf × 1/3; B, inflorescence × 1/3; C, spadix × 2/3; D, synandrium, top view × 8; E, synandrium, side view × 8; F, gynoecium with staminode, side view × 8; G, gynoecium, longitudinal section × 8. H, leaf × 1/3. Gearum sp .: A–G Araujo Dias 41 (HRB & Kew spirit collection 57663); Gearum brasiliense : H from Hatschbach 56028 (K).

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43. Spathantheum Spathantheum Schott in Bonplandia 7: 164 (1859). TYPE: S. orbignyanum Schott SYNONYM: Gamochlamys J.G. Baker in Saunders Refug. Bot. 5: t. 346 (1873). HABIT: medium-sized, seasonally dormant herbs, tuber depressed-globose, sometimes large. LEAVES: solitary. PETIOLE: sheath very short. BLADE: outline ovate to ovatecordate, entire or pinnatifid; primary lateral veins pinnate, running into margin, higher order venation reticulate. INFLORESCENCE: usually 1, rarely 2 in each floral sympodium, flowering before leaf. PEDUNCLE: short (S. intermedium) to long (S. orbignyanum), slender. SPATHE: oblong-elliptic, boat-shaped, not constricted, cuspidate, at first convolute basally, then widely gaping at anthesis, green or purple (S. intermedium), entirely persistent. SPADIX: entirely adnate to spathe (S. orbignyanum) or male part free (S. intermedium), slightly shorter than spathe, densely flowered, either female zone basal, contiguous with longer male zone and spadix fertile to apex (S. inter medium, S. orbignyanum), or female and male zones sometimes separated by a zone of mixed male and female flowers (S. orbignyanum). FLOWERS: unisexual, perigone absent. MALE FLOWER: 4–7-androus, stamens either connate into a long-stipitate, apically convex, peltate and shallowly 5–7lobed (stigmatoid) synandrium with anthers almost pendent from thickened common connective, or filaments connate, anthers free and stigmatoids elongated, overtopping anthers (S. intermedium), thecae linear-oblong and dehiscing by longitudinal slit (S. orbignyanum) or subglobose and dehiscing by apical pore (S. intermedium). POLLEN: inaperturate, ellipsoid-oblong, large (mean 54 µm.), exine nearly psilate or shallowly uneven. FEMALE FLOWER: gynoecium surrounded by whorl of 5–8 terete-clavate, short staminodes, ovary ovoid-ellipsoid, 5–8-locular, ovules 1 per locule,

43. Spathantheum

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suborthotropous, funicle distinct, placenta axile at base of septum, stylar region elongate-attenuate, stigma stellate, 6–8-lobed, lobes subtriangular. BERRY: subglobose to depressed-globose, 5–8-seeded. SEED: shortly ovoid to ± ellipsoid, testa ± smooth to slightly roughened, embryo elongate, endosperm copious. See Plates 43, 116A. CHROMOSOMES: 2n = 34. DISTRIBUTION: 2 spp.; Andean South America:– N. Argentina, Bolivia, Peru. ECOLOGY: tropical and subtropical uplands, mountain grasslands, up to 2400m; geophytes on stony ground in shade, in deposits of humus on rocks. ETYMOLOGY: Greek spathê, anthos (flower) and -eum (possessing); refers to the fusion of the spadix to the spathe. TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), Bogner (in press).

44. Spathicarpa Spathicarpa W.J. Hooker in Bot. Misc. 2: 146 (1831). TYPE: S. hastifolia W.J. Hooker SYNONYMS: [Spaticarpa Schott in Oesterr. bot. Zeitschr. 15: 34 (1865), orth. var.]; Aropsis Rojas Acosta in Bull. Acad. Int. Géogr. Bot. 28: 158 (1918). HABIT: small, seasonally dormant, sometimes evergreen herbs, tuber subterranean, rhizomatous, shortly oblong, horizontal. LEAVES: several. PETIOLE: slender, sheath often rather long. BLADE: narrowly elliptic, oblong, ovate-emarginate, ovate-cordate, cordate, cordate-sagittate, auriculate-hastate or strongly hastate to subtrisect; basal ribs short when present, primary lateral veins pinnate or mostly arising at petiole insertion, arcuate and running into margin or forming an irregular, submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: relatively long, slender. SPATHE: not constricted,

44. Spathicarpa

C

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J

E

F

G

H D

K

L

M

N

P

B

C

A Plate 43. Spathantheum. A, flowering habit × 2/3; B, juvenile leaf × 2/3; C, mature leaf × 2/3; D, inflorescence, nearside half of spathe removed × 2/3; E, synandrium × 4; F, female flower × 4; G, gynoecium, longitudinal section × 4; H, gynoecium, transverse section × 6; J, infructescence × 2/3; K, inflorescence, nearside half of spathe removed × 2/3; L, synandrium × 4; M, gynoecium with associated staminodes (= female flower) × 4; N, gynoecium, longitudinal section × 4; P, gynoecium, transverse section × 6. Spathantheum orbignyanum: A, Bang 1626 (K), Rauh 26024 (Kew spirit collection 37322); B, Bogner 900 (K); C, Cult. Kew 1883 (K); D–H, Rauh 26024 (Kew spirit collection 37322); Bogner 900 (K & Kew spirit collection 37369); S. intermedium: K–P, Munn 148 (Kew spirit collection 29047.761).

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B

C

E

K

G

D

A

F H

J

Plate 44. Spathicarpa. A, habit × 2/3; B, synandrium and gynoecium with associated staminodes (= male and female flowers) × 10; C, gynoecium, longitudinal section × 16; D, leaf × 2/3; E, habit × 2/3; F, inflorescence × 2; G, detail of spadix × 6; H, synandrium and gynoecium with associated staminodes (= male and female flowers) × 10; J, gynoecium, longitudinal section × 16; K, infructescence, nearside half of spathe removed × 1. Spathicarpa burchelliana: A, Burchell 8335(K); Gardner 2447 (K); B–C, Cult. Kew 1981–388, Cristobal & Krapovickas s.n. (Kew spirit collection 45662); S. lanceolata: D, Balansa 579 (K); S. hastifolia (S. cf. gardneri ): E–K, Tressens et al. 1569 (K); Tressens et al. 3471 (K).

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oblong-lanceolate to narrowly elliptic or oblanceolate, acuminate, fully expanded to subrevolute at anthesis, later closing, persistent, green. SPADIX: entirely adnate to spathe, recurved at anthesis, fertile to apex, ± laxly flowered, female flowers forming 2 outer longitudinal rows, enclosing 2 rows of male flowers, all in parallel. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3-4-androus, stamens connate, synandrium stipitate, apex formed by nectariferous, usually shallowly lobed stigmatoid-connective, anthers short, partly covered and overtopped by stigmatoid-connective, thecae 6-8, broadly ellipsoid, lateral, dehiscing by subapical pore. POLLEN: extruded in strands, inaperturate, ellipsoid-oblong to -elongate, medium-sized (mean 48 µm., range 45-51 µm.), exine almost perfectly psilate. FEMALE FLOWER: gynoecium partially surrounded on one side by 3 small, very shortly stipitate, discoid-umbonate, green staminodes, ovary oblong-ovoid, 1-locular, ovule 1, orthotropous, funicle very short to ± absent, placenta basal, stylar region attenuate, cylindricconoid, stigma discoid-subcapitate. BERRY: ovoid to ellipsoid, acute-attenuate, green. SEED: ovoid-obnapiform, strophiolate, testa smooth to ± rough, brownish to green, embryo axile, elongate, endosperm copious. See Plates 44, 116B. CHROMOSOMES: 2n = 34. DISTRIBUTION: ca. 5 spp.; tropical and subtropical South America:– N. Argentina, Bolivia, Brazil (Northeast, Central West, Southeast, South), Paraguay, Uruguay. ECOLOGY: tropical dry forest, humid and marshy forest, seasonally wet places; geophytes; in dry forest areas the plant has a dormant period and starts growth at the beginning of the rainy season. ETYMOLOGY: Greek spathê (spathe) and karpos (fruit); refers to the fusion of spathe and spadix (in fruit). TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), Uhlarz (1983), Croat & Mount (1988).

C

HABIT: evergreen herbs, small to gigantic, stem repent to rhizomatous, climbing, arborescent or plant rosulate and acaulescent, internodes usually long, often short to very short, intravaginal squamules present, sometimes producing flagelliform shoots. LEAVES: numerous, small to gigantic, prophylls of mature stems caducous, marcescent and deciduous or persistent and membranaceous or decomposing to net-fibrous remains. PETIOLE: sometimes warty or covered with scale-like processes, sometimes swollen, rarely geniculate apically, sheath long and slilghtly ligulate in monopodial leaves of all subgenera and in sympodial leaves of subgen. Pteromischum, otherwise very short and inconspicuous except when subtending inflorescences. BLADE: very variously shaped; simple and linear, cordate, sagittate or hastate, or trifid, trisect, pinnatifid, bipinnatifid, rarely pedatisect, resin canals linear, short to long, obscured to very distinct on abaxial surface; basal ribs sometimes well-developed, primary lateral veins pinnate, rarely pedate, running into marginal vein, secondary lateral and higher order venation parallel-pinnate, sometimes tertiaries and higher order veins transversely reticulate between secondaries, sometimes all veins slender with no distinct primary laterals. FLOWERING BRANCHES: sympodial articles of three main patterns:– subgen. Pteromischum: prophyll, many foliage leaves, 1–2(–3) inflorescences; subgen. Philodendron: prophyll, following internode suppressed, 1 foliage leaf, 1–11 inflorescences, internode to prophyll of continuation shoot elongated; subgen. Meconostigma: prophyll, following internode developed or very short, 1 foliage leaf, 1(–2) inflorescences, internode to prophyll of continuation shoot suppressed. INFLORESCENCE: 1–11 in each floral sympodium, secreting resin at anthesis, either from spathe or from spadix, rarely from both. PEDUNCLE: usually much shorter than petiole. SPATHE: erect, entirely persistent, deciduous only at ripening of fruit (caducous after anthesis in P. surinamense), fairly thick,

Tribe Philodendreae Tribe Philodendreae Schott, Syn. Aroid. 71 (1856). Laticifers present, simple, articulated, resin canals present in roots, stems, leaves and inflorescences, sclerotic hypodermis present in roots; diminutive to gigantic, climbing hemiepiphytes, epiphytes or terrestrial, stem usually epigeal, intravaginal squamules present; petiole only rarely geniculate apically; primary lateral veins pinnate (very rarely pedate) forming 1 marginal vein, higher order venation parallel-pinnate; spathe persisting until fruit maturity, then deciduous at base; flowers unisexual, perigone absent; anther lacking cell wall thickenings in endothecium (except P. goeldii, P. lealcostae); endosperm copious.

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45. Philodendron Philodendron Schott in Wiener Z. Kunst 1829 (3): 780 (1829), nom. et orth. cons. (‘Philodendrum’). LECTOTYPE: P. grandifolium (Jacq.) Schott (see Britton & Wilson 1923). SYNONYMS: Philodendrum Schott in Wiener Z. Kunst 1929 (3): 780 (1829), orth. rej.; Arosma Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Telipodus Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Thaumatophyllum Schott in Bonplandia 7: 31 (1859); Elopium Schott in Oesterr. bot. Zeitschr. 15: 34 (1865); Baursea Post & O. Kuntze, Lexicon Gen. Phanerog. 62 (1903).

45. Philodendron

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E D

C

A K B

G F

J H

Plate 45 (i). Philodendron. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2; H, leaf × 1/2; J, leaf × 1/2; K, leaf × 1/2. Philodendron callosum: A, Granville et al. 10487 (K); P. crassinervium : B; Hatschbach 45979 (K); P. fibrillosum: C, Plowman et al. 11406 (K); P. calatheifolium: D, Bunting 11659 (K); P. heterophyllum: E, Whitmore 748 (K); P. blanchetianum: F, Storr 13 (K); P. grazielae: G, Dodson 2718 (K); P. frits-wentii: H, Madison 4175 (K); P. aromaticum: J, Croat 68423 (K); P. sp. cf. verrucosum: K, André 2629 (K).

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B

C

A

D

E

Plate 45 (ii). Philodendron. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/3; D, leaf × 1/3; E, leaf × 1/6. Philodendron bipennifolium: A, Engler 237 (K); P. anisotomum: B, Heyde & Lux 4283 (K); P. goeldii: C, Stevenson 829 (K); P. angustisectum: D, Cult. Kew 1984–1076; E, P. bipinnatifidum: Mayo et al. 574 (K).

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A

C B

D

E

F

Plate 45 (iii). Philodendron. A, habit × 1/6; B, leaf × 1/12; C, habit × 1/12; D, habit × 1/3; E, detail of lower stem showing intravaginal squamules × 1/2; F, habit × 1/12. Philodendron insigne: A, Cult. Kew. 1980–2229; P. verrucosum : B, Boyce s.n. (Kew slide collection); P. bipinnatifidum: C, Cult. Kew 1983–2024; P. scandens: D, Cult. Kew 1953–43201; P. uliginosum: E, Kew Bull. 46(4): 601–681, f.3, A (1991); P. melinonii: F, Cult. Kew 1970–84.

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B

A

D

C

Plate 45 (iv). Philodendron. A, inflorescence and associated leaf × 1/2; B, infructescences and associated stem and petiole bases × 1/2; C, floral sympodium and associated stem and petiole base × 1/2; D, inflorescence and associated petiole base × 1/2. Philodendron rigidifolium: A, Sugden 585 (K); P. leal-costae: B, Harley et al. 19428 (Kew slide collection); P. verrucosum: C, Knapp 4959 (K); McPherson 9033 (K); P. bipinnatifidum: D, Boyce s.n. (Kew slide collection).

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B L

G C

N

M

D

E

H

K J Q

DD

A F

R

S

T

EE

AA V

X W FF

CC

BB

P Z

Y

U

Plate 45 (v). Philodendron. A, spadix × 1; B, stamen × 10; C, gynoecium × 10; D, gynoecium, longitudinal section × 10; E, gynoecium, transverse section × 10; F, spadix × 1; G, stamens × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, spadix × 1; L, stamen × 10; M, gynoecium × 10; N, gynoecium, longitudinal section × 10; P, spadix × 1; Q, stamen × 10; R, gynoecium × 10; S, gynoecium, longitudinal section × 10; T, gynoecium, transverse section × 10; U, spadix × 1; V, stamen × 10; W, gynoecium × 10; X, gynoecium, longitudinal section × 10; Y, gynoecium, transverse section × 10; Z, spadix × 2/3; AA, gynoecium × 5; BB, gynoecium, longitudinal section × 5; CC, spadix × 2/3; DD, stamen × 8; EE, gynoecium × 8; FF, gynoecium, longitudinal section × 8. Philodendron bipennifolium: A–E, Mayo 599 (Kew spirit collection 46286); P. melinonii: F–J, Evemy & Burgess 161 (Kew spirit collection 29047.428); P. blanchetianum: K–N, Storr 13 (Kew spirit collection 48471); P. eximium: P–T, Boyce s.n. (Kew spirit collection 51441); P. inaequilaterum: U–Y, Cult. Kew 1964–39505 (Kew spirit collection 49977); P. goeldii: Z–BB, Bogner 233 (Kew spirit collection 29047.740); P. bipinnatifidum: CC–FF, Boyce s.n. (Kew spirit collection 29047.785).

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sometimes (subgen. Meconostigma) extremely thick, usually constricted between tube and blade, tube convolute, cylindric to ventricose, often coloured purple or red within, blade usually boat-shaped, widely gaping at anthesis, later closing, usually white within, rarely red. SPADIX: sessile to stipitate, female zone free, rarely basally adnate to spathe, usually shorter than male zone and separated from it by intermediate sterile zone of staminodial flowers, intermediate sterile zone cylindric or constricted or ellipsoid and thicker than male zone, usually shorter than male zone, sometimes longer (subgen. Meconostigma), a terminal staminodial appendix sometimes also present. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–6-androus, stamens free, prismatic to obpyramidal, sometimes very elongated and slender (subgen. Meconostigma), anthers sessile to subsessile, connective thick, apically truncate, overtopping thecae, thecae ellipsoid to oblong, dehiscing by short lateral slit or by subapical pore, endothecial thickenings lacking (except P. goeldii, P. lealcostae). POLLEN: extruded in strands or mixed with resin secretion or exuded in amorphous masses, inaperturate, ellipsoid to oblong or occasionally elongate, medium-sized (mean 40 µm., range 28–54 µm.), mostly perfectly psilate, sometimes from minutely verruculate, scabrate or fossulate to clearly punctate, subfossulate, subfoveolate or subverrucate, rarely densely and coarsely verrucate (P. leal-costae). STERILE MALE FLOWERS: staminodes usually prismatic, truncate, sometimes clavate, often somewhat similar to stamens. FEMALE FLOWER: gynoecium ovoid, subcylindric, cylindric or obovoid, ovary (2–)4–8(–47)-locular, ovules 1–50 or more per locule, usually hemiorthotropous, rarely hemianatropous to nearly anatropous, funicle long to very short, placenta axile to basal, stylar region usually as broad as ovary, sometimes slightly broader, sometimes attenuate, rarely elongate, lobed in subgen. Meconostigma, stigma sometimes also lobed or discoid-hemispheric, often as broad as style. BERRY: subcylindric to obovoid, 1–many-seeded, white, whitishtranslucent, red or orange-red. SEED: tiny to fairly large, ovoid-oblong to ellipsoid, rarely arillate (in P. goeldii funicle thick, swollen, much larger than seed itself), testa thick, costate, rarely sarcotestate, embryo axile, straight, elongate, endosperm copious. See Plates 45i–v, 116C. CHROMOSOMES: 2n = 28, 30, 32, 34, 36 (26, 48). DISTRIBUTION: over 500 spp.; tropical and southern subtropical America, West Indies:– Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad & Tobago, Uruguay, Venezuela. ECOLOGY: usually tropical humid forest, more rarely in open woodland, swamps, streamsides; climbing hemiepiphytes, rosulate acaulescent epiphytes, rhizomatous terrestrials, lithophytes (also on cliffs), helophytes, mostly shade-loving, sometimes arborescent. NOTES: Mayo (1986b, 1989b, 1990a, 1991) recognized 3 subgenera: subgen. Philodendron, subgen. Pteromischum (partially revised by Grayum 1996), subgen. Meconostigma (revised by Mayo 1991). ETYMOLOGY: Greek philos (fond of) and dendron (tree); refers to the predominantly epiphytic or hemiepiphytic habit. TAXONOMIC ACCOUNTS: Krause (1913), Bunting (1968, 1977, 1980, 1984), Mayo (1986b, 1989a, b, 1990a, 1991), Grayum (1992b, 1996), Croat & Grayum (1994), Sakuragui (1994), Nadruz Coelho (1995).

Tribe Homalomeneae

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Tribe Homalomeneae (Schott) M. Hotta in Mem. Fac. Sci. Kyoto Univ., ser. Biol. 4: 89 (1970). Laticifers present, simple, articulated, resin canals present in roots, stems and leaves, sclerotic hypodermis present in roots; terrestrial or rheophytic, diminutive to robust, stem usually epigeal; petiole geniculum absent (very rarely present in Homalomena); primary lateral veins pinnate forming 1 marginal vein, higher order venation parallel-pinnate; spathe often boat-shaped, constricted or unconstricted, persistent, closing after anthesis; flowers unisexual, perigone absent; anther with cell wall thickenings in endothecium; endosperm copious.

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46. Furtadoa Furtadoa M. Hotta in Acta Phytotax. Geobot. 32: 142 (1981). TYPE: F. sumatrensis M. Hotta HABIT: small evergreen herbs, stem repent. LEAVES: several to many. PETIOLE: sheath to half as long as petiole or more. BLADE: elliptic; primary lateral veins pinnate, running into margin, higher order venation parallel-pinnate. INFLORESCENCE: 1–3 in each floral sympodium. PEDUNCLE: shorter than or subequal to petiole. SPATHE: green, ellipsoid, not constricted, boat-shaped, persistent. SPADIX: subcylindric, tapering apically, female zone a third to nearly half spadix length, male zone contiguous with female. FLOWERS: unisexual, perigone absent; each male and female flower with a stamen or staminode situated basally to pistillode or gynoecium respectively. MALE FLOWER: consisting of a single free stamen overtopped by single flask-shaped pistillode with subglobose stigmatoid apex, pistillodes absent from apical flowers, stamen apex truncate, connective thick, thecae ovoid, dehiscing by short, longitudinal slit. POLLEN: inaperturate, ellipsoid-oblong, small (mean 17 µm.), exine virtually psilate. FEMALE FLOWER: consisting of gynoecium with single, shorter, obovoid, apically truncate staminode, lowermost flowers sometimes apparently with more than one staminode, gynoecium ovoid, ovary 1-

46. Furtadoa

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Plate 46. Furtadoa. A, habit × 2/3; B, spadix × 3; C, stamens with associated pistillodes, × 15; D, stamen and associated pistillode, top view × 15; E, gynoecia with associated staminodes, × 15; F, gynoecium, longitudinal section × 15; G, habit × 2/3; H, detail of leaf tip tubule × 8; J, spadix × 3; K, stamen with associated pistillode, gynoecium in longitudinal section × 15; L, stamen and associated pistillode, top view × 15; M, gynoecia with associated staminodes, upper gynoecium in longitudinal section × 15; N, gynoecium with associated staminode, top view × 15. Furtadoa mixta: A, Nur 11091 (K); B–F, Hay 9131 (Kew spirit collection 58934); F. sumatrensis: G–H, Hotta s.n. (K); J, Bogner s.n. (Kew spirit collection 56675); K–N, Nerz s.n. (Kew spirit collection 56142).

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locular, ovules many, hemianatropous, placenta basal to intrusive basal, stylar region short and attenuate to inconspicuous, stigma discoid-subcapitate. BERRY: subcylindric, light green, bearing old stigma remains. SEED: ellipsoid to ovoid, testa smooth, thin, cream-coloured to very light green, embryo straight, endosperm copious. See Plates 46, 116D. CHROMOSOMES: 2n = 40. DISTRIBUTION: 2 spp.; Malay Archipelago:– Indonesia (Sumatra), Malaysia (Peninsula). ECOLOGY: tropical humid forest; rheophytes on rocks in streams (F. sumatrensis), or forest floor terrestrials (F. mixta). ETYMOLOGY: named after C.X. Furtado (1897–1980). TAXONOMIC ACCOUNTS: Hotta (1981, 1985).

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47. Homalomena Homalomena Schott in Schott & Endlicher, Melet. Bot. 20 (1832). LECTOTYPE: H. cordata Schott (Dracontium cordatum Houttuyn 1779, non Aublet 1775; see Nicolson in Taxon 16: 517. 1967). SYNONYMS: Homalonema Endlicher, Gen. Pl. 238 (1837), orth. var.; Spirospatha Rafinesque, Fl. Tell. 4: 8 (1838, “1836”); Cyrtocladon Griffith, Notul. Pl. Asiat. (Posthum. Pap.) 3: 147 (1851); Chamaecladon Miq. in Bot. Zeitung (Berlin) 14: 564 (1856); Adelonema Schott, Prodr. syst. Aroid. 316 (1860); Curmeria E.F. André, Ill. Hortic. 20: 45 (1873); Diandriella Engler, Nova Guinea 8: 250 (1910). HABIT: evergreen, usually aromatic (anise-scented) herbs, stem shortly aerial, more rarely arborescent or hypogeal. LEAVES: several, rarely distichous (H. geniculata). PETIOLE: rarely aculeate or pubescent, rarely geniculate apically (H. geniculata), sheath usually less than half as long as petiole. BLADE: lanceolate, elliptic, oblong, subtriangular or cordate to sagittate, rarely peltate, usually glabrous, rarely pubescent on midrib and veins; primary lateral veins pinnate, running into marginal vein, secondary and tertiary lateral veins parallelpinnate. INFLORESCENCE: 1–6 (or more) in each floral sympodium. PEDUNCLE: shorter than petiole. SPATHE: erect,

often becoming green, more rarely white or yellow-green or red, persistent, usually not constricted, ellipsoid to boatshaped, more rarely constricted between tube and blade and then tube convolute, blade gaping at anthesis and afterwards closing. SPADIX: shorter or subequal to spathe, stipitate or sessile, female zone cylindric, male zone usually entirely fertile, contiguous with and longer than female zone, rarely bearing staminodes basally, or very rarely separated by a ± naked interstice. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4-androus, rarely 5–6-androus, very rarely 1-androus (H. monandra), stamens free, truncate apically, filaments absent or distinct, connective thick, thecae ovoid, ellipsoid or oblong, opening by longitudinal slit, rarely by transversal slit. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, small (mean 22 µm., range 12–31 µm.), exine perfectly psilate in most species, rarely obscurely fossulate. STERILE MALE FLOWERS: 2–4-androus, sometimes present at base of fertile male zone, staminodia subprismatic, somewhat rounded apically. FEMALE FLOWER: gynoecium ovoid or oblong or subglobose, usually with single, anterior staminode (rarely 2, very rarely 3), equalling or half as long as ovary, sometimes absent (H. lindenii), ovary incompletely 2–4–(–5)-locular, ovules many, hemianatropous, funicle long, placenta parietal and axile, stylar region shortly narrowed or inconspicuous, stigma discoid, subhemispheric, subcapitate or slightly 2–4-lobed. BERRY: obovoid or subglobose or cylindric, locules many-seeded, rarely few-seeded. SEED: ellipsoid or elongate ellipsoid, testa thick, distinctly or only slightly costate, embryo axile, elongate, endosperm copious. See Plates 47i–ii, 117A. CHROMOSOMES: 2n = 38, 40, 42, 80. DISTRIBUTION: ca. 110 spp.; tropical southeast Asia, Malay Archipelago, tropical America:– Bangladesh, Bolivia, Brazil (Amazonia, Central-West), Brunei, Burma, Cambodia, China (Guandong, Guangxi, Hainan, Taiwan, Yunnan), Colombia, Costa Rica, Ecuador, French Guiana, Guyana, India (Assam), Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra, Sunda Is.), Laos, Malaysia (Borneo, Peninsula), Panama, Papua New Guinea, Peru, Philippines, Singapore, Solomon Is., Surinam, Thailand, ?Venezuela, Vietnam.

47. Homalomena

HOMALOMENEAE : HOMALOMENA

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Plate 47 (i). Homalomena. A, habit × 1/5; B, leaf × 1/2; C; leaf × 1/2; D, base of plant showing inflorescences × 1/2; E, leaf × 1/2; F, base of plant showing inflorescences × 1/2; G, leaf × 1/2; H, leaf × 1/2. Homalomena sagittifolia: A, Boyce 252 (Kew slide collecttion); H. picturata: B, de Granville 5467 (K); H. rubescens: C–D, Keenan s.n. (K); H. havilandii: E, Ilias & Azahari S 35677 (K); F, Hetterscheid s.n. (Kew slide collection); H. consobrina: G, de Wilde & de Wilde-Duyfjes 13495 (K); H. propinqua: H, Haviland 3134 (K).

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Plate 47 (ii). Homalomena. A, infructescence, lower part of spathe removed to reveal berries × 1/1/2; B, inflorescence × 4; C, stamen × 15; D, gynoecium and associated staminode × 15; E, gynoecium, longitudinal section × 15; F, gynoecium, transverse section × 24; G, inflorescence, spathe partly removed × 2/3; H, stamen × 10; J, gynoecium × 10; K, gynoecium, longitudinal section × 10; L, gynoecium, transverse section × 16; M, inflorescence × 1; N, stamen × 10; P, gynoecium and associated staminode × 10; Q, gynoecium, longitudinal section × 10; R, gynoecium, transverse section × 16; S, inflorescence × 1; T, stamen × 10; U, gynoecium × 10; V, gynoecium, longitudinal section × 10; W, gynoecium, transverse section × 16. Homalomena vagans: A, Poulsen & de la Motte 273 (Kew spirit collection 58113); H. humilis: B–F, Sands 180 (Kew spirit collection 32886); H. speariae: G–L, Spear s.n. (Kew spirit collection 58911); H. rubra: M–R, Burkhill & Haniff 12792 (K & Kew spirit collection 59026); H. hostifolia: S–W, Poulsen 268 (Kew spirit collection 59027).

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ECOLOGY: tropical humid forest, swamp forest, rarely in open swamps; terrestrial in leaf litter on forest floor, along forest streams, on road banks, on well-drained slopes in moist forest, rarely rheophytes. NOTES: Engler (1912) recognized 3 sections:– sect. Homalomena, sect. Chamaecladon, sect. Curmeria (tropical America); Furtado (1939) recognized a fourth section, sect. Cyrtocladon and Hotta (1967) a fifth, sect. Geniculatae. ETYMOLOGY: Greek homalos (flat) and mênê (moon), translation of a vernacular name. TAXONOMIC ACCOUNTS: Engler (1912), Furtado (1939), Hotta (1967, 1982, 1984, 1985, 1986a, 1993), Bogner (1976b, 1986a), Bogner & Moffler (1984, 1985).

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Tribe Anubiadeae Tribe Anubiadeae Engler in Nova Acta Acad. Leopold.Carol. 39: 147 (1876). Laticifers present, simple, articulated, roots with sclerotic hypodermis; helophytic or rheophytic, stem rhizomatous, creeping; petiole geniculate apically; leaf blade elliptic, lanceolate to hastate-tripartite, primary lateral veins pinnate, forming single marginal vein, higher order venation parallelpinnate, finest transverse veins ± distinct; spathe boat-shaped, unconstricted, persistent; flowers unisexual, perigone absent; stamens connate into ± prismatic synandria, fused connectives thickened, ± truncate, thecae lateral or marginal, rarely covering whole synandrium, dehiscing by longitudinal slit; ovary (1–)2–3-locular, ovules many per locule, anatropous, placenta axile, style narrower than ovary, stigma broad; berry depressed-globose to obovoid; seeds small, ± ovoid to subcylindric, endosperm copious.

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48. Anubias Anubias Schott in Oesterr. bot. Wochenbl. 7: 398 (1857). TYPE: A. afzelii Schott SYNONYM: Amauriella Rendle, Cat. Talbot’s Nigerian Pl. 115 (1913). HABIT: evergreen herbs, rhizome thick, creeping, internodes short. LEAVES: several. PETIOLE: usually smooth, rarely shortly and sparsely spiny, geniculate apically, sheath relatively short. BLADE: lanceolate, ovate, elliptic, nearly triangular to subsagittate, subcordate, auriculate, hastate to trifid, mostly coriaceous, completely glabrous or midrib and primary lateral veins densely pilose abaxially; primary lateral veins pinnate, secondary lateral veins parallel-pinnate, tertiaries transverse between them. INFLORESCENCE: 1–3 in each floral sympodium. PEDUNCLE: ± as long as petiole or shorter. SPATHE: elliptic-ovate or ovate, not constricted, only weakly differentiated into tube and blade, widely expanded at anthesis, but always slightly convolute at base, closing after anthesis and persistent to fruiting stage, uniformly coloured, mostly green or cream to reddish tinged, paler within. SPADIX: cylindric, shorter or sometimes much longer than spathe, sessile or stipitate, female zone free, usually densely flowered, rarely somewhat laxly so, shorter than male and contiguous with it or rarely with few sterile or irregular, bisexual flowers in between, male zone fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–8-androus, stamens connate, synandrium ±

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48. Anubias

obpyramidal, filaments connate, sometimes fairly long, fused connectives thick, fleshy, sometimes covered by thecae and inconspicuous, often only incompletely connate at apex, with shallow fissures in between, thecae lateral or marginal or covering nearly the whole synandrium from apex to base (A. pynaertii), dehiscing by longitudinal slit. POLLEN: inaperturate, subspheroidal to spherical, small (mean 24 µm., range 20–31 µm.), exine perfectly psilate to obscurely verrucate and/or dimpled. FEMALE FLOWER: gynoecium depressedglobose to ovoid, ovary (1–)2–3-locular, ovules many per locule, anatropous, placenta axile, stylar region narrower than ovary, stigma broad, discoid, green, pink or white. BERRY: depressed-globular to obovoid, green to pale green, many-seeded. SEED: small, irregularly ovoid to subcylindric, testa rough, thickish, embryo axile, elongate, endosperm copious. See Plates 48, 117B. CHROMOSOMES: 2n = 48, 72. DISTRIBUTION: 8 spp.; tropical west and central Africa:– Angola, ?Benin, Cabinda, Cameroon, ?Central African Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, ?Gambia, Ghana, Guinea, ?Guinea-Bissau, Ivory Coast, Liberia, Mali, Nigeria, Senegal, Sierra Leone, Togo, Zaïre. ECOLOGY: tropical humid forest; helophytes or rheophytes, forest swamps, in rocky places along streams, sometimes completely submerged. ETYMOLOGY: Classical name of an unknown herb, anoubias (according to Schott). TAXONOMIC ACCOUNTS: Engler (1915), Crusio (1979, 1987), Kasselmann (1995).

Tribe Schismatoglottideae Tribe Schismatoglottideae Nakai, Ord. Fam. Trib. Nov. 218 (1943); Hotta in Acta Phytotax. Geobot. 33: 127–139 (1982). Laticifers present, simple, articulated; terrestrial or rheophytic, stem usually epigeal; petiole sheath usually with long, marcescent, apical ligule (except most Schismatoglottis spp.); blade apex with tubular mucro (except most Schismatoglottis spp.), primary lateral veins pinnate, forming single, usually prominent marginal vein, higher order venation parallel-pin-

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Plate 48. Anubias. A, habit × 1/2; B, inflorescence × 1; C, detail of male zone of spadix × 5; D, detail of female zone of spadix × 5; E, synandrium, side view × 10; F, synandrium, top view × 10; G, gynoecium, longitudinal section × 10; H, gynoecium, transverse section × 10; J, habit × 1/2; K, habit × 1/2; L, synandrium, side view × 10; M, synandrium, top view × 10; N, gynoecium, longitudinal section × 10; P, synandrium, side view × 10; Q, synandrium, top view × 10; R, gynoecium, longitudinal section × 10; S, inflorescence × 1; T, synandrium, side view × 10; U, synandrium, top view × 10; V, gynoecium, longitudinal section × 10; W, leaf × 1/2. Anubias afzelii: A, Morton & Gledhill SL 1169 (K); B, Hepper 2504 (K); C–H, Cult. Kew 1963–02202 (Kew spirit collection 50101); A. gracilis: J, Morton & Gledhill SL 1929 (K); A. barteri var. barteri: K, Onochie & Okafor IFH 36037 (K); L–N, Brenan 9257 (Kew spirit collection 25469); A. hastifolia: P–R, Bogner s.n. (Kew spirit collection 7100); A. pynaertii: S, W. Crusio, Die Gattung Anubias Schott, p. 38 (1987); T–V, Bogner 699 (K & Kew spirit collection 56680); A. gigantea: W, Bos 1914 (K).

ANUBIADEAE : ANUBIAS

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nate; inflorescence usually 1 (except Schismatoglottis); peduncle usually elongating in fruit; spathe tube persistent, blade usually white (except Piptospatha), deciduous at anthesis, rarely marcescent (in Schismatoglottis beccariana, Hottarum lucens); flowers unisexual, perigone absent; stamens usually free, sometimes connate by fused filaments, filaments often ± elongated, thecae truncate or horned, nearly always dehiscing by apical pore; ovary 1-locular, ovules orthotropous to hemiorthotropous (except Schismatoglottis), stigma usually sessile (except Schismatoglottis); seed testa usually costate, embryo axile, endosperm copious.

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49. Schismatoglottis Schismatoglottis Zollinger & Moritzi in Moritzi, Syst. Verzeichnis Zollinger, 83 (1846). TYPE: S. calyptrata (Roxburgh) Zollinger & Moritzi (Calla calyptrata Roxburgh). SYNONYMS: Philonotion Schott in Oesterr. bot. Wochenbl. 7: 421 (1857); Apoballis Schott in Oesterr. bot. Zeitschr. 8: 318 (1858); Apatemone Schott, Gen. Aroid. t. 57 (1858); Colobogynium Schott in Oesterr. bot. Zeitschr. 15: 34 (1865); Nebrownia O. Kuntze, Rev. Gen. 2: 742 (1891). HABIT: small to large, evergreen herbs, rarely shortly, densely or long-pubescent, stem rhizomatous or epigeal, shortly erect. LEAVES: numerous, rarely distichous. PETIOLE: sheath less than half petiole length, sometimes with long apical ligule. BLADE: narrow-elliptic, elliptic, lanceolate, oblanceolate, ovate, obovate, cordate, cordate-sagittate, sometimes variegated with paler or silvery green, white or yellow; primary lateral veins pinnate, running into distinct marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation transverse-reticulate. INFLORESCENCE: 1–3 (or more) in each floral sympodium. PEDUNCLE: shorter than petiole. SPATHE: constricted between tube and blade, sometimes only slightly so, rarely not at all, tube convolute, persistent, blade thinner, erect, broadly boat-shaped, gaping and then caducous at anthesis, rarely marcescent, usually white to cream, sometimes greenish-yellow, very rarely pink, cuspidate to acuminate. SPADIX: shorter than or equalling spathe, lower part consisting of cylindric to conoid female zone, free or partially adnate to spathe, sometimes bearing sterile organs at the very base, upper part of spadix usually ± clavate, sometimes subcylindric, either fertile male below

and sterile terminally, or sterile below and fertile male terminally, or sterile below, centrally fertile male and sterile terminally, lowermost sterile zone sometimes very laxly flowered, often constricted. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–3-androus, stamens very short to long, mostly free, filaments usually well-developed, sometimes connate basally, always with distinctive tannin cells (dark in dried specimens), connective usually rather slender, sometimes thicker apically, anthers truncate, often concave apically, thecae opposite, cylindric or obconic to ovoid, dehiscing by apical, broadly elliptic, or bilobed pore. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, small (mean 20 µm., range 15–26 µm.), exine perfectly psilate, rarely rugulate to verruculate (S. spruceana). STERILE MALE FLOWERS: staminodes less compressed than stamens, obpyramidal to clavate, usually truncate, short to long. FEMALE FLOWER: gynoecium sometimes accompanied by 1–4 clavate, rarely peltate staminodes with generally distinctly swollen apices, or rarely with sterile flowers scattered among gynoecia, ovary 1-locular, ovules (1–)few to many, anatropous to hemianatropous, funicle rather long, placentae 1–4, parietal, usually extending from base to apex of locule, stylar region inconspicuous or shortly conoid, stigma discoid to capitate, small to as wide as ovary. BERRY: oblong to globose, green or dull yellow or deep red, few- to many-seeded. SEED: ellipsoid, testa costate, embryo straight, elongate, endosperm copious. See Plates 49i–iii, 117C. CHROMOSOMES: 2n = 26, 39, 52. DISTRIBUTION: ca. 120 spp.; tropical Asia, Malay Archipelago, tropical South America:– Brazil (Amazonia), Brunei, Burma, Cambodia, China (Guandong, Guangxi, Hainan, Yunnan), Colombia, French Guiana, ?Guyana, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra, Sunda Is.), Laos, Malaysia (Borneo, Peninsula), Papua New Guinea, Peru, Philippines, Solomon Is., Surinam, Thailand, Vanuatu, Venezuela, Vietnam. ECOLOGY: tropical humid forest; terrestrial, forest floor, sometimes rheophytes. NOTES: Hotta (1966a) recognized 5 informal groups in the Bornean species:– S. homalomenoidea group, S. monoplacenta group, S. barbata group, S. acutifolia group, S. calyptrata group; the neotropical species earlier recognized as the genus Philonotion now form sect. Philonotion (Bunting 1960b).

49. Schismatoglottis

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Plate 49 (i). Schismatoglottis. A, habit × 1/2; B, detail of leaf tip tubule × 8; C, habit × 1/2; D, habit × 1/2; E, habit × 1/2. Schismatoglottis gillianae: A–B, Coode 6313 (K); S. spruceana: C, Plowman 13527A (K); S. hottae: D, Johns 6872 (K); S. convolvula: E, Mamit S42102 (K).

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Plate 49 (ii). Schismatoglottis. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2. Schismatoglottis hastifolia: A, Clemens & Clemens 29493 (K); S. calyptrata: B, Beaman 10632 (K); S. ferruginea: C, Dransfield 6871 (K); D, S. hottae: Johns 6872 (K); S. crispata: E, Cult. Veitch, June & July 1881 (K); S. spruceana: F, Plowman 13527A (K); S. gillianae: G, Coode 6313 (K).

ETYMOLOGY: Greek schisma, schismatos (separating) and glôtta (tongue); refers to the deciduous spathe blade. TAXONOMIC ACCOUNTS: Engler (1912), Bunting (1960b), Hotta (1965, 1966a), Bunting & Steyermark (1969), Bogner & Hotta (1983b), Bogner (1988a).

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50. Piptospatha Piptospatha N.E. Brown in Gard. Chron., ser. 2, 11: 138 (1879). TYPE: P. insignis N.E. Brown SYNONYMS: Rhynchopyle Engler in Bot. Jahrb. 1: 183 (1880) (“1881”); Gamogyne N.E. Brown in J. Bot. 20: 195 (1882). HABIT: small to medium-sized evergreen herbs, stem erect or decumbent. LEAVES: several. PETIOLE: sheath short with long, marcescent ligule. BLADE: elongate-lanceolate to elliptic or oblanceolate, coriaceous, apex with tubular mucro; primary lateral veins pinnate, running into distinct marginal vein, secondary laterals and higher order venation parallelpinnate. INFLORESCENCE: solitary, usually nodding. PEDUNCLE: subequal to or longer than petiole. SPATHE: stoutly ellipsoid, not constricted, often pink, lower part persistent and cup-like, upper part slightly gaping at anthesis, caducous or deliquescent, cuspidate to acuminate. SPADIX: sessile with ± oblique insertion, sometimes with sterile female flowers at extreme base, female zone cylindric, shorter and contiguous with male, or separated by a short

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zone of sterile male flowers, male zone cylindric, equal in thickness to female, obtuse, fertile to apex or with a few sterile terminal flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–2-androus, stamens free, compressed, anthers truncate, connective ± flat or expanded apically or with conical beak (P. insignis), overtopping thecae, thecae oblong-ellipsoid, dehiscing by apical pore. POLLEN: inaperturate, ellipsoid, small to medium-sized (mean 25 µm.), exine psilate. STERILE MALE FLOWERS: apparently composed of a single truncate, subclavate, prismatic staminode. FEMALE FLOWER: gynoecium free or cohering to neighbouring ones, ovary 1-locular, ovules many, hemiorthotropous to almost orthotropous and erect, funicle long, placentae 2–4, parietal or parietal and basal, stigma ± sessile or stylar region inconspicuous, as broad as ovary, contiguous with adjacent ones. BERRY: obovoid, green. SEED: elongate-ellipsoid to cylindric, with long, curved micropylar appendage, testa slightly costate, embryo elongate, endosperm copious. See Plates 50, 117D. CHROMOSOMES: 2n = 26. DISTRIBUTION: 10 spp.; Brunei, Indonesia (Borneo), Malaysia (Borneo, Peninsula), Thailand. ECOLOGY: tropical humid forest; rheophytes. NOTES: sect. Piptospatha, sect. Gamogyne (N.E. Brown) M. Hotta. The latter section is only weakly defined, by the presence of superficially connate gynoecia. ETYMOLOGY: Greek piptô (I fall) and spathê (spathe); refers to the deciduous spathe blade. TAXONOMIC ACCOUNTS: Engler (1912), Hotta (1965).

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Plate 49 (iii). Schismatoglottis. A, infructescences with associated petiole × 2/3; B, inflorescence × 1; C, spadix × 2; D, staminode × 20; E, stamens × 20; F, gynoecium × 20; G, gynoecium, longitudinal section × 20; H, gynoecium, transverse section × 30; J, spadix × 2; K, staminode × 20; L, stamen × 20; M, gynoecium × 20; N, gynoecium, longitudinal section × 20; P, gynoecium, transverse section × 30; Q, inflorescence × 1; R, spadix × 2; S, staminode × 20; T, stamen × 20; U, gynoecium × 20; V, gynoecium, longitudinal section × 20; W, gynoecium, transverse section × 30; X, spadix, spathe partly removed × 2; Y, stamen × 20; Z, gynoecium × 20; AA, gynoecium, longitudinal section × 20; BB, gynoecium, transverse section × 30. Schismatoglottis neoguinensis: A, Floyd 6445 (K); S. tecturata: B–H, Bogner 1554 (Kew spirit collection 45228); S. spruceana: J–P, Plowman 13527A (K & Kew spirit collection 58039); S. crispata: Q–W, Boyce 672 (Kew spirit collection 59035); S. calyptrata: X–BB, Hay 2007, Cult. Kew 1982–04973 (Kew spirit collection 46560).

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B F

D

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Plate 50. Piptospatha. A, habit × 1/2; B, seed × 15; C, habit × 1/2; D, spadix × 2; E, stamen × 15; F, gynoecia, right hand gynoecium longitudinally sectioned × 10; G, habit × 1/2; H, detail of leaf tip tubule × 6; J, spadix × 2; K, stamen × 15; L, gynoecium × 15; M, gynoecium, longitudinal section × 15; N, staminode × 15. Piptospatha elongata: A, Chew, Corner & Stainton 2501 (K); B, Bogner 2153 (Kew spirit collection 59089); P. burbidgei: C–F, Richards 1091 (K & Kew spirit collection 58020); P. ridleyi: G, Burkill 2577 (K); H, Sinclair 10577 (K); J–M, Bogner 2120 (Kew spirit collection 57278).

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50. Piptospatha

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51. Hottarum Hottarum Bogner & Nicolson in Aroideana 1: 72 (1979, “1978”). TYPE: H. truncatum (M. Hotta) Bogner & Nicolson (Microcasia truncata M.Hotta). SYNONYM: Based on Microcasia sect. Truncatae M. Hotta in Mem. Coll. Sci. Univ. Kyoto, ser. B, Biol., 32: 21 (1965). HABIT: evergreen herbs, stem short, ± erect, epigeal, internodes short. LEAVES: numerous. PETIOLE: sheath with long, free ligule. BLADE: elliptic, oblong-elliptic, or narrowly elliptic, coriaceous, apex with tubular mucro; primary lateral veins pinnate, running into distinct marginal vein, higher order venation parallel-pinnate. INFLORESCENCE: 1, rarely 2 in each floral sympodium. PEDUNCLE: subequal to petiole in flower, elongating in fruit. SPATHE: constricted or not, uniformly coloured, or green below and white above, ellipsoid to obovoid at anthesis, lower part convolute, cup-shaped and persistent after anthesis, upper part only opening slightly at anthesis, caducous, or marcescent and then evanescent (H. lucens), apex acuminate to cuspidate. SPADIX: subcylindric, usually free or sometimes adnate to spathe for two-thirds of length (H. lucens), female zone subcylindric, sometimes with sterile flowers at base, either contiguous with male, or with a few sterile male flowers in between, male zone as thick or thicker than female, apical zone composed of sterile male flowers, in H. lucens fertile only where exposed at anthesis by spathe opening and otherwise com-

posed entirely of sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: (1–)2–3(–4)-androus, stamens truncate at apex, filaments distinct, sometimes connate basally, connective broad, thick, thecae lateral, opposite, not horned, dehiscing apically by single pore or two pores confluent into a short transverse slit. POLLEN: inaperturate, ellipsoid or globose, small (mean 14 µm., range 10–17 µm.), exine perfectly psilate or irregularly dimpled. STERILE MALE FLOWERS: composed of truncate, prismatic staminodes. FEMALE FLOWER: usually consisting of gynoecium alone, sometimes also with a clavate staminode in basal flowers (or lateral flowers when female zone adnate to spathe), ovary depressed-globose, 1-locular, ovules 10–15, orthotropous, funicle long, placenta basal, stigma sessile, discoid or subcapitate. BERRY: ± globose, many-seeded, whitish green with brownish stigma remnants. SEED: ellipsoid to elongate, testa whitish to brownish, costate, embryo straight, elongate, endosperm copious. See Plates 51, 118A. CHROMOSOMES: 2n = 26. DISTRIBUTION: 6 spp.; Brunei, Indonesia (Borneo), Malaysia (Borneo). ECOLOGY: tropical humid forest; rheophytes. NOTES: Hottarum lucens is rather different from the other species. ETYMOLOGY: named after Mitsuru Hotta (born 1935) and Arum. TAXONOMIC ACCOUNTS: Bogner (1983b,1984b), Bogner & Hotta (1983a).

51. Hottarum

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Plate 51. Hottarum. A, habit × 2/3; B, spadix × 2; C, stamen × 16; D, staminode × 16; E, gynoecium, longitudinal section × 16; F, spadix × 2; G, stamen × 16; H, staminode × 16; J, gynoecium with associated staminode; K, habit × 2/3; L, detail of leaf tip tubule × 6; M, spadix × 3; N, stamen × 16; P, gynoecium × 16; Q, gynoecium, longitudinal section × 16. Hottarum lucens: A, Elsener 184 (K); B–E, Bogner 1439 (Kew spirit collection 45223); H. sarikeense: F–J, Bogner 1553 (Kew spirit collection 45681); H. kinabaluense : K, M–Q, Edwards 2162 (Kew spirit collection 54425); L, Clemens & Clemens 29135 (K).

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52. Bucephalandra Bucephalandra Schott, Gen. Aroid. t. 56 (1858). TYPE: B. motleyana Schott SYNONYM: Microcasia Beccari in Bull. Soc. Tosc. Ortic. 4: 180 (1879). HABIT: minute to medium-sized evergreen herbs, stem creeping, apex upright. LEAVES: numerous. PETIOLE: sheath with long marcescent ligule. BLADE: elliptic, elliptic-oblong, linearoblanceolate to obovate, coriaceous, punctate below, apex with tubular mucro; primary lateral veins pinnate, running into distinct marginal vein, higher order venation parallel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole at anthesis, elongating later. SPATHE: ellipsoid, cuspidate, not constricted, lower part light green, convolute, broadly funnelshaped, persistent, enclosing developing fruits, upper part white, gaping at anthesis, caducous immediately afterwards. SPADIX: sessile, shorter than spathe, with a few pistillodes at extreme base, female zone cylindric, narrower than upper parts, with gynoecia in 2–6 spirals, separated from male zone by a few rows (usually 2) of flattened, smooth, scale-like staminodes, male zone with 2–5 rows of flowers, terminal appendix globose or ellipsoid to subcylindric, composed of truncate, obpyramidal to subcylindric, apically papillose staminodes, the uppermost ± connate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, filament distinct but short, flattened, connective ± inconspicuous, thecae ellipsoid, extrorse, dehiscing by pore at tip of conspicuous apical horn. POLLEN: extruded in a droplet, inaperturate, ellipsoid, medium-sized (mean 29 µm., range 28–30 × 20–24 µm.), exine smooth (psilate). FEMALE FLOWER: gynoecium depressedglobose, 1-locular, ovules many, orthotropous, attenuate towards micropyle, funicle distinct, placenta basal, stigma sessile, discoid, slightly concave in centre, narrower than ovary. BERRY: globose to ellipsoid with numerous seeds. SEED: narrow-ellipsoid, with long, curved micropylar appendage, testa very slightly longitudinally ribbed to scabrous, embryo straight, elongate, endosperm copious. See Plates 52, 118B. CHROMOSOMES: 2n = ca. 26. DISTRIBUTION: 3 spp.; Brunei, Indonesia (Borneo), Malaysia (Borneo). ECOLOGY: tropical humid forest; rheophytes. ETYMOLOGY: Greek bous (bull or cow), cephalê (head) and anêr, andros (man). TAXONOMIC ACCOUNTS: Engler (1912), Bogner (1980a, 1984b), Boyce (1995a), Boyce, Bogner & Mayo (1995).

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53. Phymatarum Phymatarum M. Hotta in Mem. Coll. Sci. Kyoto Imp. Univ., ser. B, 32: 29 (1965). TYPE: P. borneense M. Hotta HABIT: small evergreen herbs, stem creeping to decumbent. LEAVES: several. PETIOLE: sheath fairly short with long marcescent ligule. BLADE: narrowly elliptic, somewhat coriaceous, apex with tubular mucro; primary lateral veins pinnate, running into conspicuous marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation inconspicuously transverse-reticulate. INFLORESCENCE: solitary. PEDUNCLE: erect, shorter or subequal to petiole. SPATHE: constricted between tube and blade, tube convolute, persistent, green, blade longer, boat-shaped and gaping at anthesis, whitish, cuspidate, caducous after anthesis. SPADIX: extreme base bearing a few pistillodes or not, female zone conoid to subcylindric, basally adnate to spathe, separated from male zone by cylindric to ellipsoid zone of sterile male flowers, male zone very short and slightly narrower, terminal appendix much longer, elongate-conoid, bearing sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: apparently 1androus, free, filament short, connective inconspicuous, thecae tuberculate, ending in curved horn, dehiscing by apical pore. STERILE MALE FLOWERS: staminodes subprismatic, tuberculate, flattened or excavated, lowermost either with or without central, short, subulate projection, uppermost more slender, relatively longer, otherwise similar but never with projections. POLLEN: inaperturate, ellipsoid, small (mean 19 µm., range 17–22 µm.) exine psilate. FEMALE FLOWER: gynoecium depressed- globose, ovary 1-locular, ovules many, hemiorthotropous, funicle long, placenta basal, stigma sessile, slightly concave centrally, narrower than ovary, very thinly discoid. BERRY: many-seeded, depressed-obovoid, slightly furrowed, greenish-white. SEED: ellipsoid, with long micropylar appendage, testa costate, embryo elongate, straight, endosperm copious. See Plates 53, 118C. CHROMOSOMES: 2n = 26. DISTRIBUTION: 3 spp.; Brunei, Indonesia (Borneo), Malaysia (Borneo). ECOLOGY: tropical humid forest; rheophytes. ETYMOLOGY: Greek phyma, phymatos (tumour, growth) and Arum. TAXONOMIC ACCOUNTS: Bogner (1984a).

52. Bucephalandra

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Plate 52. Bucephalandra. A, habit × 2/3; B, habit × 2/3; C, detail of leaf tip tubule × 6; D, inflorescence × 1; E, spadix × 5; F, upper spadix staminode × 15; G, stamen × 15; H, mid–spadix staminode × 15; J, gynoecium, longitudinal section × 15; K, lower spadix pistillode × 15; L, base of plant × 2/3; M, leaf × 2/3; N, spadix × 5; P, habit × 2/3; Q, spadix × 5. Bucephalandra motleyana: A, Mamit S 33470 (K); B, Chew 1111 (K); C, Marsh & Simpson 21 (Kew spirit collection 57279); D–K, Bogner 1447 (Kew spirit collection 45261); B. gigantea: L–M, Endert 4580 (K); N, Endert 4580 (Kew spirit collection 58050); B. catherineae: P–Q, de Vogel & Cribb 9210 (Kew spirit collection 57575).

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A

B

E K

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Plate 53. Phymatarum. A, habit × 2/3; B, detail of leaf tip tubule × 6; C, spadix × 2; D, staminode from appendix × 8; E, stamen × 8; F, staminode from lower zone of sterile male flowers × 8; G, gynoecium–associated staminode × 8; H, gynoecium × 8; J, gynoecium, longitudinal section × 8; K, seed × 4. Phymatarum borneense: A–B, Bogner 1506 (K); C–J, Boyce 341 (Kew spirit collection 56097); K, Bogner 2159 (Kew spirit collection 58007).

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54. Aridarum Aridarum Ridley in J. Bot. 51: 201 (1913). TYPE: A. montanum Ridley HABIT: small to medium-sized, evergreen herbs, stem decumbent, erect distal part sometimes rather long. LEAVES: several, spiral or rarely distichous (A. annae). PETIOLE: sheath with long, marcescent ligule. BLADE: coriaceous, linear to elliptic, apex with tubular mucro; primary lateral veins pinnate, weakly or not differentiated, running into distinct marginal vein, higher order venation parallel-pinnate. INFLORESCENCE: solitary, sometimes ± nodding. PEDUNCLE: subequal or longer than petiole, sometimes ± deflexed at apex. SPATHE: stoutly ellipsoid, not constricted, convolute and gaping at apex only or broadly boat-shaped and widely gaping to base, lower part persistent, green, upper part caducous, white, cuspidate to acuminate at apex. SPADIX: sessile, cylindric, normally with a few sterile flowers at extreme base, female zone cylindric, shorter than male and separated from it by equally thick zone of sterile male flowers, male zone equally thick, with shorter apical zone of sterile flowers, apex obtuse. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–2-androus, filaments distinct to very short, free to connate, connective slightly to deeply excavated, thecae either opposite or paired

54. Aridarum

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on one side and situated inside or outside connective cavity, apically narrowed into long or short horn, dehiscing by apical pore. POLLEN: inaperturate, ellipsoid-oblong, small (mean 23 µm., range 16–31 µm.), exine psilate. STERILE MALE FLOWERS: consisting of prismatic to obpyramidal, truncate staminodes with or without a small, central, apical cavity. FEMALE FLOWER: gynoecium shallow, laterally compressed, subhexagonal to subglobose, ovary 1-locular, ovules many, orthotropous to hemiorthotropous, funicle distinct, erect, placenta basal, stigma sessile, slightly concave centrally, as broad as ovary, contiguous with neighbouring stigmas. BERRY: globose or ellipsoid to cylindric, stigma remnant persistent, infructescence globular to slightly elongate. SEED: ellipsoid, elongate, testa longitudinally costate, embryo elongate, endosperm present. See Plates 54, 118D. CHROMOSOMES: 2n = 24. DISTRIBUTION: 7 spp.; Brunei, Malaysia (Borneo). ECOLOGY: tropical humid forest; rheophytes. NOTES: 2 sections recognized by Hotta (1965):– sect. Aridarum, sect. Caulescentia. ETYMOLOGY: Latin aridus (dry) and Arum (the genus); a misnomer as Aridarum species are rheophytes and do not occur in dry habitats. TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), Hotta (1965), Bogner (1979b, 1981a, 1983a).

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Plate 54. Aridarum. A, habit × 2/3; B, spadix × 6; C, detail of stamens × 6; D, detail of gynoecia × 6; E, stamen × 10; F, stamen, three quarter view × 10; G, gynoecium × 10; H, gynoecium, longitudinal section × 10; J, habit × 2/3; K, detail of leaf tip tubule × 3; L, detail of stamens × 6; M, detail of gynoecia × 6; N, stamen × 10; P, gynoecium × 10; Q, habit × 2/3; R, detail of stamens × 6; S, detail of gynoecia × 6; T, gynoecium × 10; U, stamen × 10. Aridarum nicolsonii: A–H, Bogner 1440 (K & Kew spirit collection 56427); A. burttii: J–K, Chai S34072 (K); L–P, Burtt 5116 (Kew spirit collection 34396); A. caulescens var. caulescens : Q–U, Boyce 250 (K & Kew spirit collection 56095).

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55. Heteroaridarum Heteroaridarum M. Hotta in Acta Phytotax. Geobot. 27 (3–4): 63 (1976). TYPE: H. borneense M. Hotta HABIT: evergreen herb, stem short, erect. LEAVES: several, distichous. PETIOLE: sheath with long, narrowly triangular ligule. BLADE: narrowly elliptic-oblanceolate, coriaceous; primary lateral veins pinnate, running into marginal vein, higher order venation parallel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole. SPATHE: slightly constricted, tube convolute, green, persistent, blade white, acuminate, caducous. SPADIX: sessile, female zone short, separated from male by short zone of sterile flowers, male zone much longer, subcylindric-ellipsoid, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3-androus, filaments as long as anthers, connate, with conspicuous tannin cells, connective thick, all three stamens parallel to one another, 2 outermost much larger with shortly horned thecae, central one much smaller with hornless thecae, all thecae opening by apical pore. POLLEN: unknown. STERILE MALE FLOWERS: composed

of slender, basally connate, clavate, truncate staminodes. FEMALE FLOWER: gynoecium depressed-globose, ovary 1-locular, ovules many, hemiorthotropous, placentae 2, basal and apical, apical one bearing smaller and apparently sterile ovules, stigma sessile, as broad as ovary, concave centrally. BERRY: unknown. SEED: unknown. See Plate 55. CHROMOSOMES: unknown. DISTRIBUTION: 1 sp.; Malaysia (Borneo). ECOLOGY: tropical humid forest; rheophyte. ETYMOLOGY: Greek heteros (different) and Aridarum.

Tribe Cryptocoryneae

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Tribe Cryptocoryneae Blume, Rumphia 1: 83 (1836). Laticifers absent in foliage leaves but present in stem, roots and cataphylls (Sivadasan, pers. comm.); aquatic to amphibious, stem rhizomatous, intravaginal squamules present; primary lateral veins pinnate, forming single marginal vein, higher order venation reticulate; inflorescence 1; spathe tube with margins connate, tube usually ± swollen basally to

C

A

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D J

K

Plate 55. Heteroaridarum. A, habit × 1/3; B, leaf × 1/3; C, detail of leaf tip tubule × 4; D, spadix × 2; E, stamens, side view × 10; F, stamens, front view × 10; G, stamens, longitudinal section × 10; H, staminodes × 10; J, gynoecium × 10; K, gynoecium, longitudinal section × 10. Heteroaridarum borneense: A–K, Abbe et al. 9845 (SAR).

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55. Heteroaridarum

form a “kettle”, blade lanceolate to cordate, twisted or longspiralled, often long-caudate; spadix entirely enclosed in spathe tube “kettle”, sterile terminal appendix apically adnate to spathe at apex of kettle; flowers unisexual, perigone absent; male flower 1(–2)-androus, stamens free, anther sessile or with short filament, excavated at apex with prominent margins, connective inconspicuous, thecae ending in prominent horn, dehiscing by apical pore, pollen exine smooth (psilate); ovary 1-locular, ovules orthotropous; embryo elongate, endosperm copious.

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56. Lagenandra Lagenandra Dalzell in Hooker’s J. Bot. Kew Gard. Misc. 4: 289 (1852). TYPE: L. toxicaria Dalzell Laticifers absent in foliage leaves but present in stem, roots and cataphylls (Sivadasan, pers. comm.). HABIT: small to medium-sized evergreen herbs, rhizome creeping, thick, rarely stoloniferous (L. nairii). LEAVES: several, cataphylls conspicuous. PETIOLE: sheath fairly long. BLADE: ptyxis involute, ovate, lanceolate, elliptic to almost linear, usually glabrous (hairy in L. nairii), coriaceous; primary lateral veins pinnate, weakly differentiated, running into inconspicuous marginal vein, secondary laterals parallel to primaries, higher order venation transverse-reticulate. INFLORESCENCE: solitary. PEDUNCLE: short, rarely long (L. bogneri), erect at anthesis, deflexed in fruit. SPATHE: marcescent, very thickwalled, outer surface green to purple, completely smooth or very warty on blade, margins basally connate into cylindric or ellipsoid tube (“kettle”), tube sometimes narrowing apically within (by thickening of walls, thus not apparent from outside), inner surface of lower region usually longitudinally furrowed, mouth of tube occluded by transverse, centrally perforated septum with separate transverse flap situated immediately below septum and partly or completely covering male zone of spadix, blade straight or twisted, opening only slightly by narrow longitudinal or spiral slit, or widely gaping, apex acute to long-caudate, inside smooth, uniformly warty or with warts in transverse bands, or cov-

ered with shortly branched processes, distinct “collar” around mouth of kettle present or absent. SPADIX: slender, female zone usually with 3–5(–7) spirals of gynoecia, rarely in pseudowhorl (L. nairii) or true whorl (L. gomezii), separated from male zone by slender naked axis, male zone cylindric to ellipsoid, terminal appendix small, conoid, apically adnate to spathe behind spathe flap; olfactory bodies present or absent above female flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1(–2)-androus, stamens free, anthers sessile or with short thick filaments, apex somewhat excavated with prominent thickened margin, connective inconspicuous, thecae opposite, each narrowed into an erect horn usually exceeding, rarely equalling thickened margin and dehiscing by apical pore. POLLEN: inaperturate, ellipsoid to ellipsoid-oblong, mediumsized (mean 36 µm., range 35–38 µm.), exine perfectly

56. Lagenandra

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K

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Plate 56. Lagenandra. A, habit × 2/3; B, infructescence × 1; C, habit × 2/3; D, leaf × 2/3; E, inflorescence with spadix detail viewed through opened kettle × 2; F, inflorescence with spadix detail viewed through opened kettle × 1; G, spadix × 3; H, stamen × 20; J, gynoecium × 10; K, gynoecium, longitudinal section × 10. Lagenandra toxicaria: A, Barnes 641 (K); Meebold 8877 (K); B, Barnes 1003 (Kew spirit collection 22447); L. koenigii: C–D, Bogner 564 (K & Kew spirit collection 57394); L. nairii: E, Bogner 1847 (Kew spirit collection 53950); L. dewitii: F–K, Bogner 1749 (Kew spirit collection 49709).

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psilate. FEMALE FLOWER: gynoecium free, broadly ellipsoid to globose, ovary 1-locular, ovules 1–12(–15), orthotropous, tapering towards micropyle, funicle short, bearing very long trichomes, placenta basal, stylar region thick, usually short, rarely long, stylar region and upper part of ovary often conspicuously warty, stigma discoid to hemispheric, sometimes oblique, relatively broad. BERRY: free, obovoid to prismatic-ellipsoid, often apically warty, rarely smooth, at maturity pericarp splitting and revolute basally to release seeds, infructescence usually globose, deflexed, prostrate. SEED: ellipsoid to narrowly ellipsoid or subcylindric, testa longitudinally costate, dark brown, embryo elongate, endosperm copious. See Plates 56, 119A. CHROMOSOMES: 2n = 36, 72. DISTRIBUTION: 14 spp.; tropical south Asia:– Bangladesh, India (Assam, south India), Sri Lanka. ECOLOGY: tropical humid forest; usually helophytes, rarely rheophytes, streams, marshes. NOTES: L. nairii and L. gomezii are quite distinct from the other species. ETYMOLOGY: Greek lagenos or lagynos (flask, bottle) and anêr, andros (man). TAXONOMIC ACCOUNTS: Engler (1920a), de Wit (1978,1990), Sivadasan (1982), Bogner & Jacobsen (1987), Nicolson (1988a), Kasselmann (1995).

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57. Cryptocoryne Cryptocoryne Fischer ex Wydler in Linnaea 5: 428 (1830). TYPE: C. spiralis (Retzius) Fischer ex Wydler (Arum spirale Retzius). Laticifers absent in foliage leaves but present in stem, roots and cataphylls (Sivadasan, pers. comm.). HABIT: small or mediumsized herbs, evergreen or rarely dormant in dry season (C. nevillii, C. consobrina), rhizome creeping, stoloniferous. LEAVES: many, cataphylls conspicuous when present. PETIOLE: sheath fairly long. BLADE: ptyxis convolute, cordate, ovate, lanceolate, elliptic, linear, rarely filiform (seasonally so in C. retrospiralis), mostly glabrous, rarely pubescent, sometimes bullate; primary lateral veins pinnate, usually weakly differentiated, running into inconspicuous marginal vein, higher order venation transverse-reticulate. INFLORESCENCE: solitary, usually appearing with the leaves (except C. consobrina). PEDUNCLE: short, elongating in fruit. SPATHE: tube with connate margins, at base swollen into ellipsoid to cylindric “kettle”, kettle rarely constricted, sometimes alveolar within, entrance with a lateral flap partially or completely covering male zone of spadix, upper part of tube long or short (in some species length depending on water depth), rarely absent (C. spiralis), sometimes widening or narrowing at mouth, blade cordate to lanceolate, erect or reflexed at anthesis, spreading, slightly twisted or long-spiralled, apex cuspidate to long-caudate, inner surface smooth, rough, uniformly warty or with warts in transverse bands, or covered with shortly branched protuberances, white, yellow, red, purple, brown, sometimes spotted, junction with tube usually marked by inconspicuous to very prominent “collar” which normally differs in colour and texture from blade. SPADIX: slender, female zone consisting of a single basal whorl of 4–8 upright gynoecia, with single whorl of rounded, subclavate or irregular olfactory bodies (?pistillodes) usually present immediately above, female and male zones separated by a long naked axis , or rarely fertile zones

almost contiguous, male zone ellipsoid to subcylindric, terminal appendix short, conical or irregularly oblong to subclavate, apically adnate to spathe behind flap, often breaking free during anthesis. FLOWERS: unisexual, perigone absent. MALE FLOWER: apparently 1-androus, stamens free, anthers sessile or with short filaments, apex excavated with very prominent, thickened margins, connective inconspicuous, thecae opposite, ending in short to long, attenuate horn usually projecting beyond thickened margins, dehiscing by apical pore. POLLEN: extruded in a droplet, inaperturate, ellipsoid to oblong, medium-sized (mean 34 µm., range 27–42 µm.), exine psilate. FEMALE FLOWER: ovaries connate, 1-locular, ovules 5 to many, orthotropous, funicle short, usually bearing numerous long trichomes extending between ovules, placenta subbasal to obliquely parietal (adaxial, morphologically basal), stylar region free, usually curved away from spadix axis, stigma discoid, rounded or elongated, often emarginate, often centrally concave. FRUIT: not a berry, all connate into an ovoid, smooth to verrucose syncarp dehiscing apically at maturity and becoming star-shaped. SEED: ellipsoid-oblong, straight or slightly curved, testa brown, not very thick, rough to slightly costate, or smooth, embryo elongate, straight, endosperm copious; seed sometimes viviparous and then seedling with many filiform cataphylls (C. ciliata). See Plates 57, 119B. CHROMOSOMES: 2n = 20, 22, 28, 30, 33, 34, 36, 42, 54, 66, 68, 72, 85, 88, 90, 102, 132. DISTRIBUTION: ca. 50 spp.; tropical Asia, Malay Archipelago:– Bangladesh, Brunei, Burma, Cambodia, China (Guandong, Guangxi, Guizhou, Yunnan), India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Laos, Malaysia (Borneo, Peninsula), Papua New Guinea, Philippines, Singapore, Sri Lanka, Thailand, Vietnam. ECOLOGY: tropical humid forest, freshwater tidal zone, rarely in brackish water (C. ciliata), submerged or emergent aquatics, helophytes or facultative rheophytes. NOTES: Jacobsen (1982) recognized 26 informal subgroups. C. spiralis is distinguished from other species of the genus in its spathe, which lacks the tube between kettle and blade and

57. Cryptocoryne

C RY P TO C O RY N E A E : C RY P TO C O RY N E

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D

F

E

J

B

G H

K A

L C

M

Plate 57. Cryptocoryne. A, habit × 2/3; B, inflorescence × 1; C, inflorescence with spadix detail viewed through opened kettle × 2; D, stamen × 30; E, female zone of spadix × 6; F, gynoecium, longitudinal section × 6; G, syncarp × 1; H, habit × 2/3; J, inflorescence × 1; K, inflorescence × 1; L, habit × 2/3; M, syncarp × 1. Cryptocoryne spiralis: A, Sivadasan CU 21406 (K); Sivadasan CU 21435 (K); B–G, Bogner 1829 (Kew spirit collection 53996); C. affinis: H, Bogner 1706 (K); J, Bogner 227 (Kew spirit collection 45235); C. ciliata: K, Cult. Kew 1953–203 (Kew spirit collection 16895); C. longicauda: L, Dransfield s.n. (Kew slide collection); M, Bogner 1735 (Kew spirit collection 50159).

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has a transverse perforated septum at the mouth of the kettle. It thus strongly resembles Lagenandra in this respect. However, C. spiralis shares with the other species of Cryptocoryne the connate gynoecia borne in a single whorl, the syncarpous infructescence and convolute ptyxis. ETYMOLOGY: Greek kryptos (hidden) and korynê (club); the spadix is entirely enclosed in the spathe tube and thus hidden from view. TAXONOMIC ACCOUNTS: Engler (1920a), Rataj (1975), Jacobsen (1982, 1985, 1988, 1991), Sivadasan (1982), Jacobsen & Bogner (1987), Jacobsen, Sivadasan & Bogner (1989), de Wit (1990), Kasselmann (1995).

C

Tribe Zomicarpeae Tribe Zomicarpeae Schott, Syn. Aroid. 33 (1856). Laticifers anastomosing; small, evergreen (Ulearum) or seasonally dormant (Zomicarpa, Zomicarpella); leaf blade cordate-sagittate (except Zomicarpa); primary lateral veins often arising at petiole insertion, higher order venation reticulate; spadix longer than spathe (except Zomicarpa, Zomicarpella amazonica), with sterile terminal appendix, female spadix zone adnate to spathe; flowers unisexual, perigone absent; stamens free; gynoecia few, ovary 1-locular, ovules anatropous, placenta basal.

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58. Zomicarpa Zomicarpa Schott, Syn. Aroid. 33 (1856). TYPE: Z. pythonium (Martius) Schott (Arum pythonium Martius). HABIT: small, seasonally dormant herbs, tuber subglobose to depressed-globose. LEAVES: few. PETIOLE: sheath short. BLADE: entire ovate-cordate when juvenile, adult blade trisect to pedatisect, lobes oblong-elliptic or oblong-lanceolate,

58. Zomicarpa

outer ones smaller; primary lateral veins of lobes pinnate, joined into submarginal collective vein at least in upper half of lobe, 1–2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: 3–5 in each floral sympodium, appearing before or with leaves. PEDUNCLE: longer than petiole, slender. SPATHE: somewhat constricted, entirely persistent, tube only laxly convolute, slightly gaping at anthesis, obliquely held, blade longer than tube, expanded at anthesis, distinctly or slightly bent forward at constriction, ± erect, sometimes fornicate, ovate-lanceolate or lanceolate, acuminate. SPADIX: shorter than spathe, female zone few- and laxly flowered, adnate to spathe, contiguous with male zone, male zone cylindric, longer than female, laxly to densely flowered, appendix erect or bent forward near base, clavate to subcylindric, smooth or ± covered with sterile flowers (Z. steigeriana). FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–2-androus, stamens free, filaments distinct or very short, connective distinctively coloured, thecae ellipsoid, opposite, dehiscing by apical pore. POLLEN: inaperturate, spherical to subspheroidal, medium-sized (mean 32 µm., range 32–33 µm.), exine spinose. STERILE FLOWERS: sparse when present, conic and acuminate towards base of appendix, otherwise wartlike. FEMALE FLOWER: gynoecium cylindric or ovoid to subglobose, ovary 1-locular, ovules 6–11, anatropous, funicle distinct, placenta basal, stylar region shortly attenuate, stigma broad, discoid-hemispheric. BERRY: globose to depressed-globose, whitish below, darker at apex, few–several-seeded. SEED: ovoid to ellipsoid, testa smooth, thin, transparent, micropyle shortly rostrate, funicle strophiolate, swollen, white, embryo axile, elongate, endosperm copious. See Plates 58, 119C. CHROMOSOMES: 2n = 20. DISTRIBUTION: 3 spp.; Brazil (Northeast). ECOLOGY: tropical humid forest, upland gallery forest, deciduous forest; geophytes on forest floor. ETYMOLOGY: Greek zômê (woman’s girdle) and karpos (fruit); refers to spathe constriction. TAXONOMIC ACCOUNTS: Peyritsch (1879), Engler (1920a), Bogner (1980d).

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59. Zomicarpella Zomicarpella N.E. Brown in Gard. Chron., ser.2, 16: 266 (1881). TYPE: Z. maculata N.E. Brown HABIT: small herbs with creeping rhizome. LEAVES: 1–3. PETIOLE: sheath short. BLADE: cordate-sagittate to hastatesagittate, often variegated with pale green blotches; primary lateral veins mostly arising at petiole insertion, running into margin or forming a submarginal collective vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing with leaves. PEDUNCLE: slender, longer or shorter than petiole. SPATHE: oblong-lanceolate, fully expanded and unconstricted (Z. maculata), or constricted between tube and blade, tube convolute, blade expanded, erect at first, then reflexed, persistent in fruit (Z. amazonica). SPADIX: longer or shorter than spathe, female zone adnate to spathe, contiguous or separated from male by short axis (Z. maculata), male zone subequal to female, cylindric, terminal appendix more slender, ± elongate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, stamens densely arranged, free, anthers subsessile, filaments short but distinct, connective inconspicuous, thecae opposite,

ZOMICARPEAE : ZOMICARPELLA

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B

C

E

F

A G

D

Plate 58. Zomicarpa. A, habit with mature leaf × 1; B, juvenile leaf × 1; C, semi–mature leaf × 1; D, spadix × 5; E, stamen × 20; F, gynoecium, longitudinal section × 20; G, infructescence, nearside half of spathe removed × 2. Zomicarpa riedeliana: A, Noblick, Lemos & Valdomiro 3202 (K); B, Cult. Berkley (K); C, Mayo s.n. (K); D–F, Bogner 1213 (Kew spirit collection 42464); G, Schott, Icones Aroideae 3760 (K).

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A B D

F

H

G M C

L

J

K

E

Plate 59. Zomicarpella. A, leaf × 1; B, inflorescence, nearside half of spathe folded back × 3; C, habit × 1; D, leaf × 1; E, inflorescence, nearside half and upper part of spathe removed × 3; F, stamen, top view × 15; G, stamen, side view × 15; H, stamen, longitudinal section × 15; J, gynoecium × 15; K, gynoecium, longitudinal section × 15; L, fruit × 3; M, seed × 4. Zomicarpella maculata: A–B, Cult. Linden (K); Z. amazonica: C–D, Bogner 1985 (K & Kew spirit collection 57529 & 57400); E–M, Bogner 1985 (Kew spirit collection 57529 & 57400).

ZOMICARPEAE : ZOMICARPELLA

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59. Zomicarpella

subglobose, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, spherical, small (18–20 µm. diam.), exine spinose. FEMALE FLOWER: gynoecium ellipsoid-oblong, or depressed-bottle-shaped, somewhat laterally compressed, ovary 1-locular, ovules 1–6, anatropous, funicle short or very short, placenta basal, stylar region short, distinct, narrower than ovary, stigma small, discoid-hemispheric. BERRY: depressed-globose, whitish, stigma remnant persistent, 1–3seeded. SEED: irregularly ellipsoid, somewhat compressed laterally, testa thin, smooth, whitish, raphe conspicuous, hilum somewhat sunken, purple, embryo ellipsoid to pear-shaped, small, endosperm copious. See Plates 59, 119D. CHROMOSOMES: 2n = 26. DISTRIBUTION: 2 spp.; Brazil (western Amazonia), ?Colombia. ECOLOGY: tropical humid forest (“terra firme”); geophytes on forest floor. NOTES: Zomicarpella maculata N.E. Br. was described from a cultivated plant known to have originated in tropical America, but without a more exact provenance. No other collection of this species has been seen. ETYMOLOGY: Zomicarpa and Latin -ella (diminutive); refers to similarity to Zomicarpa. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1979a), Bogner (in press).

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insertion, forming a submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: slender, subequal to or longer than petiole, base enclosed by a single relatively long cataphyll. SPATHE: oblong-lanceolate, greenish, unconstricted, completely expanded, margins revolute, persistent and dark green in fruit. SPADIX: slender, subequal to spathe, female zone adnate to spathe, separated from male zone by a longer sterile zone, lower part of sterile zone naked or only sparsely covered with sterile flowers, upper part with short dense zone of staminodes, male zone short, composed of 6–7 whorls of stamens, terminal appendix digitiform, base composed of a few whorls of apically rounded staminodes, becoming smooth above. FLOWERS: unisexual, perigone absent. MALE FLOWER: apparently 1androus, stamens sessile, free, subtruncate, connective ± broad, thecae ellipsoid to globose, dehiscing by apical pore. POLLEN: inaperturate, spherical, medium-sized (diam. 26–32 µm.), exine spinose. STERILE FLOWERS: depressed globose or cylindric; staminodes below male zone in 2–3 whorls, rounded apically. FEMALE FLOWER: gynoecium narrowly subcylindric, ovary 1-locular, ovule 1, anatropous, funicle short, placenta basal, stylar region ± as thick as ovary, stigma discoid, as broad as style. BERRY: subcylindric to ellipsoid, stigmatic remnant persistent, 1seeded, dark green and red-spotted. SEED: ellipsoid, testa thin, white, smooth, transparent, embryo ellipsoid, dark green with red spots externally, whitish internally, undifferentiated, endosperm absent. See Plates 60, 120A. CHROMOSOMES: 2n = 14. DISTRIBUTION: 1 sp.; Brazil (western Amazonia), Peru (Amazonia). ECOLOGY: tropical humid forest (“terra firme”); geophytes on forest floor. ETYMOLOGY: named after Ernst Ule (1854–1915) and Arum. TAXONOMIC ACCOUNTS: Engler (1920a), Boyce (1995e).

60. Ulearum Ulearum Engler in Bot. Jahrb. 37: 95 (1905, “1906”). TYPE: U. sagittatum Engler HABIT: small evergreen herb, rhizome creeping horizontally just below the soil surface. LEAVES: 1–several, 1 marcescent cataphyll enclosing base of petiole. PETIOLE: sheath very short. BLADE: subreniform to broadly sagittate, often variegated; primary lateral veins arising from petiole

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60. Ulearum

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F

E

C

D B A

Plate 60. Ulearum. A, habit × 2/3; B, inflorescence, nearside half of spathe removed × 3; C, gynoecium × 20; D, gynoecium, longitudinal section × 20; E, staminode × 20; F, stamen, side view × 10. Ulearum sagittatum: A, Ule 6323 (K); B, Jangoux et al. INPA 138864 (Kew spirit collection 56426); C–F, Bogner 1947 (K & Kew spirit collection 56424).

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61. Filarum Filarum Nicolson in Brittonia 18: 348 (1967). TYPE: F. manserichense Nicolson (“manserichensis”).

Tribe Caladieae Schott in Schott & Endlicher, Melet. Bot. 18 (1832).

HABIT: small herb, tuber subglobose. LEAVES: few. PETIOLE: sheath short. BLADE: entire, cordate; primary lateral veins 3 per side, arising from petiole insertion, lower 2 pairs retrorse, all arcuately curving upwards forming submarginal collective vein, marginal vein distinct, higher order venation reticulate. INFLORESCENCE: 1–3 in each floral sympodium, appearing with leaves. PEDUNCLE: slender, longer or shorter than petiole. SPATHE: persistent, oblonglanceolate, pale green. SPADIX: very slender, much longer than spathe, female zone adnate to spathe, few-flowered, separated from male zone by longer sterile zone, male zone elongate, purple, sparsely flowered, with a very few sterile rudimentary flowers occurring above and below male zone, appendix slender, naked, elongate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, filaments short, connective extremely long and drawn out into a filiform flexuose thread, thecae globose, dehiscing by apical pore. POLLEN: inaperturate, spherical, small to mediumsized (mean 21 µm.), exine spinose. FEMALE FLOWER: ovary oblong, 1-locular, ovule 1, anatropous, funicle short, placenta basal, stylar region attenuate, stigma small, discoid. BERRY: ellipsoid, smooth, bearing the style and stigma remains. SEED: ellipsoid, testa thin, smooth, punctate, embryo ellipsoid, endosperm absent. See Plate 61. CHROMOSOMES: 2n = 28. DISTRIBUTION: 1 sp.; Peru (Amazonia). ECOLOGY: tropical humid forest; geophytes on forest floor. ETYMOLOGY: Latin filum (thread) and Arum; named after the thread-like spadix and stamen connectives.

Laticifers anastomosing; terrestrial, geophytic, rarely aquatic (Jasarum) or climbing hemiepiphytes (Syngonium), stem tuberous, rhizomatous or aerial; basal ribs well-developed, primary lateral veins of leaf pinnate, forming submarginal collective vein, 1 or more marginal veins also usually present, higher order venation reticulate; spadix: female zone often separated from male by subconical to attenuate zone of synandrodes (sterile male flowers), male zone subcylindric to subclavate, sterile appendix absent (except some Hapaline); flowers unisexual, perigone absent; stamens connate into synandria, synandria truncate, margins sinuous, thecae dehiscing by short apical slit, pollen extruded in strands (except Scaphispatha); ovules anatropous to hemianatropous; endosperm copious (except Jasarum, Syngonium, Hapaline).

61. Filarum

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THE GENERA OF ARACEAE

62. Scaphispatha Scaphispatha Brongniart ex Schott, Prodr. syst. Aroid. 214 (1860). TYPE: S. gracilis Brongniart ex Schott HABIT: seasonally dormant herb, tuber globose to depressed-globose. LEAVES: usually solitary, sometimes 2. PETIOLE: slender, sheath very short. BLADE: ovate-subcordate to ovate-sagittate, peltate, apex cuspidateacuminate; basal ribs well-developed, primary lateral veins pinnate, forming submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate. INFLO-

62. Scaphispatha

C

C

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E

B D

F

G

A

H

C

Plate 61. Filarum. A, habit × 1; B, leaf × 1; C, inflorescence, nearside spathe base folded back × 4; D, stamen, side view × 16; E, stamen, view from below × 16; F, gynoecium × 16; G, gynoecium, longitudinal section × 16; H, fruit × 10. Filarum manserichense A, D–G, Wurdack 2402 (K); B–C, H, Mexia 6353 (K).

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H

E

F

D

B

C

G

A

Plate 62. Scaphispatha. A, habit × 1; B–C, inflorescences × 1; D, spadix × 5; E, synandrium, top view × 30; F, synandrium, side view × 30; G, gynoecium, longitudinal section × 30; H, infructescence × 1. Scaphispatha gracilis: A–C, Hatschbach & Koczicki 33081 (K); D–H, Bogner 1211 (Kew spirit collection 42451).

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RESCENCE: solitary, appearing well before leaf. PEDUNCLE: very long, slender. SPATHE: decurrent on peduncle, slightly constricted between tube and blade, light green without, white within, tube convolute before and after anthesis, gaping at anthesis, persistent to fruiting stage, blade fully expanded at anthesis, at first erect, later reflexed, marcescent after anthesis. SPADIX: sessile, cylindric, hardly constricted, shorter than spathe, fertile to apex, densely flowered, male and female zones contiguous. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4-androus, stamens connate into truncate, deeply lobed synandrium, connective not greatly thickened, thecae lateral, dehiscing apically by broad, pore-like slit. POLLEN: inaperturate, spherical to subspheroidal, medium-sized (mean 45 µm.), exine coarsely verrucate with large polygonal flat-topped verrucae. FEMALE FLOWER: ovary ovoid, 1-locular, ovules 3–5, anatropous, funicle short, placenta basal, stylar region distinct, attenuate, much narrower than ovary, stigma small, discoid-subcapitate, only slightly broader than style. BERRY: subglobose to obovoid, always 1-seeded, stigma remnants persistent, fruiting very rapidly (ca.10 days after anthesis), whitish-grey. SEED: subglobose, testa smooth and thin, greyish to brown, covered with minute brownish spots, raphe pronounced, swollen, embryo elongate, straight, endosperm copious. See Plates 62, 120B. CHROMOSOMES: 2n = 28. DISTRIBUTION: 1 sp.; Bolivia, Brazil (southern Amazonia, Central West, Northeast). ECOLOGY: tropical seasonal and dry forest, open woodland; geophytes, temporarily wet places, flowering at beginning of rainy season after first rains. ETYMOLOGY: Greek skaphê (boat, bowl) and -ion (diminutive) and “spathê” (spathe); refers to spathe shape. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1980d).

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63. Caladium Caladium Ventenat, Descript. Pl. Nouv. Jard. Cels, 30 (1801). LECTOTYPE: C. bicolor (Aiton) Ventenat (Arum bicolor Aiton, see Hubbard & Rehder, Bot. Mus. Leafl. 1: 3. 1932). SYNONYMS: Phyllotaenium André in Ill. Hort. 19: 3 (1872); Aphyllarum S. Moore in Trans. Linn. Soc. London, Bot. ser. 2, 4: 501 (1895). HABIT: seasonally dormant or evergreen herbs, stem tuberous, subglobose. LEAVES: several. PETIOLE: sheath distinct. BLADE: usually peltate, sometimes not (C. lindenii), often variegated, cordate-sagittate or sagittate, rarely trisect (C. ternatum); basal ribs well-developed, primary lateral veins pinnate, forming submarginal collective vein, 1–2 marginal veins also present, secondary and tertiary laterals arising from the primaries at a wide angle, forming interprimary collective vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing with or before leaves. PEDUNCLE: rather long, often as long as petiole. SPATHE: constricted, tube with convolute margins, usually ventricose-globose, persistent, green, eventually splitting in fruit, blade white, boat-shaped, gaping, marcescent after anthesis and deciduous. SPADIX: a little shorter than spathe, stipitate or sessile, densely flowered, female zone cylindric-conoid or ellipsoid, separated from male by longer, subconic to attenuate, basally thicker zone of sterile male flowers, male zone fertile to apex, subcylindric to subclavate,

63. Caladium

more than twice as long as female zone. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–5-androus, stamens connate into obpyramidal, truncate to ± convex, sinuously subhexagonal synandrium, common connective thick, thecae lateral, extending nearly to base of synandrium, oblonglanceolate, dehiscing by short apical slit. POLLEN: extruded in strands, shed in monads, inaperturate, spherical to subspheroidal, medium-sized (mean 41 µm., range 38–43 µm.), exine psilate or obscurely dimpled or verruculate. STERILE MALE FLOWERS: synandrodes depressed-obpyramidal, compressed, truncate, lowermost often larger and prismatic, uppermost narrow and elongated. FEMALE FLOWER: gynoecium cylindric to obconic, ovary 1–2(–3)-locular, ovules 1–20, anatropous, funicle short, placentae subbasal, pseudoaxile or parietal, stylar region free or rarely coherent (C. paradoxum), as broad as ovary or broader in obconic gynoecia, stigma nearly as wide as ovary. BERRY: 1–many-seeded (C. ternatum usually 1-seeded), white. SEED: ovoid to ellipsoid, raphe somewhat prominent, integument succulent, testa thick, longitudinally costate, costae angled, embryo axile, elongate, endosperm copious. See Plates 63, 120C. CHROMOSOMES: 2n = 22, 26, 28, 30, 32. DISTRIBUTION: ca. 12 spp.; tropical America, West Indies:– N. Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Lesser Antilles, Panama, Peru, Puerto Rico, Surinam, Venezuela. ECOLOGY: tropical humid forest, open woodlands; geophytes, forest floor, river margins and damp sites. NOTES: Caladium bicolor and its many cultivated varieties are cultivated throughout the world as ornamental plants. This species occurs throughout tropical America. ETYMOLOGY: Latinization of Malay word keladi, a vernacular name in Malaysia for several colocasioid genera.

CALADIEAE : CALADIUM

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S

B

N

J

Q

R

P M A

D

G

E L H

F

K

C

Plate 63. Caladium. A, leaf × 1/3; B, base of plant × 1/2; C, spadix × 1; D, synandrium × 10; E, upper synandrode × 5; F, lower synandrode × 5; G, gynoecium × 10; H, gynoecium, transverse section × 10; J, infructescence × 2/3; K, leaf × 1/3; L, spadix × 1; M, inflorescence × 2/3; N, synandrium × 10; P, synandrode × 10; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, habit × 1/5. Caladium aristeguietae: A–B, Bunting 2201 (K); C–H, Bunting 2201 (Kew spirit collection 58022); J, Bunting 4521 (K); C. coerulescens: K, Bunting 4980 (K); L, Bunting 4865 (K); C. lindenii: M–R, Cult. Kew 1982–02775 (Kew spirit collection 39521); C. bicolor: S, Lowe 3.1962, 7220 (Kew slide collection).

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TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), Madison (1976b, 1981), Bogner (1984d), Grayum (1986c), Mayo & Bogner (1988).

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64. Jasarum Jasarum Bunting in Acta Bot. Venezuelica 10: 264 (1977, “1975”). TYPE: J. steyermarkii Bunting HABIT: evergreen aquatic herb, stem thick, hypogeal, upright, annulate, terete, surface brown, white within, usually unbranched, roots thick, somewhat spongy. LEAVES: 6–12, dark green in all parts except whitish hypogeal lower part of petiole. PETIOLE: sheath long, fleshy. BLADE: linear, somewhat bullate, acute to cuspidate, cuneate at base, midrib pronounced; primary lateral veins pinnate, numerous, 35–60 per side, joined into submarginal collective vein, marginal vein also present, higher order venation reticulate. INFLORESCENCE: solitary, held above water level. PEDUNCLE: much longer than petiole. SPATHE: erect, constricted between tube and blade, tube convolute, persistent, blade long-acuminate, never reflexed, marcescent after anthesis and later deciduous, bronze-green outside, cream-coloured within. SPADIX: shorter than spathe, subcylindric, basal half of female zone adnate to spathe, separated from male by zone of sterile flowers (synandrodes), male zone fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4-androus, stamens connate into synandrium, synandrium truncate, irregularly 4–6-angled, thecae 6–8, lateral, elongate, dehiscing by apical pore. POLLEN: ellipsoid, medium-sized (mean 44 µm.),

64. Jasarum

exine foveolate-reticulate. FEMALE FLOWER: ovary cylindric, 1-locular, ovules (1–)2–3(–4), anatropous, funicle distinct, placenta basal, stylar region broader than ovary, stigma discoid-hemispheric. BERRY: subovoid, tightly packed, dull maroon- green apically, stigma remains persistent, pericarp fleshy, usually 1-seeded, sometimes 2-seeded. SEED: large, obovoid, compressed when paired, testa smooth, embryo globular, large, green, plumule welldeveloped, raphe prominent, endosperm nearly absent. See Plates 64, 120D. CHROMOSOMES: 2n = 22. DISTRIBUTION: 1 sp.; tropical South America: Venezuela, Guyana. ECOLOGY: tropical uplands; submerged aquatic in oligotrophic (blackwater) streams. ETYMOLOGY: named after Julian A. Steyermark (1909–1988). TAXONOMIC ACCOUNTS: Bunting (1977), Bogner (1985b).

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65. Xanthosoma Xanthosoma Schott in Schott & Endlicher, Melet. Bot. 19 (1832). LECTOTYPE: X. sagittifolium (L.) Schott (“sagittaefolium”; Arum sagittaefolium L.; see Nicolson 1975). SYNONYMS: Acontias Schott in Schott & Endlicher, Melet. Bot. 19 (1832); Cyrtospadix K. Koch in Index Sem. Hort. Berol. 1853 App.: 13 (1853). Latex milky. HABIT: small to gigantic, sometimes arborescent, evergreen or seasonally dormant herbs, stem either a thick, subcylindric, hypogeal tuber often producing smaller tubers on stoloniferous side branches or often with a distal, epigeal, massive, arborescent upper part, or an entirely hypogeal subglobose tuber, bearing many very small tubercles in some spp. (X. pubescens, X. viviparum), tuber sometimes yellow within. LEAVES: several, rarely pubescent. PETIOLE: sheath usually rather long. BLADE: cordate, sagittate, hastate, trifid, trisect, pedatifid or pedatisect, rarely linear-lanceolate to ovate with emarginate base, rarely peltate; basal ribs well-developed, often denuded proximally, primary lateral veins pinnate, forming submarginal collective vein, 1 or more distinct marginal veins also present, secondary and tertiary laterals arising from the primaries at a wide angle, forming interprimary collective vein, higher order venation reticulate. INFLORESCENCE: 1 to many in each floral sympodium, always appearing with leaves. PEDUNCLE: usually rather short, rarely long. SPATHE: strongly constricted, tube with convolute margins, ovoid to ellipsoid, usually ventricose, rather thick-walled in large species, persistent, blade boat-shaped-oblong to oblong-lanceolate, gaping and erect or sometimes reflexed at anthesis, marcescent after anthesis and then deciduous. SPADIX: shorter than spathe, densely flowered, female zone cylindric-conoid, often obliquely inserted onto peduncle, separated from male zone by longer, conoid to attenuate, basally thicker zone of sterile male flowers, male zone cylindric-conoid, longer than female zone, usually fertile to apex, rarely with a few sterile flowers at extreme apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4–6-androus, stamens connate into a truncate-obpyramidal, subpentagonal or hexagonal synandrium, anthers lateral, nearly reaching base of synandrium, common connective thick, thecae oblong or tapering basally, dehiscing by subapical pore or short slit. POLLEN: extruded in strands, shed in

CALADIEAE : XANTHOSOMA

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Plate 64. Jasarum. A, habit, peduncle partly removed × 1/2; B, detail of leaf venation × 2/3; C, inflorescence × 2/3; D, spadix × 2; E, synandrium, side view × 15; F, gynoecium × 15; G, gynoecium, longitudinal section × 15; H, infructescence × 2/3; J, berry, side view × 2. Jasarum steyermarkii: A, Bunting 4628 (K); Herkner & Jeschke s.n. (Kew spirit collection 29047.618); Aroideana 8(2): 55–63, f.4 (1985); B–J, Cult. Munich (Kew spirit collection 57524); Herkner & Jeschke s.n. (Kew spirit collection 29047.618).

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tetrads, inaperturate, ellipsoid to subspherical, mediumsized (mean 42 µm., range 35–49 µm., tetrads:– mean 76 µm., range 62–97 µm.), exine minutely punctate- or fossulate-verruculate. STERILE MALE FLOWER: composed of obpyramidal truncate, laterally compressed synandrodes, lowermost larger, uppermost narrower and more elongated (in sense of spadix axis). FEMALE FLOWER: ovary ovoid, 2–4-locular, more rarely 1-locular, ovules (12–)20 to very numerous, anatropous or hemianatropous, funicles rather long, placentae usually pseudoaxile, or sometimes parietal or axile basally, stylar region broader than ovary, usually discoid-thickened and coherent to weakly connate with those of neighbouring flowers, rarely free and swollen (X. plowmanii), stigma subhemispheric or 2–4-lobed, yellow, narrower than style. BERRY: cylindric, somewhat furrowed apically, whitish to orange, many-seeded. SEED: ovoid, testa costate, embryo axile, subequal to endosperm, endosperm copious. See Plates 65, 121A. CHROMOSOMES: 2n = 22, 26, 39, 52. DISTRIBUTION: ca. 57 spp.; tropical and southern subtropical America, West Indies:– N. Argentina, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad, Venezuela. ECOLOGY: tropical moist and humid forest, subtropical forest; geophytes on forest floor, in wet places, swamps, river banks, seasonally flooded sites, grassy places, plantations, some species are weedy. NOTES: Engler in Engler & Krause (1920) recognized 2 sections:– section Xanthosoma, section Acontias.

65. Xanthosoma

VERNACULAR NAMES AND USES: “cocoyam”, “tannia”; X. sagittifolium and its many varieties are important subsistence food plants throughout the humid tropics of the world due to their starch- and protein-rich tuberous stems (see chapter 16). ETYMOLOGY: Greek “xanthos” (yellow) and “soma, somatos” (body); refers to the yellow colour of the stem tissue present in several species. TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), Madison (1976b, 1981), Bogner (1986b), Grayum (1986c), Mayo & Bogner (1988), Okeke (1992).

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66. Chlorospatha Chlorospatha Engler in Gartenflora 27: 97 (1878). TYPE: C. kolbii Engler SYNONYM: Caladiopsis Engler in Bot. Jahrb. 37: 139 (1905). Latex milky. HABIT: small to medium, usually evergreen herbs, stem epigeal to ± subterranean, acaulescent to elongate, decumbent to erect. LEAVES: 1 to several. PETIOLE: sheath long. BLADE: cordate, sagittate, hastate, trifid, trisect, pedatifid or pedatisect; basal ribs well-developed, primary lateral veins pinnate, forming submarginal collective vein, 1 or more distinct marginal veins also present, secondary and tertiary laterals arising from the primaries at a wide angle, forming interprimary collective vein, higher order venation reticulate. INFLORESCENCE: 3–8 in each floral sympodium. PEDUNCLE: very slender, much shorter than petiole and supported by sheath. SPATHE: constricted, tube

66. Chlorospatha

CALADIEAE : CHLOROSPATHA

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G Plate 65. Xanthosoma. A, leaf × 1/4; B, base of plant × 1/5; C, inflorescence × 1/5; D, detail of synandria, top view × 5; E, detail of gynoecia, top view × 5; F, synandrium, side view × 5; G, gynoecia, left hand gynoecium in longitudinal section × 5; H, infructescence × 1/4; J, leaf × 1/3; K, leaf × 1/2; L, inflorescence, nearside half of spathe removed × 2/3; M, detail of synandrodes, top view × 15; N, synandrode, side view × 15; P, synandrium, side view × 15; Q, gynoecium × 15; R, gynoecium, longitudinal section × 15; S, ovules × 25. Xanthosoma violaceum: A–B, Engler 245 (K); C, Vollesen (Kew slide collection); D–H, Skvortzov (Kew spirit collection 29047.189); X. striatipes: J, Krapovickas & Cristóbal 35351 (K); X. helleborifolium: K, Broadway 6340 (K); X. plowmanii: L–S, Plowman et al. 8460 (Kew spirit collection 49708).

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Plate 66. Chlorospatha. A, habit × 1/2; B, spadix × 2; C, synandrium, side view × 15; D, synandrode, side view × 15; E, gynoecium, side view × 15; F, gynoecium, longitudinal section × 15; G, leaf × 1/4; H, inflorescences with associated leaf × 1/2; J, inflorescence × 2; K, gynoecium, longitudinal section × 15. L, spadix × 2; M, gynoecium, side view × 15. Chlorospatha longipoda: A, Plowman 14063 (K); B–F, Plowman 3979 (Kew spirit collection 49717); C. croatiana subsp. enneaphylla: G, Lawrence 794 (K); C. atropurpurea: H, Dodson 5911 (K); C. corrugata: J–K, Spear s.n. (Kew spirit collection 42452); C. kolbii : L–M, Franke s.n. (Kew spirit collection 62897).

CALADIEAE : CHLOROSPATHA

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with convolute margins, narrow, elongate, ± cylindric to narrowly ellipsoid, persistent, blade boat-shaped to more widely expanded, sometimes fornicate, rather narrowly elliptic to lanceolate, marcescent after anthesis and deciduous. SPADIX: female zone free or adnate to spathe, laxly or densely flowered, separated from male zone by longer or shorter laxly or densely flowered sterile zone, male zone densely flowered, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–5-androus, stamens connate into truncate synandrium, deeply or shallowly-lobed, fused connectives thickened, thecae extending almost to base of synandrium, oblong, dehiscing by short longitudinal slit. POLLEN: extruded in strands, shed in tetrads, inaperturate, spherical or subspheroidal, medium-sized (mean 26 µm., range 24–29 µm., tetrad mean 45 µm., range 41–48 µm.), exine psilate or very obscurely punctate to obscurely verruculate to foveolate-reticulate. STERILE FLOWERS: staminodes either free or partially or completely connate into irregular, fungiform or 3–4 lobed synandrodes, rarely prismatic. FEMALE FLOWER: gynoecium ± ovoid to subhemispheric, ovary (1–)2–4(–5)- locular, ovules several per locule, anatropous to hemianatropous, funicle rather long, placentae pseudoaxile or axile, rarely basal (C. longipoda), stylar region short to relatively long, usually expanded into thin, spreading, evanescent mantle contiguous with neighbouring ones, usually containing numerous red chromoplasts, more rarely style with only a short, rimlike lateral outgrowth (C. longipoda, C. mirabilis) or with soft tuberculate outgrowth (C. kolbii), stigma subhemispheric or subcapitate or slightly lobed. BERRY: depressed-globose, somewhat 3–5-furrowed, stigma remnant persistent, manyseeded, white to yellowish. SEED: minute, ovoid to ellipsoid, white, testa longitudinally costate, embryo elongate, axile, endosperm copious. See Plates 66, 121B. CHROMOSOMES: 2n = 26. DISTRIBUTION: ca. 16 spp.; tropical America:– Colombia, Costa Rica, Ecuador, Panama, Peru. ECOLOGY: tropical humid forest; terrestrial on forest floor, well shaded creek beds or in boggy areas. NOTES: Phylogenetically, Chlorospatha may be a derived offshoot of Xanthosoma, and C. longipoda is somewhat intermediate between the two genera. ETYMOLOGY: Greek chlôros (green) and spathê (spathe). TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), Madison (1981), Bogner (1985a), Grayum (1986c, 1991a), Mayo & Bogner (1988).

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67. Syngonium Syngonium Schott in Wiener Z. Kunst 1829 (3): 780 (1829). TYPE: S. auritum (L.) Schott (Arum auritum L.). SYNONYM: Porphyrospatha Engler in A. & C. de Candolle, Monogr. Phan. 2: 289 (1879). Latex milky. HABIT: root-climbing or shortly creeping, evergreen herbs, internodes short to elongate, green. LEAVES: numerous. PETIOLE: sheath long. BLADE: at maturity cordate, oblong-cordate, lanceolate, sagittate, trifid to trisect, pedatifid to pedatisect, rarely pinnatifid, when juvenile entire, ovate to sagittate or cordate-sagittate or hastate; basal ribs usually well-developed, primary lateral veins pinnate, usually forming a submarginal collective vein, 1–2 distinct marginal veins also present, or sometimes primaries long-

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67. Syngonium

arcuate and running into margin (e.g. S. schottianum), secondary laterals ± parallel to primaries, higher order venation reticulate to transverse-reticulate. INFLORESCENCE: 1–8 in each floral sympodium. PEDUNCLE: much shorter than petiole, erect at anthesis, pendent in fruit. SPATHE: strongly constricted between tube and blade, tube convolute, ovoidellipsoid, or cylindric or globose, sometimes ventricose, persistent, blade usually cream, boat-shaped to expanded at anthesis, erect to spreading, usually marcescent after anthesis, later deciduous. SPADIX: sessile, much shorter than spathe, female zone cylindric to conoid, separated from male zone by zone of sterile flowers, sterile zone constricted, male zone clavate to cylindric or ellipsoid, longer than female zone, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4-androus, stamens connate into synandrium, synandrium obpyramidal, truncate to rounded at apex, sometimes prominent above thecae, often deeply lobed, common connective thick, overtopping anthers, thecae oblong, dehiscing by apical pore or short slit. POLLEN: extruded in strands, inaperturate, ellipsoidoblong to spherical or subspheroidal, medium-sized (mean 48 µm., range 34–75 µm.), exine minutely fossulate or verruculate, narrowly rugulate and minutely scabrate, spinose or tuberculate. STERILE MALE FLOWERS: synandrodes ± prismatic. FEMALE FLOWER: gynoecia connate, rarely ± free at anthesis and becoming connate later (S. steyermarkii), ovary obovoid or oblong-obovoid, (1–)2(–3)-locular, ovules 1(–2) per locule, anatropous, funicle short, placenta axile near septum base to parietal-basal, stylar region as broad as ovary and ± truncate or very slightly attenuate, stigma discoid or 2-lobed, rarely globose, discoid-capitate or cup-shaped, narrower than ovary. BERRIES: connate, forming ovoid to ellipsoid, usually brown, sometimes white, fleshy syncarp, usually exposed at maturity by opening of persistent orange, yellow, red or violet spathe tube. SEED:

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Plate 67. Syngonium. A, leaf × 1/3; B, inflorescence × 1/2; C, habit × 1/2; D, infructescences and associated stem × 1/2; E, leaf × 1/3; F, inflorescence × 1/2; G, leaf × 1/3; H, spadix × 1; J, synandrium, side view × 8; K, detail of gynoecia, top view × 4; L, gynoecium, longitudinal section × 6; M, gynoecium, transverse section × 6. Syngonium steyermarkii: A, 201/12 (Kew slide collection); B, Croat 47201 (Kew spirit collection 56140); S. armigerum: C, Grayum 6677 (K); S. podophyllum: D, Silverstone-Sopkin et al. 5468 (K); S. neglectum: E–F, Bunting 1658 (K); S. schottianum: G, Cult. Kew June 1878 (K); H–K, Sugden 425 (Kew spirit collection 29047.347); S. vellozianum: L–M, Harley et al. 17874 (Kew spirit collection 46608).

CALADIEAE : SYNGONIUM

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ovoid to ellipsoid, rather large, testa smooth, thin, black or dark brown, shiny, embryo large, ellipsoid to subglobose, endosperm absent. See Plates 67, 121C. CHROMOSOMES: 2n = 28 (24, 26). DISTRIBUTION: 35 spp.; tropical America, West Indies:– Belize, Bolivia, Brazil, Colombia, Costa Rica, ?Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, ?Lesser Antilles, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad, Venezuela. ECOLOGY: tropical humid forest and disturbed areas; climbing epiphytes and hemiepiphytes, creeping when juvenile. NOTES: Croat (1982) recognizes 4 sections:– sect. Oblongata, sect. Cordata, sect. Pinnatiloba, sect. Syngonium. ETYMOLOGY: Greek syn- (together), gonê (womb) and -ion (diminutive); refers to connate gynoecia. TAXONOMIC ACCOUNTS: Engler & Krause (1920), Croat (1982).

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68. Hapaline Hapaline Schott, Gen. Aroid. t. 44 (1858), nom. cons. TYPE: H. benthamiana Schott SYNONYM: Hapale Schott in Oesterr. bot. Wochenbl. 7: 85 (1857). HABIT: small to moderate, slender, seasonally dormant or evergreen herbs, tubers small, depressed-globose. LEAVES: 1–few, usually solitary. PETIOLE: sheath short. BLADE: cordate-sagittate, sagittate or hastate, sometimes pale green or silvery variegated; primary lateral veins pinnate or mostly arising at petiole insertion, forming arching submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing with leaf. PEDUNCLE: subequal or longer than petiole, slender. SPATHE: slender, not distinctly constricted, tube very slender, persistent, tightly convolute around female flowers, blade longer, oblong-lanceolate, erect to reflexed and ± revolute at anthesis, marcescent. SPADIX: subequal or longer than spathe, slender, female zone adnate to spathe, few-flowered (2–7), ± biseriate, separated from male zone by short sterile zone, male zone subulate to cylindric, fertile to apex or with a few sterile flowers at apex or with a long, terminal appendix (H. appendiculata). FLOWERS: unisexual, perigone absent. MALE FLOWER: 3-androus, synandrium peltate, truncate, hexagonal, elongated in sense of spadix axis, shallow, stipitate, connective strongly dilated, thecae short, subglobose, remote, almost pendent from margin, dehiscing by pore. POLLEN: inaperturate, spherical, mediumsized (mean 40 µm.), exine rather densely spinose. STERILE FLOWER: lower ones very few, ± remote, apiculiform, upper ones very few, consisting of tiny, peltate synandrodes. FEMALE FLOWER: gynoecium oblong to lageniform, ovary 1locular, ovule 1, anatropous, funicle very short, placenta parietal to subbasal (morphologically basal), stylar region very short, stigma subcapitate. BERRY: ellipsoid to globose, style persistent, pericarp thin, 1-seeded, white. SEED: ellipsoid, testa smooth, very thin, embryo large, ellipsoid, light green, endosperm absent. See Plates 68, 121D. CHROMOSOMES: 2n = 26, 28. DISTRIBUTION: 7 spp.; tropical southeast Asia, Malay Archipelago:– Brunei, Burma, ?Cambodia, China (Yunnan), Laos, Malaysia (Borneo, Peninsula), Thailand, Vietnam. ECOLOGY: tropical humid forest; geophytes on forest floor or humus deposits on limestone or basalt rocks.

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68. Hapaline

ETYMOLOGY: modification of the feminine form of Greek hapalos (soft), referring to the small tender nature of this genus. TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), Li (1979), Bogner (1984a), Boyce (1996).

Tribe Nephthytideae

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Tribe Nephthytideae Engler in Engler & Prantl, Nat. Pflanzenfam. II (3): 112, 128 (1887). Laticifers simple, articulated; geophytic, stem tuberous to rhizomatous; petiole geniculate apically; leaf blade sagittate or trifid to trisect and highly divided, primary lateral veins of ultimate lobes or divisions pinnate, higher order venation reticulate; spathe boat-shaped or fully expanded, not clearly differentiated into tube and blade; spadix cylindric, female and male zones usually contiguous, sterile flowers ± absent (except Pseudohydrosme); flowers unisexual, perigone absent; stamens free with filaments lacking or very short (except Nephthytis); ovules 1 per locule, anatropous, placenta basal (except Pseudohydrosme); berries and seeds large, testa thin to absent, endosperm absent (fruit unknown in Pseudohydrosme). Chromosomes large to very large.

69. Nephthytis Nephthytis Schott in Oesterr. bot. Wochenbl. 7: 406 (1857). TYPE: N. afzelii Schott SYNONYM: Oligogynium Engler in Bot. Jahrb. 4: 64 (1883). HABIT: evergreen, rarely seasonally dormant, small to medium-sized herbs, rhizome creeping, subepigeal, rarely hypogeal, slender to thick, internodes very short, rarely long.

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Plate 68. Hapaline. A, habit × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium, top view × 10; E, synandrium, side view × 10; F, gynoecium × 15; G, gynoecium, longitudinal section × 15; H, infructescence × 2; J, habit × 2/3; K, leaf × 2/3; L, leaf × 2/3. Hapaline celatrix: A–H, Boyce 417 (Kew spirit collection 57283 & Kew slide collection); H. benthamiana: J, Kerr 614 (K); H. ellipticifolia: K, Sun s.n. (YUKU); H. colaniae: L, Pételot 2919 (K).

CALADIEAE : HAPALINE

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LEAVES: several, rarely solitary. PETIOLE: geniculate apically (geniculum very long in N. bintuluensis), usually smooth, rarely sparsely aculeate, uniformly green, sheath very short. BLADE: cordate-sagittate to sagittate-trifid or subtriangular, posterior divisions usually longer than anterior, all three divisions often long-acuminate; basal ribs usually well-developed, primary lateral veins of both anterior and posterior divisions pinnate forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, usually flowering with leaves, rarely without (N. bintuluensis). PEDUNCLE: usually relatively slender, shorter or equalling petiole. SPATHE: ovate- to oblong-elliptic, fully expanded, spreading to reflexed at anthesis, not constricted, decurrent on peduncle, green, persistent or marcescent. SPADIX: short- to long-stipitate, cylindric, female zone contiguous with male, rarely with a few sterile flowers in between, male zone longer than female, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4androus, stamens free, obpyramidal, filaments thick, well developed (except N. hallaei, N. mayombensis), connective thick, thecae lateral, dehiscing by apical pore. POLLEN: presented in amorphous mass, inaperturate, ellipsoid to subsphaeroidal, medium-sized (mean 46 µm.), exine verrucate. FEMALE FLOWER: ovary ovoid, 1-locular, ovule 1, anatropous, funicle very short, placenta basal, stylar region shortly attenuate or inconspicuous, stigma discoid-hemispheric. BERRY: large, subglobose to ellipsoid, orange, pericarp thick. SEED: obovoid or subglobose to ellipsoid, smooth, testa absent at maturity, embryo large, outer cell layers with chlorophyll, endosperm absent. See Plates 69, 122A. CHROMOSOMES: 2n = 36, 40, 60, karyology very similar to that of Anchomanes. DISTRIBUTION: 10 spp.; tropical west and central Africa:– ?Benin, Cameroon, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Ghana, Ivory Coast, Liberia, Malaysia (Sarawak), Nigeria, Sierra Leone, ?Togo. ECOLOGY: tropical humid forest; geophytes on forest floor. ETYMOLOGY: after mythic Nephthys, mother of Anubis and wife of Typhon.

69. Nephthytis

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TAXONOMIC ACCOUNTS: Bogner (1980c), Knecht (1983), Ntépé-Nyame (1988), Namur & Bogner (1994), Hay, Bogner & Boyce (1994).

70. Anchomanes Anchomanes Schott in Oesterr. bot. Wochenbl. 3: 314 (1853). TYPE: A. hookeri Schott, nom. illeg. (= Caladium petiolatum W.J. Hooker, A. petiolatus (W.J. Hooker) Hutchinson). HABIT: herbs, often very robust, tuber small to gigantic, erect or grossly rhizomatous, seasonally dormant. LEAF: solitary, often gigantic. PETIOLE: very long, terete, aculeate, rarely smooth, sheath very short. BLADE: sagittate when juvenile, becoming dracontioid at maturity: i.e. trisect, primary divisions each further divided ± dichotomously or pinnately, secondary divisions irregularly pinnatifid, ultimate lobes very variable in size and shape, distal ones larger, trapezoid, apically broader, truncate or shallowly bifid, decurrent to sessile, proximal lobes ovate and acuminate; primary lateral veins of ultimate lobes pinnate, long-arcuate, mostly running into margin, sometimes forming irregular submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, usually appearing before leaf. PEDUNCLE: aculeate, rarely smooth, shorter than petiole. SPATHE: erect, broadly ovate to narrowly oblong-lanceolate or oblong-elliptic, boat-shaped, not constricted, convolute basally or not at all, apex sometimes fornicate, marcescent. SPADIX: much shorter or subequal to spathe, sessile to subsessile, cylindric, female zone subequal to male zone or much shorter, male zone contiguous with female, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: stamens free, anthers sessile or filaments short, compressed, connective slender below, thickened and dilated apically, thecae ovate-oblong, opposite, dehiscing by apical slit. POLLEN: extruded in strands, inaperturate, ellipsoid to ellipsoid-oblong, large (mean 64 µm., range 37–94

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Plate 69. Nephthytis. A, habit × 1/2; B, stamen × 10; C, stamen, longitudinal section × 10; D, gynoecium × 10; E, gynoecium, longitudinal section × 10; F, habit × 1/2; G, leaf × 1/2; H, spadix × 1; J, stamen × 10; K, stamen, longitudinal section × 10; L, gynoecium × 10; M, gynoecium, longitudinal section × 10; N, habit × 1/2; P, habit × 1/5; Q, inflorescence × 1; R, stamen × 10; S, stamen, longitudinal section × 10; T, gynoecium × 10; U, gynoecium, longitudinal section × 10; V, infructescence × 1; W, seed, hilum view × 1. Nephthytis hallaei: A, Bogner 750 (K & Kew spirit collection 46624); B–E, Bogner 750 (Kew spirit collection 54219); N. swainei: F, Swaine et al. GC 44621 (K); N. poissonii: G, Bogner 605 (K); H–M, Cult. Kew 1957–43801 (Kew spirit collection 51366); N. constricta: N, Brenan 8527 (K); N. afzelii: P, V–W, Knecht 6 (Cult. Kew 1982–4608); Q, Knecht 6 (Kew spirit collection 46572); R–U, Knecht 6 (Kew spirit collection 46103).

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E F

DD D

R P

EE C

K

FF AA H

Q

J

CC BB

M

S T W X L

N

G A

B

Z

Y

V

U

Plate 70. Anchomanes. A, habit in flower × 1/10; B, habit in fruit × 1/10; C, habit × 1/20; D, part of mature leaf × 2/3; E, seedling leaf × 1/3; F, juvenile leaf × 1/3; G, inflorescence × 2/3; H, stamen, top view × 6; J, stamen, longitudinal section × 6; K, gynoecium, longitudinal section × 6; L, infructescence × 1/2; M, seed × 2; N, inflorescence × 2/3; P, stamen, top view × 6; Q, stamen, longitudinal section × 6; R, gynoecium × 6; S, inflorescence × 2/3; T, stamen, top view × 6; U, stamen, longitudinal section × 6; V, gynoecium × 6; W, inflorescence × 2/3; X, stamen, top view × 6; Y, stamen, longitudinal section × 6; Z, gynoecium × 6; AA, stamen, top view × 6; BB, stamen, longitudinal section × 6; CC, gynoecium × 6; DD, stamen, top view × 6; EE, stamen, longitudinal section × 6; FF, gynoecium × 6. Anchomanes welwitschii: A, Bullock 1215 (Kew photograph collection); B, Cult. Kew 14/12/59 (Kew slide collection); A. difformis: C, Croat s.n. (Kew photograph collection); D, Lowe 2116 (K); A. welwitschii: E–F, Fanshawe 8549 (K); A. difformis: G–K, Cult. Kew 1961–38601 (Kew spirit collection 29047.636); L–M, Milne-Redhead 2676 (Kew spirit collection 18904); A. nigritianus: N–R, Cult. Kew 1976.00789 Bogner 640 (Kew slide collection & Kew spirit collection 29047.1); Bogner 662 (Kew spirit collection 37350); A. abbreviatus: S–V, Cult. Kew 1951–556 Greenway s.n. (Kew spirit collection 21975); Cult. Kew 1952.51801 Faulkner 878 (Kew slide collection); A. boehmii: W–Z, Bally 7558 (K & Kew spirit collection 56560); Shabani 11 (K); A. petiolatus: AA–CC, Boughey 163 (Kew spirit collection 16869); A. welwitschii: DD–FF, Cult. Kew (Kew spirit collection 22258).

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70. Anchomanes

µm.), exine psilate or obscurely verruculate, very thin. FEMALE FLOWER: ovary 1-locular, ovule 1, erect, anatropous, funicle very short, placenta basal, stylar region shortly conic or absent, sometimes strongly deflexed towards spadix base, stigma either 2-lobed and reniform to V-shaped or discoid or depressed-globose. BERRY: large, oblong-ellipsoid, fleshy, borne in cylindric spike, red, purplish or partly white. SEED: obovoid to oblong-ovoid, testa very thin, smooth, transparent, embryo large, green, endosperm absent. See Plates 70, 122B. CHROMOSOMES: 2n = 40, very large. DISTRIBUTION: 7–8 spp.; tropical Africa: Angola, Benin, Burkina Faso, ?Cabinda, Cameroon, Central African Republic, Chad, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Gambia, Ghana, ?Guinea, ?Guinea-Bissau, Ivory Coast, Kenya, Liberia, ?Malawi, Mozambique, Nigeria, Senegal, Sierra Leone, Sudan, Tanzania, Togo, Uganda, Zaïre, Zambia. ECOLOGY: tropical humid forests, savannas (A. welwitschii), near swamp (A. boehmii); geophytes, in leaf litter between rocks or on forest floor, seasonally dormant. NOTES. The only clear cut distinction between Anchomanes and Pseudohydrosme is locule number. ETYMOLOGY: Greek agchein (to strangle) and mainesthai (to rage); old plant name by Appuleius. TAXONOMIC ACCOUNTS: Engler (1911), Knecht (1983), Mayo & Bogner in Mayo (1985a), Ntépé-Nyame (1988).

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71. Pseudohydrosme Pseudohydrosme Engler in Bot. Jahrb. 15: 455 (1892, “1893”). LECTOTYPE: P. gabunensis Engler (see N.E. Brown in Thiselton-Dyer, Fl. Trop. Africa 8: 160. 1901, “1902”). SYNONYM: Zyganthera N.E. Brown in Thiselton-Dyer, Fl. Trop. Africa 8: 160 (1901, “1902”). HABIT: large, seasonally dormant herbs, tuber subglobose, subterranean, with annular leaf scars, growing continuously and not renewed with each growing period. LEAF: solitary, large. PETIOLE: long, aculeate, sheath very short. BLADE: dracontioid: i.e. trisect, primary divisions pinnatifid or bip-

71. Pseudohydrosme

innatifid to pinnatisect, ultimate (distal) lobes mostly truncate to shallowly bifid, sessile to decurrent, proximal lobes acuteacuminate; primary lateral veins of ultimate lobes pinnate, forming irregular submarginal collective vein or running into margin, higher order venation reticulate. INFLORESCENCE: solitary, appearing before leaf. PEDUNCLE: aculeate, very short, much shorter than petiole. SPATHE: large, somewhat resembling the horn of a euphonium, unconstricted, bright yellow, dark purple within and at mouth of tube, tube convolute, fleshy, obconic, blade very broad, thinner, with flaring, auriculate margins, fornicate. SPADIX: very short, sessile, female zone subcylindric, male zone cylindric, obtuse, subequal to or longer than female, fertile to apex (P. gabunensis) or with appendix covered with sterile flowers (P. buettneri). FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–5-androus, stamens free, subprismatic, compressed, anthers sessile, connective thick, broad, overtopping thecae, thecae oblong, long, lateral, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, ellipsoid-oblong, very large (mean 106 µm., range 93–114 µm.), exine psilate to slightly scabrous. STERILE MALE FLOWERS: composed of subprismatic, free staminodes. FEMALE FLOWER: ovary globose to broadly ellipsoid, sometimes compressed, 2(–3)-locular, ovules 1 per locule, anatropous, funicle short, placenta axile at base of septum, stylar region attenuate to cylindric, narrower than ovary, stigma thick, 2(–3)-lobed, concave centrally. BERRY: unknown. SEED: unknown. See Plates 71, 122C. CHROMOSOMES: 2n = ca. 40. DISTRIBUTION: 2 spp.; tropical Africa:– Gabon. ECOLOGY: tropical humid forest; geophytes on forest floor. NOTES: Engler (1911) recognized 2 sections:– sect. Pseudohydrosme (“Chorianthera”), sect. Zyganthera. Pseudohydrosme buettneri (sect. Zyganthera) has never been recollected and the type no longer exists. N.E. Brown considered this species to be sufficiently different to warrant generic recognition. ETYMOLOGY: Greek pseudo (false) and Hydrosme, itself derived from hydra (water snake) and osmê (smell); in reference to its similarity to Hydrosme (now Amorphophophallus). TAXONOMIC ACCOUNTS: Engler (1911), Bogner (1981b).

NEPHTHYTIDEAE : PSEUDOHYDROSME

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F

G

E H

A

J

C

B D

Plate 71. Pseudohydrosme. A, habit × 1/10; B, portion of leaf × 1/2; C, section of petiole × 2/3; D, habit in flower × 1/2; E, spadix × 2/3; F, stamen × 6; G, stamen, longitudinal section × 6; H, gynoecium × 6; J, gynoecium, longitudinal section × 6. Pseudohydrosme gabunensis: A, Aroideana 4(1): 32, fig.2 (1981); B–D, Bogner 664 (K); Aroideana 4(1): 33, fig.6, 34, fig.8 (1981); E–J, Bogner 664 (Kew spirit collection 45184).

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C

Tribe Aglaonemateae Tribe Aglaonemateae Engler in Nova Acta Acad. Leopold.Carol. 39: 148 (1876, “Aglaonemeae”). Laticifers simple, articulated; leaf blade (linear-) ovate to elliptic-oblong, base sometimes subcordate, primary lateral veins pinnate, forming single marginal vein, higher order venation parallel-pinnate; spathe boat-shaped, unconstricted; spadix with contiguous female and male zones; flowers unisexual, perigone absent; stamens free, thecae dehiscing by apical pore, pollen grains large, ovary 1-locular, ovule 1, anatropous, style short, narrower than ovary, stigma broad, discoid, concave; berry large; seed large, ellipsoid, testa smooth, thin, endosperm absent.

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72. Aglaonema Aglaonema Schott in Wiener Z. Kunst 1829 (3): 892 (1829). TYPE: A. oblongifolium Schott, nom. illeg. (Arum integrifolium Link, Aglaonema integrifolium (Link) Schott). HABIT: evergreen herbs, sometimes robust, stem epigeal, erect and unbranched or creeping and often branched, internodes green, smooth, often rooting at the nodes. LEAVES: several, forming an apical crown. PETIOLE: sheath usually long. BLADE: ovate-elliptic, narrowly elliptic, rarely broadly ovate or sublinear, base often unequal, attenuate to rounded, rarely cordate, often with striking, silvery and pale green patterns of leaf variegation; primary lateral veins pinnate, often weakly differentiated, running into marginal vein, higher order venation parallel-pinnate. INFLORESCENCE: 1–9 in each floral sympodium. PEDUNCLE: shorter or longer than petiole, deflexing in fruit. SPATHE: ovate to ± globose, erect, boat-shaped to convolute, not differenti-

ated into tube and blade, often apiculate, green to whitish, slightly to strongly decurrent, marcescent. SPADIX: cylindric to clavate, shorter or longer than spathe, stipe long to almost absent, female zone rather few-flowered, contiguous with and much shorter than male zone, male zone fertile to apex; rarely with staminodes. FLOWERS: unisexual, perigone absent. MALE FLOWER: stamens free, not forming clear floral groups, filaments usually distinct, connective thickened, thecae opposite, obovoid, short, dehiscing by apical pore or reniform transverse slit. POLLEN: inaperturate, ellipsoid, large (mean 52 µm.), range 37–67 µm.), exine essentially psilate. FEMALE FLOWER: ovary subglobose, 1-locular, ovule 1, anatropous, shortly ovoid, funicle very short, placenta basal, stylar region short, thick, stigma broad, discoid, concave centrally. BERRY: ellipsoid, outer layer fleshy green but turning yellow, rarely white and finally red. SEED: ellipsoid, almost as large as berry, testa thin, ± smooth, tegmen inconspicuous, embryo large, endosperm absent. See Plates 72, 122D. CHROMOSOMES: 2n = 40, 60, 80, 100, 120 (70, 110). DISTRIBUTION: 21 spp.; tropical Asia, Malay Archipelago, Papuasia:– Bangladesh, Brunei, Burma, Cambodia, China (Guandong, Guangxi, Hainan, Yunnan), India (NE), Indonesia (Borneo, Java, Moluccas, Sulawesi, Sumatra), Laos, Malaysia (Borneo, Peninsula), Papua New Guinea, Philippines, Thailand, Vietnam. ECOLOGY: tropical humid forest; terrestrial on forest floor, occasionally in deciduous forest or regrowth, also in humus deposits on limestone and in peat deposits. COMMON NAMES AND USES: chinese evergreen, very widely cultivated as an ornamental plant. ETYMOLOGY: Greek aglaos (splendid) and nema, nêmatos (stamen). TAXONOMIC ACCOUNTS: Engler (1915), Nicolson (1969), Jervis (1980).

72. Aglaonema

AGLAONEMATEAE : AGLAONEMA

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E

C

D

F

B

J

H

G

A

K

L

Plate 72. Aglaonema. A, habit × 1/4; B, leaf × 1/3; C, detail of leaf venation × 4; D, spadix × 1; E, stamen × 10; F, inflorescence × 1; G, staminode × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, habit × 1/4; L, inflorescence × 1. Aglaonema simplex : A–C, Cult. Kew 1963–43405; D–E, Giles & Woolliams PB109 (Kew spirit collection 37985); A. simplex : F–J, Hay 2001 (Kew spirit collection 59067); A. modestum: K, Cult. Kew 1968–38227; A. hookerianum: L, Cult. Kew (Kew spirit collection 22425).

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C

73. Aglaodorum

Tribe Culcasieae

C

Aglaodorum Schott, Gen. Aroid. t. 58 (1858). TYPE: A. griffithii (Schott) Schott (Aglaonema griffithii Schott).

Tribe Culcasieae Engler in Engler & Prantl, Nat. Pflanzenfam. II (3): 112, 116 (1887).

HABIT: evergreen herb, stem rhizomatous, creeping, aerenchymatous stem, much-branched, . LEAVES: several to many. PETIOLE: terete, sheath short. BLADE: oblanceolate to oblong, thick; midrib very thick, primary lateral veins pinnate, weakly differentiated, higher order venation parallel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: long, subequal to petiole, erect in fruit. SPATHE: oblong, cuspidate, convolute, erect. SPADIX: stipitate, a little shorter than spathe, female zone very short with a single whorl of flowers, male zone subcylindric, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: stamens free, short, prismatic, filaments distinct, connective thickened, thecae oblong, adjacent or opposite, dehiscing by apical pore. POLLEN: inaperturate, ellipsoid, large (mean 59 µm.), exine apparently psilate or verruculate. FEMALE FLOWER: gynoecium surrounded by whorl of 1–3 shorter prismatic staminodes, ovary 1–2-locular, ovule 1 per locule, anatropous, funicle very short, placenta parietal in 1-locular ovaries, stigma broad, discoid, 4-lobed, concave at centre. BERRY: obovoid to ellipsoid, large, pericarp very thick, green. SEED: large, ellipsoid, testa thin, smooth, embryo large, plumule well-developed, endosperm absent. See Plates 73, 123A. CHROMOSOMES: 2n = 40. DISTRIBUTION: 1 sp.; Indonesia (Sumatra), Malaysia (Borneo, Peninsula), Vietnam. ECOLOGY: tropical open swamps, especially freshwater tidal zones and brackish water, often growing with Nypa fruticans and Cryptocoryne ciliata; creeping rhizomatous helophyte. ETYMOLOGY: Greek aglaos (splendid) and dôron (gift), a poetic name. TAXONOMIC ACCOUNTS: Engler (1915), Nicolson (1969).

Laticifers present, simple, articulated (Cercestis) or absent (Culcasia), resin canals present in roots, stems and leaves, sclerotic hypodermis present in roots; usually climbing hemiepiphytes with slender stems (rarely robust), sometimes erect or prostrate terrestrial plants; petiole geniculate apically; primary lateral veins pinnate, higher order venation reticulate; female zone of spadix contiguous with male or separated by zone of sterile male flowers; flowers unisexual, perigone absent; male flower 2–4 androus, stamens free, anthers subsessile, lacking endothecial thickenings, connective usually strongly thickened, broad, truncate, thecae dehiscing by short apical slit or pore; ovules 1 per locule, stigma sessile, large, discoid; berries ± globose, usually red; seed testa thin, endosperm absent.

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74. Culcasia Culcasia Palisot de Beauvois, Fl. Oware 1: 3 (1805), nom. cons. TYPE: C. scandens Palisot de Beauvois (typ. cons.). SYNONYM: Denhamia Schott in Schott & Endlicher, Melet. Bot. 19 (1832). Laticifers absent. HABIT: erect, repent or climbing herbs, rooting at least from lower nodes, branches slender. LEAVES: many, often forming terminal crown in terrestrial species. PETIOLE: geniculate apically, geniculum often inconspicuous, sheath persistent, rather long. BLADE: lanceolate or oblanceolate to ovate or ovate-oblong, rarely rounded, rarely pubescent below, resin canals pellucid, linear or punctate; primary lateral veins pinnate, often forming submarginal collective vein, otherwise running into marginal vein, higher order venation reticulate.

73. Aglaodorum

CULCASIEAE : CULCASIA

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F

E

B

D

A G

C

Plate 73. Aglaodorum. A, habit × 2/3; B, inflorescence × 1; C, spadix × 2; D, stamen × 8; E, gynoecium with associated staminode × 8; F, gynoecium, longitudianal section × 8; G, fruit × 1. Aglaodorum griffithii: A, C–G, Bogner 1767 (M); Bogner 1672 (Kew spirit collection 49830); B, Boyce (Kew slide collection).

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A

B K

T C

M

L

F

H

G

P

N

J R

S

Q D

E

Plate 74. Culcasia. A, habit × 1/3; B, juvenile habit × 1/3; C, habit × 1/3; D, leaf × 1/3; E, habit × 1/3; F, inflorescence × 1; G, inflorescence × 1; H, stamens × 10; J, gynoecium, longitudinal section × 10; K, spadix × 5; L, detail of stamens, top view × 4; M, detail of gynoecia × 4; N, stamens × 10; P, gynoecium, longitudinal section × 10; Q, habit × 2/3; R, stamens × 10; S, gynoecium, longitudinal section × 10; T, seed × 1 1/2. Culcasia parviflora: A, Keay IFH 37529 (K); B, Bogner 697 (K); C. angolensis: C, Johnson s.n. (K); Robyns 1140 (K); C. panduriformis: D, Zenker 4572 (K); C. longevaginata: E, Baldwin 9226 (K); C. striolata: F, Milne-Redhead 5155 (Kew spirit collection 22442); C. orientalis: G–J, Bogner 140 (Kew spirit collection 22712); C. seretii: K–M, Cult. Kew 1982–04604 (Kew spirit collection 46485); C. saxatilis: N–P, Meikle 1275 (Kew spirit collection 25627); C. rotundifolia: Q–S, Bogner 749 (K & Kew spirit collection 56676); C. liberica: T, Cult. Kew 1982–04601 (Kew spirit collection 51491).

CULCASIEAE : CULCASIA

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INFLORESCENCE: 1–12(–20) in each floral sympodium, internodes of floral sympodium sometimes relatively elongated. PEDUNCLE: short to relatively long. SPATHE: erect, green to whitish, boat-shaped, not or hardly constricted, convolute basally, gaping apically at anthesis, deciduous to marcescent. SPADIX: subsessile to stipitate, cylindric-clavate, equal to or somewhat longer than spathe, female zone usually densely flowered, rarely laxly, shorter than male zone and either contiguous with it or separated by a laxly flowered zone of sterile male flowers, male zone fertile to apex, axis often persistent in fruit. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4-androus, stamens free, short, obpyramidal, truncate apically, anthers subsessile, connective thick, thecae oblong, dehiscing by short apical slit. POLLEN: inaperturate, subspheroidal, medium-sized (mean 32 µm., range 27–40 µm.), exine verrucate to rugulate or subreticulate, usually with psilate patches of variable size, rarely spinose. STERILE MALE FLOWERS: with 3–4 obpyramidal, depressed staminodia. FEMALE FLOWER: ovary depressed, 1–3-locular, ovules 1 per locule, anatropous, funicle short, placentae subbasal, stigma sessile, hemispheric-discoid, relatively large, sometimes weakly lobed. BERRY: globose to ellipsoid, 1–3-seeded, mostly red, sometimes orange to greenish-yellow, infructescence subglobose to cylindric. SEED: ovoid to ellipsoid, testa thin, smooth, brown, plumule lateral, superficial, with leaf primordia (C.liberica), endosperm absent. See Plates 74, 123B. CHROMOSOMES: 2n = 42, 84. DISTRIBUTION: ca. 27 spp.; tropical Africa:– Angola, Benin, Burkina Faso, ?Burundi, Cabinda, Cameroon, Central African Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, ?Gambia, Ghana, Guinea, ?Guinea-Bissau, Ivory Coast, Kenya, Liberia, Mali, Niger, Nigeria, ?Rwanda, Senegal, Sierra Leone, Tanzania, Togo, Uganda, Zaïre. ECOLOGY: tropical moist and humid forest; usually climbing hemiepiphytes, sometimes terrestrial, terrestrial species tend to occur in damp forest sites. ETYMOLOGY: variant of Middle Eastern name qolqas, (also the origin of the name Colocasia). TAXONOMIC ACCOUNTS: Engler (1905), Bogner (1980b), Knecht (1983), Mayo (1985a), Ntépé-Nyame (1988), Boyce (1995b).

74. Culcasia

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75. Cercestis Cercestis Schott in Oesterr. bot. Wochenbl. 7: 414 (1857). TYPE: C. afzelii Schott SYNONYMS: Alocasiophyllum Engler in Bot. Jahrb. 15: 449 (1892, “1893”); Rhektophyllum N.E. Brown in J. Bot. 20: 194 (1882). Laticifers present, simple, articulated. HABIT: climbing herbs, sometimes creeping, small to robust, stem long, producing flagelliform shoots with long internodes and bearing only cataphylls, followed by thicker, flowering stems with several foliage leaves and short internodes. LEAVES: many. PETIOLE: geniculate apically, geniculum sometimes inconspicuous, sheath short to long. BLADE: oblong-lanceolate or oblong to cordate, sagittate, or hastate, or trifid with acuminate segments, sometimes laciniate-pinnatifid with slit-like perforations extending to margin forming large rhomboid to obtriangular lobes (C. camerunensis, C. mirabilis), resin canals pellucid, linear or punctate; primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: usually 1–4 in each floral sympodium (up to 16 in C. camerunensis). PEDUNCLE: shorter than spathe. SPATHE: erect, thick, boat-shaped to subcylindric, convolute basally into a tube, gaping apically at anthesis, not or hardly constricted, persistent to marcescent. SPADIX: sessile, shorter than spathe, female zone shorter and contiguous with male. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4androus, stamens free, prismatic, slightly narrowed basally, usually short and relatively broad, sometimes rather long and narrow (C. camerunensis, C. mirabilis), filaments very short, connective broad, thecae shortly oblong to almost linear, dehiscing by small apical slit. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong or spherical to subspheroidal, medium-sized (mean ca. 40 µm., range 36–55 µm.), exine obscurely fossulate-verrucate, or almost perfectly psilate (C. camerunensis, C. mirabilis). FEMALE FLOWER: ovary obovoid, 1-locular, ovule 1, hemianatropous to anatropous, rarely campylotropous (C. taiensis), funicle very short, placenta parietal to subbasal, stigma sessile to subsessile, often rather broad, discoid, often somewhat concave centrally.

75. Cercestis

C

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J

L

N

M

K

B

Q

A

P

C

H

F

G

D

E

Plate 75. Cercestis. A, flowering shoot arising from leafless continuation shoot, attached to twig by aerial roots × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, inflorescences with associated stem and petiole base × 1/2; E, male flower × 6; F, gynoecium × 6; G, gynoecium, longitudinal section × 6; H, spadix, J, detail of male zone of spadix × 6; K, detail of female zone of spadix × 6; L, male flower × 9; M, gynoecium × 9; N, gynoecium, longitudinal section × 9; P, habit × 1/2; Q, fruiting habit × 1/2. Cercestis stigmaticus: A, Coombe 169 (K); C. kamerunianus: B, Jones 19526 (K); C. mirabilis: C, Bogner 601 (K); D, Bogner 705 (K); E–G, Bogner s.n. (K & Kew spirit collection 29047.471); C. dinklagei: H–N, Hepper 7508 (Kew spirit collection 49930); C. afzelii: P, Jones & Onochie FHI 18753 (K); Q, Adams 3729 (K).

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BERRY: infructescence ellipsoid to oblong, berries obovoid to subglobose or ellipsoid, red, pericarp thick. SEED: obovoid to ellipsoid, testa smooth, thin, embryo large, outer cell layer green, endosperm absent. See Plates 75, 123C. CHROMOSOMES: 2n = 42, morphologically similar to those of Culcasia. DISTRIBUTION: 13 spp.; tropical Africa:– Angola, Benin, ?Burundi, ?Cabinda, Cameroon, Central African Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Nigeria, ?Rwanda, Senegal, Sierra Leone, ?Togo, Uganda, Zaïre. ECOLOGY: tropical humid forest; climbing hemiepiphytes and terrestrial plants. ETYMOLOGY: Greek Cercestes, one of the fifty sons of mythic Aegyptus. TAXONOMIC ACCOUNTS: Engler (1911), Knecht (1983), Mayo (1985a), Bogner (1986a), Ntépé-Nyame (1988), Bogner & Knecht (1995).

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Tribe Montrichardieae Tribe Montrichardieae Engler in Nova Acta Acad. Leopold.Carol. 39: 144 (1876). Laticifers simple, articulated, sclerotic hypodermis present in roots; arborescent helophytes; leaf blade cordate-sagittate to trifid, primary lateral veins pinnate, higher order venation ± reticulate; spathe deciduous after anthesis, slightly constricted, differentiated into lower tube and upper, ± boat-shaped blade; spadix: female and male zone contiguous, sterile flowers absent; flowers unisexual, perigone absent; male flower 3–6androus, stamens free, ± sessile, connective overtopping thecae, thecae dehiscing by short, apical slit, pollen large; ovary 1-locular, ovules 1–2, anatropous, placenta subbasal, style prismatic, truncate, longer than ovary, ± excavated apically; berry and seed large, plumule well-developed, endosperm absent.

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76. Montrichardia Montrichardia H. Crüger in Bot. Zeitung (Berlin) 12: 25 (1854), nom. cons. TYPE: M. aculeata (G.F.W. Meyer) Schott (Caladium aculeatum G.F.W. Meyer). SYNONYM: Pleurospa Rafinesque, Fl. Tell. 4: 8 (1838, “1836”). HABIT: robust to sometimes gigantic arborescent evergreen herbs, stem erect, multiplying at base from hypogeal rhizomes, internodes well-developed, smooth or aculeate, ± slender to massive. LEAVES: several, borne in terminal crown. PETIOLE: sheath half as long as petiole or longer, with free apical ligule. BLADE: cordate-sagittate to sagittate, hastate to trifid, rarely trisect, posterior divisions often longer than anterior division; basal ribs well-developed, primary lateral veins pinnate, running into marginal vein, secondary laterals ± parallel-pinnate, higher order venation reticulate. INFLORESCENCE: 1(–2) in each floral sympodium. PEDUNCLE: shorter than petiole. SPATHE: erect, thick, entirely deciduous after anthesis, ± constricted between tube and blade, tube convolute, blade longer than tube, widely gaping at anthesis, ± boat-shaped. SPADIX: sessile, erect, subequal to spathe, female zone cylindric, male zone subconoid, contiguous with and much longer than female zone, fertile to apex. FLOWERS: unisexual, perigone absent. MALE

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FLOWER: 3–6-androus, stamens free, obpyramidal-prismatic, truncate at apex, anthers sessile, connective thick, overtopping thecae, thecae oblong-ellipsoid, dehiscing by longitudinal slit. POLLEN: inaperturate, spherical to subspheroidal, large (mean 94 µm., range 92–96 µm.), exine psilate. FEMALE FLOWER: gynoecia prismatic-cylindric, ovary 1-locular, ovules 1–2, anatropous, funicle short, placenta subbasal to basal, stylar region prismatic, thick, excavated and rugulose at apex, stigma small, on low central boss, irregular to elliptic or roundish. BERRIES: free, large, subcylindric, somewhat compressed, pericarp spongiose, at apex excavated and radiately furrowed, 1-seeded. SEED: large, obovoid to ellipsoid, testa smooth or rough, brown, embryo large, endosperm absent. See Plate 76, 123D. CHROMOSOMES: 2n = 48. DISTRIBUTION: 2 spp.; tropical America, West Indies:– Belize, Brazil (Amazonia, Northeast, South East), Colombia, Costa Rica, French Guiana, Guatemala, Guyana, Honduras, Lesser Antilles, Nicaragua, Panama, Peru, Puerto Rico, Surinam, Trinidad, Tobago, Venezuela. ECOLOGY: tropical humid forest; helophytes in tranquil freshwater habitats, forming dense, often extensive stands along river margins. ETYMOLOGY: named after G. de Montrichard. TAXONOMIC ACCOUNTS: Engler (1911), Jonker-Verhoef & Jonker (1953), Lins (1994).

Tribe Zantedeschieae Tribe Zantedeschieae Engler in Engler & Prantl, Nat. Pflanzenfam. II (3): 113, 136 (1887). Laticifers simple, articulated; stem a depressed-globose tuber or short rhizome (Z. aethiopica); leaf blade lanceolate to hastate-sagittate, primary lateral veins pinnate, forming single marginal vein, higher order venation parallel-pinnate; peduncle long, sometimes longer than leaves; spathe ± obconic,

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Plate 76. Montrichardia. A, habit × 1/10; B, flowering shoot × 1/2; C, base of stem showing rhizome × 1/2; D, mid–portion of stem × 1/2; E, portion of stem showing prickles × 1/2; F, spadix × 1/2; G, stamens, top view × 3; H, stamen, side view × 5; J, gynoecium, top view showing irregular surface with stigma × 5; K, gynoecium, side view × 5; L, gynoecium, top view showing irregular surface with stigma × 3; M, gynoecium, longitudinal section × 5; N, infructescence × 1/2; P, seed × 2; Q, leaf × 1/2; R, leaf × 1/2. Montrichardia linifera: A, Mayo s.n. (Kew slide collection); M. arborescens: B, Philcox 8032 (K); Philcox et al. 8413 (Kew spirit collection 45821); C–E, Bogner 2188 (Kew spirit collection 29047.760 & Kew illustration collection); Nelson 1729 (K); F, Simmons s.n. (K & Kew spirit collection 58906); G-M, Bogner 2188 (Kew spirit collection 29047.760); M. linifera: N–P, Stannard & Arrais (Kew spirit collection 51643 & 51644); Q, Harley et al. 24747 (K); M. arborescens R, Nelson 1733 (K).

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unconstricted, tube subcylindric-obconic, blade gaping widely; spadix shorter than spathe, male and female zones contiguous; flowers unisexual, perigone absent; stamens free, connective thickened apically, thecae dehiscing by apical pore, pollen extruded in strands; ovary 1–5 locular, ovules (1–)4(–8) per locule, anatropous, placenta axile to subapical; berry green, orange or yellow; seed testa striate, endosperm copious.

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77. Zantedeschia Zantedeschia K. Sprengel, Syst. Veg. 3: 756, 765 (1826), nom. cons. TYPE: Z. aethiopica (L.) K. Sprengel (Calla aethiopica L.), typ. cons. SYNONYMS: ? Aroides Heister ex Fabricius, Enum., ed. 2, 42 (1763); Houttinia Necker, Elementa Botanica 3: 291 (1790); Colocasia Link, Diss. Bot. 77 (1795), nom. rej.; Richardia Kunth in Mém. Mus. Hist. Nat. 4: 433, 437 (1818, non L. 1753); Otosma Rafinesque, Fl. Tell. 4: 8 (1838, “1836”); [Arodes O.Kuntze, Rev. Gen. 2: 739 (1891), orth. var.]; Pseudohomalomena A.D. Hawkes in Madroño 11: 147 (1951). HABIT: seasonally dormant, sometimes evergreen (Z. aethiopica) herbs, tuber depressed-globose, or with thick rhizome (Z. aethiopica). LEAVES: several, radical. PETIOLE: spongiose, sheath long. BLADE: lanceolate, narrowly elliptic, cordate, cordate-sagittate, sagittate, hastate, often variegated by aggregations of pale diaphanous necrotic patches; primary lateral veins pinnate, running into distinct marginal vein, secondary and tertiary laterals parallel-pinnate, higher order venation transverse-reticulate. INFLORESCENCE: solitary in each floral sympodium, appearing with leaves. PEDUNCLE: long, subequal or longer than petiole. SPATHE: unconstricted, persistent, pure white, cream, yellow, pink or rosy purple, lower part convolute forming a short, stout, subcylindric to obconic tube, often dark purple within at base, upper part widely gaping, shorter than or subequal to tube, suberect or recurved with somewhat revolute margins. SPADIX: erect, digitiform,

shorter than spathe, sessile or stipitate, male and female zones contiguous, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–3-androus, stamens free, anthers oblong, somewhat compressed, subsessile, connective truncate at apex, separating thecae, thecae opposite, oblong, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, medium-sized (mean 40 µm., range 31–49 µm.), exine perfectly psilate to obscurely dimpled. FEMALE FLOWER: either a naked gynoecium or rarely (Z. aethiopica, Z. odorata) surrounded by whorl of ca. 3 spathulate to clavate and apically truncate staminodes, ovary ovoid, 1–5-locular, ovules (1–)4(–8) per locule, anatropous, placenta axile to subapical, stylar region short- to long-attenuate, stigma rather small, discoid-subcapitate. BERRY: obovoid or depressed-globose, 1–several-seeded, usually green, or orange, rarely yellow, and mucilaginous within (Z. aethiopica). SEED: ovoid to ellipsoid, strophiolate, testa costate, embryo axile, elongate, endosperm copious. See Plates 77, 124A. CHROMOSOMES: 2n = 32. DISTRIBUTION: 8 spp.; southern Africa:– Angola, Botswana, Lesotho, Malawi, ?Mozambique, Namibia, South Africa, Swaziland, Tanzania, ?Zaire, Zambia, Zimbabwe. ECOLOGY: tropical and subtropical damp open habitats; helophytes, seasonally wet places, along streams, swampy ground along forest edges, dry rocky hills. NOTES: Zantedeschia aethiopica is morphologically distinct from the other species; it is naturalized in various parts of tropical America, in southern Europe, the Philippines, New Zealand and elsewhere. ETYMOLOGY: named after G. Zantedeschi (1773–1846). TAXONOMIC ACCOUNTS: Engler (1915), Letty (1973), Perry (1989), Singh, Wyk & Baijnath (1996).

Tribe Callopsideae

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Tribe Callopsideae Engler in Engler & Prantl, Nat. Pflanzenfam., Nachtr. 3: 29, 34 (1906). Laticifers simple, articulated; small, evergreen herbs, rhizome slender, hypogeal; leaf blade cordate-ovate, primary lateral veins forming single marginal vein, higher order venation reticulate; spathe completely expanded at anthesis, persistent; spadix fertile to apex; female zone entirely adnate to spathe, contiguous with male zone or with short naked zone in between; flowers unisexual, perigone absent; stamens free, anthers sessile, thecae opposite, dehiscing by apical pore; gynoecia few, slender, ovary 1-locular, ovule 1, anatropous, placenta basal, style long-attenuate, stigma small; embryo elongate, endosperm copious.

78. Callopsis Callopsis Engler in Notizbl. Königl. Bot. Gart. Berlin 1: 27 (1895). TYPE: C. volkensii Engler

77. Zantedeschia

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Laticifers simple, articulated. HABIT: small evergreen herb, stem a slender hypogeal rhizome, internodes very short. LEAVES: usually minutely hispid on petiole and abaxial surface of blade. PETIOLE: sheath very short. BLADE: cordate-ovate; primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 2 in each floral sympodium, appearing with the

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Plate 77. Zantedeschia. A, habit × 1/3; B, leaf × 2/3; C, inflorescence × 2/3; D, spadix × 2; E, stamens, top view × 9; F, stamen × 9; G, gynoecium × 9, H, gynoecium, longitudinal section × 9; J, infructescence × 2/3; K, leaf × 2/3; L, gynoecium with associated staminodes × 9. Zantedeschia albomaculata subsp. albomaculata: A, Scheepers 36 (K); B, Codd 10688 (K); C–H, Milne-Redhead 3812 (Kew spirit collection 199162); J, Prosser 1827 (K); Z. rehmannii: K, Cult. Kew 1958–11001 (K); Z. aethiopica: L, Cult. Kew (Kew spirit collection 7467).

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Plate 78. Callopsis. A, habit with spathe of infructescence flattened out by pressing × 1; B, inflorescence, nearside and upper part of spathe removed × 5; C, gynoecium, longitudinal section × 20; D, stamen × 20; E, berry, side view with hilum at top × 4; F, seed, side view × 4. Callopsis volkensii: A, Faulkner 1709 (K); Greenway 4631 (K); Robertson & Luke 6468 (K); B–D, Bogner 218 (Kew spirit collection 56422); E–F, Robertson & Luke 6468 (K).

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leaves. PEDUNCLE: slender, equalling or longer than petiole, sometimes exceeding entire leaf. SPATHE: broadly ovate-elliptic to subcircular, acuminate, decurrent, white, ± fully expanded and sometimes arching backwards at anthesis, persistent, closing at fruiting stage. SPADIX: shorter than spathe, female zone entirely adnate to spathe, ± laxly flowered with gynoecia ± biseriately arranged, contiguous with male zone or with short, naked axis in between, male zone equalling female, cylindric, densely flowered, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: probably 1-androus (2–3-androus according to Engler 1920a), stamens free, depressed subquadrate, anthers sessile, thecae opposite with subovoid microsporangia dehiscing by apical pore. POLLEN: inaperturate, spherical to subspheroidal, medium-sized (mean 37 µm.), exine papillate-spinose. FEMALE FLOWER: gynoecium slender, flask-shaped, falcate, yellowish, ovary 1-locular, ovule 1, anatropous, placenta basal, stylar region rather long-attenuate, stigma small, discoid-subhemispheric, slightly wider than style. BERRY: ovoid-ellipsoid and slightly angled, style forming persistent mucro, green. SEED: ovoid-ellipsoid, testa ± smooth, thin, embryo elongate, straight to slightly curved, endosperm copious. See Plates 78, 124B. CHROMOSOMES: 2n = 36. DISTRIBUTION: 1 sp.; Kenya, Tanzania. ECOLOGY: tropical humid forest; creeping, rhizomatous geophytes, in leaf litter. NOTES: Callopsis resembles Ulearum in various respects but lacks anastomosing laticifers and has a narrower connective. In Callopsis the thecae open by a pore which later becomes a very broad slit. The two microsporangia of each theca thus have a single common opening. The stamens appear to be free and uniformly separated to a depth which reaches to the base of the microsporangia. This makes it impossible to say from mature material how many stamens form a male flower. Developmental studies will probably show that the male flower is unistaminate. ETYMOLOGY: Calla and the Greek opsis (appearance). TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1980c), Mayo (1985a).

78. Callopsis

Tribe Thomsonieae

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Tribe Thomsonieae Blume, Rumphia 1: 138 (1837); Bogner et al. in Aroideana 8 (1): 14–25 (1985). Laticifers simple, articulated; seasonally dormant, stem tuberous, occasionally rhizomatous, hypogeal; leaf usually 1; petiole long, terete, variously maculate; blade dracontioid, first foliage leaf of seedling divided, never entire, the three primary leaf divisions upright in bud, ultimate lobes usually elliptic-lanceolate, acuminate, primary lateral veins of ultimate lobes pinnate, forming submarginal collective vein, higher order venation reticulate; inflorescence 1, subtended by cataphylls; spadix with sterile terminal appendix, female zone usually contiguous with male; flowers unisexual, perigone absent; pollen extruded in strands, exine various; ovules 1 per locule, anatropous; seed testa smooth, embryo large, endosperm absent.

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79. Amorphophallus Amorphophallus Blume ex Decaisne in Nouv. Ann. Mus. Hist. Nat. 3: 366 (1834), nom. cons. TYPE: A. campanulatus Decaisne (= A. paeoniifolius (Dennstedt) Nicolson). SYNONYMS: Thomsonia Wallich, Pl. Asiat. Rar. 1: 83 (1830), Bogner in Pl. Syst. Evol. 125 (1): 15–20 (1976), nom. rej.; Pythion Martius in Flora 14: 458 (1831), nom. rej.; Candarum Reichenb. ex Schott & Endlicher, Melet. Bot. 17 (1832), nom. illeg.; Pythonium Schott in Schott & Endlicher, Melet. Bot. 17 (1832), nom. illeg.; Kunda Rafinesque, Fl. Tell. 2: 82 (1837, “1836”), nom. illeg.; Brachyspatha Schott, Syn. Aroid. 35 (1856); Conophallus Schott, Syn. Aroid. 34 (1856); Plesmonium Schott, Syn. Aroid. 34 (1856), Bogner in Adansonia 20 (3): 305–308 (1980); Corynophallus Schott in Oesterr. bot. Wochenbl. 7: 389 (1857); Allopythion Schott, Gen. Aroid. t. 24 (1858); Hansalia Schott in Oesterr. bot. Zeitschr. 8: 82 (1858); Hydrosme Schott, Gen. Aroid. t. 33 (1858); Rhaphiophallus Schott, Gen. Aroid. t. 27 (1858); Synantherias Schott, Gen. Aroid. t. 28 (1858); Proteinophallus J.D. Hooker in Bot. Mag. 101: t. 6195 (1875). HABIT: seasonally dormant (sometimes irregularly so) or rarely semi-evergreen herbs, often large, sometimes gigantic, tuber usually depressed-globose, sometimes irregularly ± elongate-cylindric, napiform or carrot-shaped, rarely rhizomatous or stoloniferous. LEAVES: usually solitary (rarely 2–3) in adult plants, sometimes 2–3 in seedlings. PETIOLE: long, usually smooth, rarely verrucose to asperate, sometimes very thick, usually conspicuously spotted and marked in a variety of patterns, sheath very short. BLADE: dracontioide: i.e. trisect, primary divisions pinnatisect, bipinnatisect or dichotomously further divided, tubercles rarely present at junction of divisions, secondary and tertiary divisions ± regularly pinnatifid to pinnatisect, ultimate lobes oblongelliptic to linear, acuminate, decurrent, rarely petiolulate; primary lateral veins of ultimate lobes pinnate, forming distinct submarginal collective vein, higher order venation reticulate. INFLORESCENCE: always solitary, preceded by cataphylls, usually flowering without leaves, rarely with the leaves. PEDUNCLE: very short to long, similar to petiole. SPATHE: variously coloured, marcescent and finally deciduous, boat-shaped and not or hardly convolute, or clearly differentiated into tube and blade, sometimes constricted between them; tube convolute, rarely connate (A. pusillus,

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Plate 79 (i). Amorphophallus. A, habit × 1/20; B, habit × 1/18; C, habit × 1/0; D, part of leaf × 1/6; E, detail of leaf × 2/3; F, part of leaf × 1/6; G, part of leaf × 1/6; H, detail of petiole × 2/3; J, detail of petiole × 2/3. Amorphophallus decus-silvae: A, Java (Kew slide collection); A. maculatus: B, Bogner 600 (Kew illustration collection); A. sp.: C, Richards 741–64 (Kew slide collection); A. galbra: D–E, Brown 5801 (K); A. sp.: F, Cult. Kew. (K); A. dracontioides: G, Johnson 662 (K); A. paeoniifolius : H, Cult. Kew. (Kew slide collection); A. krausei : J, Cult. Kew (Kew slide collection).

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Plate 79 (ii). Amorphophallus. A, inflorescence × 1/3; B, inflorescence × 1/3; C, spadix × 2/3; D, inflorescence × 1/3; E, inflorescence × 2/3; F, detail of fertile part of spadix × 3; G, inflorescence × 1/8; H, inflorescence × 1/6; J, detail of spadix appendix × 2/3; K, inflorescence × 2/3; L, inflorescence × 2/3; M, detail of fertile part of spadix and base of appendix × 3; N, inflorescence × 1/6; Amorphophallus pendulus: A, Cult. Kew 1968–200 (Kew spirit collection 7237); A. krausei: B, H.AM. 40 (Kew slide collection); A. dracontioides: C, I.F.H.19156 (Kew spirit collection 21971); A. lewallei: D, H.AM. 20B (Kew slide collection); A. sumawongii: E–F, Bogner 372 (Kew spirit collection 26332.44); A. paeoniifolius: G, Bogner s.n. (Kew spirit collection 34673); H.AM. 41A (Kew slide collection ); A. hirtus: H, H.AM. 132A (Kew slide collection); J, (Kew spirit collection 58005); A. pusillus: K, H.AM. 246 (Kew slide collection); A. pygmaeus: L–M, Bogner 1948 Cult. Hetterscheid (Kew spirit collection 59079); A. gomboczianus: N, H.AM. 3 (Kew slide collection).

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Plate 79 (iii). Amorphophallus. A, stamen × 10; B, gynoecium × 10; C, gynoecium, longitudinal section × 10; D, stamen × 10; E, gynoecium × 10; F, gynoecium, longitudinal section × 10; G, stamen × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, inflorescence, nearside half of spathe removed × 4; L, stamen × 10; M, gynoecium × 10; N, gynoecium, longitudinal section × 10; P, stamen × 10; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, gynoecium, transverse section × 10; T, stamen × 10; U, gynoecium × 10; V, gynoecium, longitudinal section × 10; W, stamen × 10; X, gynoecium × 10; Y, gynoecium, longitudinal section × 10; Z, stamen × 10; AA, gynoecium × 10; BB, gynoecium, longitudinal section × 10. Amorphophallus albispathus: A–C, H.AM 16 (Kew slide collection); A. pygmaeus: D–F, Bogner 1948 Cult. Hetterscheid (Kew spirit collection 59079); A. pendulus: G–J, Cult. Kew 1968–200 (Kew spirit collection 7237); A. pusillus: K, H.AM. 246 (Kew slide collection); A. gomboczianus: L–N, Ash 1434 (Kew spirit collection 15077.14); A. hirtus: P–S, (Kew spirit collection 58005); A. sumawongii: T–V, Bogner 372 (Kew spirit collection 26332.44); A. paeoniifolius: W–Y, Bogner s.n. (Kew spirit collection 34673); A. dracontioides: Z–BB, I.F.H.19156 (Kew spirit collection 21971).

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Plate 79 (iv). Amorphophallus. A, tuber × 1/6; B, tuber × 1/6; C, spathe base interior sculpturing × 4; D, spathe base interior sculpturing × 4; E, spathe base interior sculpturing × 4; F, spathe base interior sculpturing × 4; G, spathe base interior sculpturing × 4. Amorphophallus longituberosus: A, H.AM.120 (Kew illustration collection); A. corrugatus: B, H.AM. 229 (Kew illustration collection); A. johnsonii: C, H.AM. 41 (Kew slide collection); A. maculatus: D, Bogner 600 (Kew slide collection); A. parvulus: E, H.AM. 26A (Kew slide collection); A. henryi: F, H.AM. 97 (Kew slide collection); A. bulbifer: G, Bogner s.n., no other data (Kew slide collection).

A. elliotii), campanulate to cylindric or ventricose, inner surface smooth or longitudinally ribbed or near base verruculose, scabrate or densely covered with scale- or hairlike processes or smooth; blade erect to spreading, smooth, ribbed or variously undulate or frilled at margins. SPADIX: sessile or stipitate, shorter or much longer than spathe; female zone shorter, equalling or longer than male zone; male zone cylindric, ellipsoid, conoid or obconoid, usually contiguous with female, sometimes separated by a sterile zone which may be naked, or bear prismatic, subglobose or hair-like sterile flowers; terminal appendix usually present, rarely absent or reduced to a stub, erect, sometimes horizontal, rarely pendent, very variable in shape, usually ± conoid or cylindric, rarely ± globose, sometimes ± stipitate or basally narrowed, usually smooth or bearing staminodelike structures near base or entirely covered with staminodes, sometimes corrugate or densely to sparsely hirsute, or grossly and irregularly crumpled. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–6-androus, stamens free or sometimes connate in basal flowers or throughout male zone, short, filaments absent or distinct, connective fairly thick, sometimes projecting beyond thecae, thecae obovoid or oblong, opposite, dehiscing apically by an apical (rarely lateral) pore or transverse slit. POLLEN: extruded in strands, inaperturate, mostly ellipsoid to ellipsoid-oblong, occasionally spherical or subsphaeroidal, medium-sized to large (mean 53 µm., range 34–82 µm.), exine striate, striate-reticulate, psilate, scabrate, areolate, fossulate, punctate-foveolate, verrucate, or spinose. FEMALE FLOWER: gynoecia usually crowded, sometimes ± distant, ovary subglobose to ovoid or obovoid, 1–4-locular, ovules 1 per locule, anatropous, funicle very short to distinct, erect, placenta axile to basal, stylar region absent, short or very long, conoid to cylindric, stigma variably shaped, entire and subglobose or 2–4-lobed or stellate or rarely punctiform, sometimes large and brightly coloured. BERRY: sometimes very large, 1- to few-seeded, orange to red, rarely blue or white, infructescence ± cylindric. SEED: ellipsoid, testa

smooth, thin, embryo large, somewhat green superficially, endosperm absent. See Plates 79i–iv, 124C–D. CHROMOSOMES: 2n = 26, 28, 39. DISTRIBUTION: ca. 170 spp. (W. Hetterscheid, pers. comm.); tropical Africa, Madagascar, tropical Asia, Malay Archipelago, Melanesia, Australasia:– Angola, Australia, Bangladesh, Benin, Bhutan, ?Botswana, Brunei, Burkina Faso, Burma, ?Burundi, ?Cabinda, Cambodia, Cameroon, Central African Republic, Chad, S. China (incl. Taiwan), Congo, Equatorial Guinea (Bioko, Rio Muni), Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, India, Indonesia, Ivory Coast, Japan, Kenya, Laos, Liberia, Madagascar, Malawi, Malaysia, Mali, Mozambique, ?Namibia, Nepal, Niger, Nigeria, Papua New Guinea, Philippines, ?Rwanda, Senegal, Sierra Leone, Somalia, South Africa, Sri Lanka, Sudan, Tanzania, Thailand, Togo, Uganda, Vietnam, Zaire, Zambia, Zimbabwe. ECOLOGY: tropical humid forest, seasonal forest, open woodlands; geophytes, sometimes in humus deposits on rocks (limestone), also in waste places or areas of human habitation (e.g. A. paeoniifolius). NOTES: About 14 sections or informal sectional groups are currently recognized, although many of these may be unnatural (W. Hetterscheid, pers. comm.). USES: The tubers of A. paeoniifolius and A. konjac are widely used sources of carbohydrate foods in tropical Asia and Japan respectively. ETYMOLOGY: Greek amorphos (deformed) and phallus (penis), referring to the spadix appendix (especially of A. paeoniifolius). TAXONOMIC ACCOUNTS: Engler (1911), Gagnepain (1942), Hepper (1968), Hu (1968), Bogner (1976a), Li (1979), Sivadasan (1982, 1986, 1989), Knecht (1983), Bogner, Mayo & Sivadasan (1985), Mayo (1985a), Bogner (1986a), Nicolson (1988a), NtépéNyame (1988), Hay (1990a, c), Bogner (1989b), Bogner & Hetterscheid (1992), Hetterscheid (1994), Hetterscheid & Serebryanyi (1994), Hetterscheid, Yadav & Patil (1994), Sivadasan, Mohanan & Rajkumar (1994), Bogner (1995), Hetterscheid & Peng (1995), Hetterscheid & Ittenbach (1996).

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79. Amorphophallus

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80. Pseudodracontium Pseudodracontium N.E. Brown in J. Bot. 20: 193 (1882). LECTOTYPE: P. anomalum N.E. Brown (= P. lacourii (Lind. & André) N.E. Brown)(see Nicolson in Taxon 16: 518. 1967). HABIT: seasonally dormant herbs, tuber depressed-globose, napiform or irregularly elongate. LEAVES: often more than 1. PETIOLE: long, usually mottled or otherwise variegated, sheath very short and inconspicuous. BLADE: dracontioid: i.e. trisect with 3 ± equal primary divisions, anterior division sometimes smaller than posterior divisions, primary divisions mostly pinnatisect or partly bipinnatisect, ultimate lobes decurrent, sessile or shortly petiolulate, narrowly elliptic, acuminate; primary lateral veins of ultimate lobes pinnate, forming submarginal collective vein, 1 marginal vein also present, higher order venation reticulate. INFLORESCENCE: solitary, usually appearing with leaves. PEDUNCLE: long, longer, shorter or subequal to petiole, similar in appearance to petiole. SPATHE: erect,

boat-shaped, convolute basally, slightly fornicate, not constricted, green to yellow. SPADIX: sessile, shorter to subequal to spathe, female zone shorter than male, densely flowered, male zone laxly flowered, contiguous with female zone, terminal appendix usually shorter than male zone, with naked basal stipe, ± conic, covered with sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–6-androus, flowers distinct, ± distant, filaments relatively long, connate to ± free, thecae subglobose, dehiscing by short slit or pore. POLLEN: inaperturate, ellipsoid to oblong, medium-sized (mean 48 µm.), exine narrowly striate. STERILE MALE FLOWERS: composed of clavate, irregularly ± connate staminodes, not forming distinct floral groups. FEMALE FLOWER: ovary ovoid to subglobose, 1-locular, ovule 1, anatropous, funicle short, placenta basal, stylar region shortly attenuate, stigma discoid-subcapitate. BERRY: ellipsoid, with stigma remnant persisting, 1-seeded. SEED: ellipsoid, testa smooth, brown, raphe conspicuous, embryo large, ellipsoid, somewhat truncate at each end, endosperm absent. See Plates 80, 125A.

80. Pseudodracontium

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D

E

C

F

A

B

Plate 80. Pseudodracontium. A, habit × 1/4; B, leaflet × 1/2; C, spadix × 1; D, stamen × 8; E, gynoecium × 8; F, gynoecium, longitudinal section × 8. Pseudodracontium latifolium: A–B, Dransfield 6219 Cult. Kew. 1984–4105; C–F, Dransfield 6219 (Kew spirit collection 58899).

THOMSONIEAE : PSEUDODRACONTIUM

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C

F

A

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B J

K

G

E

L

H

M

Plate 81. Arophyton. A, habit × 1/6; B, leaf × 2/3; C, detail of leaf venation × 4; D, seedling × 2/3; E, inflorescence × 2/3; F, infructescence × 2/3; G, habit × 2/3; H, spadix × 3; J, synandrium, top view × 10; K, synandrode, side view × 10; L, gynoecium with associated synandrode × 10; M, gynoecium, longitudinal section × 10. Arophyton buchetii: A, 7542 (Kew slide collection); B–D, Bogner 207 (K); E–F, Bogner 207 (Kew spirit collection 32138 & 34394). A. tripartitum var. tripartitum: F, Bogner 205 (Kew spirit collection 34400); G, Bogner 205 (K!); H–M, Morat 3626 (K).

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CHROMOSOMES: 2n = 26. DISTRIBUTION: 7 spp.; tropical southeast Asia:– Cambodia, Laos, Thailand, Vietnam. ECOLOGY: tropical humid and deciduous forest, bamboo forest; geophytes, on forest floor. NOTE: Closely related to Amorphophallus, and characterized by the staminodial, stipitate appendix, constantly 1-locular ovaries and male flowers with long, partially connate filaments. ETYMOLOGY: Greek pseudos (false) and Dracontium ; refers to the similarity to Dracontium. TAXONOMIC ACCOUNTS: Engler (1911), Gagnepain (1942), Serebryanyi (1995).

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Tribe Arophyteae Tribe Arophyteae Bogner in Bot. Jahrb. 92: 9 (1972). SYNONYM: Tribe Synandrodieae Buchet in Bull. Soc. Bot. France 86: 279 (1939) Laticifers simple, articulated; seasonally dormant, stem tuberous (except most Arophyton spp.); primary lateral veins pinnate forming 1 submarginal collective vein and 1–2 marginal veins, higher order venation parallel-pinnate to reticulate; spadix fertile to apex (except some Arophyton spp.); flowers unisexual, perigone absent; male flower a synandrium of partially or completely connate stamens (sometimes reduced to 1 stamen), pollen spinose; gynoecium surrounded by cup-like synandrodium, ovary 1-locular, ovule 1, orthotropous, placenta basal; testa thin, smooth, embryo large, endosperm absent.

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81. Arophyton Arophyton Jumelle in Ann. Musée Colon. Marseille, 36 année, sér. 4, 6 (2): 23 (1928). TYPE: Arophyton tripartitum Jumelle SYNONYMS: Synandrogyne S. Buchet in Bull. Soc. Bot. France 86: 69 (1939); Humbertina S. Buchet in Bull. Soc. Bot. France 88: 848–849 (1941). HABIT: seasonally dormant herbs, rhizomatous or tuberous. LEAVES: 1–few. PETIOLE: sheath short. BLADE: cordate, hastate, trifid to trisect or pedatifid; primary lateral veins pinnate, forming submarginal collective vein, secondary laterals sometimes parallel-pinnate, otherwise higher order venation reticulate. INFLORESCENCE: (1–)2–3 in each floral sympodium. PEDUNCLE: subequal or shorter than petiole. SPATHE: not or slightly constricted between tube and blade, tube persistent at maturity or entire spathe marcescent, blade widely spreading, white or cream to greenish within. SPADIX: fertile to apex, or with short appendix, entirely free or female zone partly adnate to spathe, female zone contiguous with male, or separated by synandrodes or a few bisexual flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–7-androus, stamens connate, synandrium elongate-ellipsoid, often irregularly shaped, shallow, truncate, sometimes with a central slit, common connective broad, thecae marginal, broad-ellipsoid to globular, dehiscing by slit or pore on upper surface. POLLEN: inaperturate, spherical to subspheroidal, medium-sized (mean 32 µm., range 27–36 µm.), exine spinose. FEMALE FLOWER: gynoecium surrounded by cuplike synandrode, ovary 1-locular, ovule 1, orthotropous,

81. Arophyton

funicle usually short, placenta basal, stylar region short to relatively long and cylindric-conoid, or ± absent, stigma discoid. STERILE FLOWERS: consisting of a truncate synandrode with central cavity. BISEXUAL FLOWERS: as for female flowers, but surrounded by synandrium bearing a few scattered thecae. BERRY: ellipsoid to fusiform or clavate, red or green. SEED: ellipsoid to globular, testa thin, smooth, embryo large, globular or ellipsoid, plumule subapical, endosperm absent. See Plates 81, 125B. CHROMOSOMES: 2n = 38, 54, 76 (40). DISTRIBUTION: 7 spp.; Madagascar. ECOLOGY: tropical humid, seasonal forest or deciduous forest on limestone; geophytes or epiphytes, litter-filled crevices and holes. ETYMOLOGY: Greek aron (Arum) and phyton (plant). TAXONOMIC ACCOUNTS: Bogner (1972, 1975).

C

82. Carlephyton Carlephyton Jumelle in Ann. Musée Colon. Marseille, 27 année, ser.3, 7: 187 (1919). TYPE: C. madagascariense Jumelle HABIT: seasonally dormant herbs, tuber depressed-globose. LEAVES: 1–2(–3). PETIOLE: sheath short. BLADE: cordate; primary lateral veins pinnate, forming submarginal collective vein, 1–2 marginal veins also present, secondary and tertiary laterals mostly parallel-pinnate, higher order venation reticulate. INFLORESCENCE: 1–2(–3) in each floral sympodium, appearing before or with leaf. PEDUNCLE: shorter than petiole. SPATHE: not constricted, upper half marcescent, lower part persistent, ± fully expanded at anthesis, later closing, cream to dull purple within. SPADIX: fertile to apex, lower part of female zone adnate to spathe, male and female zones contiguous or with a few bisexual flowers between them, sometimes (C. glaucophyllum) basal zone of spadix composed almost entirely of bisexual flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–6-androus, stamens connate, synandrium somewhat angled, truncate apically, common connective broad, thecae either marginal, projecting laterally and inverted, thus

AROPHYTEAE : CARLEPHYTON

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E

H

K

D

G

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C L

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A

N

P

M

Plate 82. Carlephyton. A, habit in flower with leaf emerging × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium, side view × 15; E, synandrium, top view × 15; F, gynoecium with associated synandrode × 15; G, gynoecium, longitudinal section × 15; H, bisexual flower × 15; J, bisexual flower, longitudinal section × 15; K, infructescence × 1; L, habit × 2/3; M, inflorescence × 2; N, synandrium, top view × 10; P, gynoecium with associated synandrode × 15. Carlephyton madagascariense: A, Bot. Jahrb. Syst. 92: 50, Fig. 4 (1972); B, Bogner 169 (K); C–J, Bogner 169 (Kew spirit collection 56421); K, Bogner 279 (Kew spirit collection 34032); C. diegoense: L, 6449 (Kew slide collection); M–P, Bogner 234 (Kew spirit collection 7700).

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83. Colletogyne

82. Carlephyton

dehiscing by slit on lower surface, or stamens not completely connate, upper part of filaments ± free and thickened with thecae dehiscing by apical slit, sometimes remote. POLLEN: inaperturate, spherical or subspheroidal, medium-sized (mean 34 µm., range 33–35 µm.), exine spinose. FEMALE FLOWER: gynoecium surrounded by cup-like synandrode, ovary 1-locular, ovule 1, orthotropous, funicle short, placenta basal, stylar region, short, stigma discoid to subhemispheric. BISEXUAL FLOWERS: gynoecium surrounded by synandrium bearing 1–4 thecae, ovary as above. BERRY: ellipsoid to fusiform, orangered. SEED: ellipsoid, testa thin, smooth, embryo ellipsoid, plumule lateral, endosperm absent. See Plates 82, 125C. CHROMOSOMES: 54, 108. DISTRIBUTION: 3 spp.; northern Madagascar. ECOLOGY: tropical deciduous forest on limestone or basalt; geophytes, in rock crevices and holes with leaf litter. NOTES: Bogner (1972) recognized 2 sections:– sect. Carlephyton, sect. Pseudocolletogyne. ETYMOLOGY: named for G. Carle with the Greek suffix -phyton (plant). TAXONOMIC ACCOUNTS: Bogner (1972, 1975).

C

gated, anther apical-oblique, thecae ellipsoid, dehiscing by slit. POLLEN: inaperturate, spherical, medium-sized (mean 39 µm.), exine spinose, obscurely verruculate between spines, spines long, stout. FEMALE FLOWER: gynoecium surrounded by a cup-like, variegated synandrode, ovary ovoid-ellipsoid, 1-locular, ovule 1, orthotropous, funicle short, placenta basal, stylar region shortly attenuate, stigma discoid-capitate. BERRY: fusiform, stigma remnant persistent, reddish-spotted. SEED: ellipsoid, testa thin, smooth, embryo with lateral plumule, endosperm absent. See Plates 83, 125D. CHROMOSOMES: 2n = 54. DISTRIBUTION: 1 sp.; northern Madagascar. ECOLOGY: tropical deciduous forest, on limestone; geophytes in holes or crevices with leaf litter. NOTES: The 1-androus male flowers are thought to have been derived phylogenetically from multistaminate synandria (Bogner 1972). ETYMOLOGY: Greek kollêtos (glued together) and gynê (woman); the female part of the spadix is adnate to the spathe. TAXONOMIC ACCOUNTS: Bogner (1972, 1975).

Tribe Peltandreae

C

83. Colletogyne Colletogyne Buchet in Bull. Soc. Bot. France 86: 23 (1939). TYPE: C. perrieri Buchet HABIT: seasonally dormant herb, tuberous. LEAVES: 1 (–2). PETIOLE: sheath short. BLADE: cordate; primary lateral veins pinnate and also arising from petiole insertion, forming submarginal collective vein, 1–2 marginal veins also present, secondary and tertiary laterals mostly parallel-pinnate, higher order venation reticulate. INFLORESCENCE: 1–3 in each floral sympodium, appearing before or with leaves. PEDUNCLE: shorter than petiole. SPATHE: obovate, not constricted, erect, fully expanded at anthesis, closing later and persisting to fruiting stage, red-spotted on greenish background. SPADIX: fertile to apex, female zone adnate to spathe, female and male zones contiguous. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, filament conoid, varie-

Tribe Peltandreae Engler in Nova Acta Acad. Leopold.Carol. 39: 146 (1876). SYNONYM: Tribe Typhonodoreae Engler in Nova Acta Acad. Leopold.-Carol. 39: 146 (1876). Laticifers simple, articulated; rhizome hypogeal; primary lateral veins of leaf blade pinnate, forming submarginal collective vein, 1–2 marginal veins also present, higher order venation ± parallel-pinnate; inflorescences usually 2, infructescence pendent; spathe ± constricted centrally, tube ellipsoid to oblong, persistent, blade gaping at anthesis, later marcescent; flowers unisexual, perigone absent; male flower a truncate, prismatic synandrium of connate stamens, fused connectives very thick, thecae lateral, dehiscing by apical pore or short slit; ovary 1-locular, ovules orthotropous, style narrower than ovary; berry and seeds large, embryo large, plumule well-developed, endosperm vestigial to absent.

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B

A

D

E

F

C

Plate 83. Colletogyne. A, habit × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium × 16; E, gynoecium with associated synandrode × 16; F, gynoecium, longitudinal section × 16. Colletogyne perrieri: A, 5489 (Kew slide collection); B–F, Bogner 165 (Kew spirit collection 42467 & photograph (M)).

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84. Peltandra Peltandra Rafinesque in J. Phys. Chim. Hist. Nat. Arts 89: 103 (1819), nom. cons. TYPE: P. undulata Rafinesque (= P. virginica (L.) Rafinesque). SYNONYMS: Lecontia W. Cooper ex Torrey, Compend. fl. n. middle stat. 358 bis (1826); Rensselaeria L.C. Beck, Bot. N. Middle U.S. 382 (1833). HABIT: seasonally dormant herbs, rhizome short, hypogeal, sometimes branching. LEAVES: several. PETIOLE: sheath rather long, up to at least half as long as petiole. BLADE: lanceolate to broadly ovate, sagittate, hastate or rarely cordate; basal ribs well-developed, primary lateral veins pinnate, sometimes only weakly differentiated, forming submarginal collective vein, 2 or more distinct marginal veins also present, secondary laterals parallel-pinnate, higher order venation parallel-pinnate near midrib, becoming reticulate towards margin. INFLORESCENCE: 1–2 in each floral sympodium. PEDUNCLE: subequal to or shorter than petiole, erect at anthesis, bent downwards in fruit. SPATHE: constricted between tube and blade, tube convolute, externally green to yellow-green, ellipsoid, persistent, blade green to white, erect, widely expanded (P. sagittifolia) or only gaping (P. virginica), later deciduous. SPADIX: sessile, cylindric, female zone shorter than male, partially adnate to spathe, either contiguous with male zone or with a short zone of sterile flowers in between, male zone with short terminal appendix of sterile flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 4–5androus, truncate, often slightly excavated centrally, anthers lateral, thecae adjacent, oblong-ellipsoid, dehiscing by apical pore. POLLEN: inaperturate, ellipsoid or spherical, mediumsized (mean 33 µm., range 28–37 µm.), exine spinose (P. virginica) or almost psilate (P. sagittifolia). FEMALE FLOWER: gynoecium closely surrounded by 3–5 free staminodes or some staminodes free and others connate or all connate into a truncate-urceolate synandrode, ovary ovoid, 1-locular, ovules 1–few, hemiorthotropous, funicle short, placenta parietal to basal, stylar region shortly attenuate, narrower than

84. Peltandra

ovary, stigma discoid-hemispheric. BERRY: obconical to irregularly subglobose, green to blackish-purple (P. virginica) or red (P. sagittifolia), 1–3-seeded, with mucilaginous contents. SEED: large, ovoid to subglobose, laterally flattened, embryo large, plumule well-developed with 6–7 leaf primordia, endosperm vestigial or absent. See Plates 84, 126A. CHROMOSOMES: 2n = 56, 112. DISTRIBUTION: 2 spp.; eastern North America:– Canada (Ontario, Quebec), USA (Alabama, Arkansas, Carolinas, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Hampshire, New Jersey, New York, Ohio, Pennsylvania, Rhode Is., Tennessee, Texas, Vermont, Virginia, West Virginia, Wisconsin). ECOLOGY: temperate wetland habitats; helophytes, marshes, along watercourses, brackish water. ETYMOLOGY: Greek peltê (small shield) and aner, andros (man). TAXONOMIC ACCOUNTS: Engler (1915), Barkley (1944), Huttleston (1953), Blackwell & Blackwell (1975).

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85. Typhonodorum Typhonodorum Schott in Oesterr. bot. Wochenbl. 7: 70 (1857). TYPE: T. lindleyanum Schott SYNONYM: Arodendron Werth in Mitt. Sem. Orient. Sprachen 4 (Abt. 3): 112 (1901). HABIT: evergreen, robust to gigantic herb (to 4m), rhizome thick, subterranean, spreading, terminal shoot forming massive banana-like pseudostem formed by the lower part of the petiole sheaths. LEAVES: several in terminal crown. PETIOLE: very long, free apical part relatively short, sheath extending for most of petiole length. BLADE: very large, subtriangular to sagittate, acute; basal ribs well-developed, primary lateral veins pinnate, forming submarginal collective vein very near to margin, 3–4 marginal veins also present, secondary lateral

85. Typhonodorum

PELTANDREAE : TYPHONODORUM

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B

D

E

A

G F

C

H

Plate 84. Peltandra. A, habit, rhizome cut through × 1/2; B, detail of leaf venation × 5; C, spadix × 1; D, synandrium, top view × 8; E, synandrium, side view × 8; F, gynoecium with associated staminodes, three quarter view × 8; G, gynoecium, longitudinal section × 8; H, berry × 2. Peltandra virginica: A, Curtis 4572 (K); Cult. Kew 1969–19636 (Kew spirit collection 29047.739); B–G, Cult. Kew 1969–19636 (Kew spirit collection 29047.739); H, Herb. Careyanum s.n. (K).

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D A

B

F

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A J

E

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Plate 85. Typhonodorum. A, habit in × 1/30; B, habit in flower and fruit × 1/25; C, leaf × 1/5; D, detail of leaf venation × 4; E, spadix × 2/3; F, staminodes from upper sterile male zone × 5; G, synandrium × 5; H, synandrode from lower sterile male zone × 5; J, gynoecium with associated staminodes × 5; K, gynoecium, longitudinal section × 5; L, infructescence, spathe partly removed × 1/2. Typhonodorum lindleyanum: A, 5497 (Kew slide collection); B, 4M 4.91 (Kew slide collection); C, Cult. Kew 1979–4434; D, Vaughn 445 (K); E–L, Cult. Kew 1979–4434 (Kew spirit collection 29047.658); Lourance s.n. (Kew spirit collection 29047.356).

PELTANDREAE : TYPHONODORUM

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and higher order venation densely parallel-pinnate. INFLORESCENCE: 1–2 in each floral sympodium. PEDUNCLE: erect, bent downwards in fruit. SPATHE: erect, constricted between tube and blade, tube convolute, oblong-ellipsoid, green, persistent, blade longer than tube, narrowly boatshaped, acuminate, gaping at anthesis, cream-coloured, marcescent, later deciduous. SPADIX: shorter than spathe, sessile, free, differentiated into 4 zones, basal female zone with staminodes scattered between gynoecia, separated from male zone by narrower zone of sterile male flowers, male zone thicker, terminal zone composed of sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4–6-androus, stamens connate into truncate synandrium, connective very broad, thecae lateral, sessile, dehiscing by short longitudinal slit. POLLEN: inaperturate, ellipsoid, medium-sized (mean 45 µm.), exine subverrucate or obscurely foveolate or punctate. LOWER STERILE MALE FLOWERS: synandrode truncate, subhexagonal becoming narrowly rhomboid to trapezoid towards male zone. UPPER STERILE MALE FLOWERS: composed of 3–6 obpyramidal, truncate staminodes, irregularly connate or merely crowded together. FEMALE FLOWER: gynoecia surrounded by irregularly scattered, obpyramidal, truncate staminodes, ovary shortly ovoid to globose, 1-locular, ovule 1(–2), orthotropous, funicle thick, short, placenta basal, stylar region very shortly attenuate, stigma rather broad, shallowly 3–6-lobed. BERRY: borne in subcylindric, pendent infructescence enclosed in persistent spathe tube, berries very large, compressed-obovoid to -globose, bright yellow, pericarp fleshy. SEED: large, compressed-obovoid, testa thin, ± smooth, embryo very large, plumule exceptionally welldeveloped with several leaf primordia, endosperm absent. See Plates 85, 126B. CHROMOSOMES: 2n = 112. DISTRIBUTION: 1 sp.; Comores, Madagascar, Mauritius, Tanzania (Pemba, Zanzibar). ECOLOGY: tropical wetland habitats; helophytes, freshwater swamps, along rivers, coastal lagoons in brackish water, forming huge populations.

86. Arisarum

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ETYMOLOGY: Greek typhôn (personification of stormy winds) and doron (gift); a poetic name. TAXONOMIC ACCOUNTS: Engler (1915), Bogner (1975), Mayo (1985a).

Tribe Arisareae

C

Tribe Arisareae Dumortier, Fl. Belg. 162 (1827). Laticifers simple, articulated; seasonally dormant, stem a tuber or rhizome; leaves 1–3; petiole usually relatively long; primary lateral veins forming submarginal collective vein, higher order venation reticulate; inflorescence 1, appearing with leaves; spathe not constricted, tube margins connate, blade fornicate, gaping; spadix with usually smooth, sterile, terminal appendix, female zone adnate to spathe, very short, contiguous with male, male zone laxly flowered; flowers unisexual, perigone absent; male flower 1-androus, filament distinct, anther peltately attached, circular, thecae apically confluent, dehiscing by 1 continuous slit, pollen ellipsoid, striate-reticulate; gynoecia 2–5, ovary 1-locular, ovules several, orthotropous, placenta basal; endosperm copious.

86. Arisarum Arisarum P. Miller, Gard. Dict. abr. ed., [121] (1754). LECTOTYPE: A. vulgare Targioni-Tozzetti (Arum arisarum L., see Nicolson in Taxon, 24: 467. 1975). SYNONYMS: Arisaron Adanson, Fam. 2: 470 (1763); Balmisa M. Lagasca, Gen. Sp. Pl. Nov. 17 (1816). HABIT: small, seasonally dormant herbs, stem an ovoid to cylindric tuber or slender rhizome (A. proboscideum) with stolons. LEAVES: 1–2(–3). PETIOLE: often sparsely spotted, sheath short. BLADE: cordate-sagittate to subhastate; primary lateral veins pinnate and also arising at petiole insertion, forming submarginal collective vein, 2 marginal

C

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F

A

B

C

D

G

E

J

H

K

Plate 86. Arisarum. A, habit × 2/3; B, spadix × 2; C, stamen × 15; D, gynoecium × 15; E, gynoecium, longitudinal section × 15; F, infructescence × 2; G, spadix × 2; H, habit × 2/3; J, spadix × 2; K, infructescence × 2. Arisarum proboscideum: A, Sievercori s.n. (K); B–E, Cult. Boyce (Kew spirit collection 53920); F, Cult. Meikle (Kew spirit collection 28255); A. simorrhinum: G, Cult. Salmon (Kew spirit collection 53914); A. vulgare : H, Mitchell, Cheese & Watson 3880 (K); J, Cook & Keesing (Kew spirit collection 51085); K, Cult. Kew Marr 1996 (Kew spirit collection 51128).

ARISAREAE : ARISARUM

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veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: shorter or equalling leaf, often spotted. SPATHE: eventually evanescent, tube erect, margins connate, cylindric to subventricose, slightly constricted at apex, white or with white to pale green longitudinal stripes, blade fornicate, gaping, sometimes subtomentose, apex cuspidate or drawn out into a very long, erect to twisted thread, greenish or brownish or purple-brown. SPADIX: female zone adnate to spathe, 2–5-flowered, contiguous with male zone, male zone laxly flowered, extending for more than half spathe tube length, terminal appendix naked, either stipitate with massive apical knob, or stipitate with thick, clavate, basally truncate, fungoid, apical region, or not stipitate and slenderly clavate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, filament terete, as long or longer than anther, anther peltately attached, circular, connective slender, thecae apically confluent, dehiscing by single continuous slit. POLLEN: pollen shed in amorphous mass, inaperturate, ellipsoid-elongate, medium-sized (mean 45 µm., range 43–46 µm.), exine striate to foveo-reticulate. FEMALE FLOWER: ovary 1-locular, depressed-globose, ovules many, orthotropous, funicle short, placenta basal, stylar region ± abruptly narrowed, stigma small, subhemispheric. BERRY: hemispheric, flattened at apex with elevated angled margins, few-seeded, pericarp carnose-leathery, style base persistent. SEED: ovoid, with large, irregularly conoid strophiole (aril), testa longitudinally rugose, embryo terete, straight, axile, endosperm copious. See Plates 86, 126C. CHROMOSOMES: 2n = 28, 42, 56. DISTRIBUTION: 3 spp.; Mediterranean Europe, Macaronesia:– Albania, Algeria, Bosnia-Hercegovina, Croatia, Cyprus, Egypt, France (incl. Corsica), Greece (incl. Crete), Israel, Italy (incl. Sardinia, Sicily), Lebanon, Libya, Macedonia, Morocco, Portugal (incl. Azores), Serbia, Slovenia, Spain (incl. Balearics, Canary Is.), Syria, Tunisia, Turkey. ECOLOGY: warm temperate scrub and woodland; geophytes, on stony ground in macchie (“maquis”), between rocks or under trees and shrubs. ETYMOLOGY: Greek word arisaron (as used by Dioscorides in reference to aris, aridos, the name of a small herb mentioned by Pliny, possibly Arisarum itself and aron (Arum)). TAXONOMIC ACCOUNTS: Engler (1920a), Boyce (1990).

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Tribe Ambrosineae Tribe Ambrosineae Schott in Schott & Endlicher, Melet. Bot. 16 (1832 “Ambrosinieae”). Laticifers simple, articulated; very small, seasonally dormant, stem a rhizomatous tuber; leaves several, blade ovate-elliptic, primary lateral veins mostly arising at petiole insertion, arcuate, running into apex, higher order venation reticulate; inflorescence 1, on recurved peduncle; spathe boat-shaped, unconstricted externally, margins free, basal part convolute, apical part narrowly gaping, rostrate, internally divided into 2 chambers, an upper, morphologically ventral one and a lower, morphologically dorsal one; spadix shortly appendiculate, adnate to spathe by longitudinal, septum-like lateral dilations, ventral chamber containing female zone, dorsal chamber containing 8–10 male flowers arranged in 2 longitudinal rows; flowers unisexual, perigone absent; male flower a 2-androus, sessile synandrium, thecae confluent, dehiscing by single continuous slit, pollen exine striate-reticulate;

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THE GENERA OF ARACEAE

gynoecium 1, ovary 1-locular, ovules many, orthotropous, placenta basal, style long, curved, stigma discoid; berry manyseeded; seed subglobose, testa costate, strophiolate, endosperm copious.

87. Ambrosina Ambrosina Bassi, Ambrosina Nov. Pl. Gen. 3 (1763); Bononiensi Sci. Inst. Acad. Comment. 5 (2): 82–86 (1766, “1767”). TYPE: A. bassii L. (“Ambrosinia ”). SYNONYM: [Ambrosinia L., Gen. pl., ed. 6, 579 (1764), orth. var.]. HABIT: very small herb, seasonally dormant, stem a rhizomatous tuber. LEAVES: 2–4. PETIOLE: sheath short. BLADE: ovate or ovate-elliptic, obtuse, often spotted; primary lateral veins 2–3 on each side, mostly arising at petiole insertion, arcuate, running into apex, higher order venation reticulate. INFLORESCENCE: solitary, lying horizontally on ground. PEDUNCLE: short, hypogeal, elongating in fruit. SPATHE: ellipsoid, boat-shaped, not constricted, basally convolute, gaping above, interior surfaces bearing hair-like processes, stellate hairs occurring on inner and outer surfaces of ventral (pistillate) chamber, apex forming curved beak. SPADIX: enclosed by the spathe, shortly appendiculate, adnate to internal wall of spathe by septum-like lateral dilations thus forming two longitudinal chambers separating male flowers from female; ventral chamber (held uppermost) containing single female flower, dorsal chamber (held lowermost) containing usually 16 thecae arranged in two parallel rows of 8 each. FLOWERS: unisexual, perigone absent. MALE FLOWER: each row of 8 thecae represents 4 sessile stamens (see note), thecae oriented transversely, opening by longitudinal slit. POLLEN: extruded in irregular masses, inaperturate, ellipsoid-oblong, medium-sized (44 µm.), exine striate-reticulate, with narrow undulate striae in “football” pattern, breaking up into verrucae at ends. FEMALE FLOWER: ovary 1-locular, ovules many, orthotropous, funicles rather long, placenta discoid, basal, stylar region long, attenuate, curved towards spadix axis, stigma discoid, flattened, held ± parallel to spadix axis. BERRY: depressed-globose, many-seeded, style and old

87. Ambrosina

C

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G B

C

A

D

E

F

Plate 87. Ambrosina. A, habit × 1; B, inflorescence, nearside half of spathe removed × 2; C, inflorescence, three quarter view, nearside half of spathe removed × 2; D, stamens × 8; E, gynoecium × 8; F, gynoecium, longitudinal section × 8; G, infructescence, nearside half of pericarp removed × 2. Ambrosina bassii: A, Besler s.n. (K); B–G, Bogner 2101 (Kew spirit collection K 57302).

stigma persistent, whitish with reddish tinge. SEED: subglobose to ellipsoid, testa costate with weak reticulation, brown, hard, with large, white, conical strophiole (aril), embryo straight, elongate, endosperm copious. See Plates 87, 126D. CHROMOSOMES: 2n = 22. DISTRIBUTION: 1 sp.; western Mediterranean Europe:– Algeria, Italy (incl. Lampedusa Is., Sardinia, Sicily). ECOLOGY: warm temperate scrub and woodland; geophytes, usually in macchie (maquis) scrub, forest floor, humus deposits between rocks or in open stony ground. NOTES: The morphological interpretation of the male flower remains uncertain. Benzing (1969: Tafel 27.I) presented an illustration which shows that in each longitudinal row, the thecae are associated in pairs by their vascular supply. Both Engler (1920a) and Benzing state that the male flowers are 2-androus. However, in our opinion the evidence for this conclusion remains weak. ETYMOLOGY: named after B. Ambrosini (1588–1657), botanist at Bologna. TAXONOMIC ACCOUNTS: Engler (1920a), Killian (1929, 1933), Benzing (1969).

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Tribe Areae Tribe Areae Laticifers simple, articulated; minute to robust, seasonally dormant, stem a subglobose to rhizomatous tuber; leaves several (often 1 in Sauromatum), primary lateral veins of leaf blade or lobes or leaflets forming submarginal collective vein, higher order venation reticulate; inflorescence usually 1, appearing with or after leaves (except Sauromatum and some Biarum spp.); spathe usually strongly constricted, tube with convolute margins (partially to fully connate in Sauromatum and in most Biarum spp.), blade gaping, erect

to reflexed; spadix with usually smooth, terminal, sterile appendix; flowers unisexual, perigone absent; stamens usually free, connective slender, pollen exine spinose (except Dracunculus, Biarum ditschianum; scabrous in Arum korolkowii); ovary 1-locular, ovules orthotropous, style usually as broad as ovary (except some Biarum and Arisaema spp.), stigma hemispheric; berry subglobose; seed usually with rather large, fleshy strophiole (aril), testa usually rugosereticulate (except Sauromatum and some Biarum spp.), endosperm copious.

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88. Arum Arum L., Sp. Pl. 964 (1753). LECTOTYPE: Arum maculatum L. (see M.L. Green, Brit. Bot. 186. 1929). SYNONYMS: [Aron Adanson, Fam. 2: 470 (1763), orth. var.]; Gymnomesium Schott in Oesterr. bot. Wochenbl. 5: 17 (1855). HABIT: seasonally dormant herbs, tuber hypogeal, subglobose or rhizomatous and horizontal. LEAVES: 3–4(–6) in each sympodial unit. PETIOLE: sheath long to short. BLADE: cordate (A. pictum) or hastate-sagittate to sagittate; primary lateral veins pinnate, forming submarginal collective vein, at least 1 marginal vein also present, higher order venation reticulate. INFLORESCENCE: 1, rarely 2 in each floral sympodium, usually appearing with leaves, rarely just before (A. pictum), hidden within foliage or fully exposed above leaves. PEDUNCLE: much shorter to much longer than petiole. SPATHE: marcescent, usually strongly constricted between tube and blade, rarely not, tube convolute, cylindric to ellipsoid, blade ovate- or oblong-lanceolate to lanceolate, ± acuminate, at anthesis erect and boat-shaped or widely expanded or patent and ± revolute. SPADIX: shorter to longer than spathe, sessile, female zone cylindric, interstice sepa-

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rating male and female zones usually short, rarely absent, usually covered with sterile flowers (pistillodes), rarely naked, male zone cylindric, conoid, ellipsoid or subglobose, interstice separating appendix and male zone usually covered with sterile flowers (staminodes), rarely absent, terminal sterile appendix usually stipitate, then gradually or suddenly dilated into a conoid or cylindric club, sometimes slender. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4androus, stamens free, filaments very short but distinct, connective slender, thecae shortly obovoid, opposite or subopposite, dehiscing by apical pore-like slit, rarely porose (A. pictum). POLLEN: usually presented in a loose, powdery mass, sometimes extruded in strands (A. pictum, A. nigrum), grains inaperturate, spherical to subspheroidal, medium-sized (mean 32 µm., range 29–34 µm.), exine spinose, rarely scabrous (A. korolkowii). STERILE FLOWERS: (staminodes and pistillodes) consisting of basal, ± hemispheric, swollen, verrucose to smooth portion bearing usually one (sometimes more) erect, ± straight, subulate to filiform processes. FEMALE FLOWER: gynoecium cylindric, ovary 1-locular, ovules 6–more, orthotropous, lageniform, biseriate, funicle short, placenta parietal to subbasal, stylar region short, as broad as ovary or absent, stigma subhemispheric, exuding nectar droplet at anthesis. BERRY: obovoid, pericarp juicy, many-seeded, bright glossy orange-red. SEED: ellipsoid to ovoid, testa rugose, especially towards base, with large, succulent obconic strophiole, embryo axile, straight, endosperm copious. See Plates 88, 127A. CHROMOSOMES: 2n = 28, 42, 56, 70, 84. DISTRIBUTION: 25 spp.; Central Asia, Europe, Macaronesia, Mediterranean, Middle East:– Afghanistan, Albania, Algeria, Andorra, Armenia, Austria, Azerbaijan, Belgium, Belorussia (S), Bosnia–Hercegovina, Bulgaria, China (Tien Shan), Croatia, Cyprus, Czech Republic, Denmark, France (incl. Corsica), Georgia, Germany, Greece (incl. islands), Hungary, India (NW), Iran, Iraq, Ireland, Israel, Italy (incl. Sardinia, Sicily), Jordan, Kazakhstan, Kirghizia, Lebanon, Libya, Liechtenstein, Luxembourg, Macedonia, Moldavia, Morocco, Nepal (W), Netherlands, Pakistan, Poland, Portugal (incl. Azores, Madeira), Romania, Russia (far S), San Morino, Serbia, Slovak Republic, Slovenia, Spain (incl. Canary Is., Balearics), Sweden (S), Switzerland, Syria, Tajikistan, Tunisia, Turkey, Turkmenestan, Ukraine (S), United Kingdom (incl. Channel Is.), Uzbekistan.

88. Arum

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ECOLOGY: temperate and warm temperate woodland, up to 4400m alt.; geophytes, forest floor, hedges, orchards, stony open ground, along rivers, open scrub, pastures, abandoned areas. NOTES: Boyce (1989, 1993) recognized 2 subgenera, subgen. Arum (with 2 sections and 6 subsections) and subgen. Gymnomesium. Arum italicum is naturalized in Argentina and New Zealand. ETYMOLOGY: Latin form of the classical Greek name aron. TAXONOMIC ACCOUNTS: Engler (1920a), Arrigoni et al. (1982), Boyce (1989, 1993a,1995).

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89. Eminium Eminium (Blume) Schott, Aroideae 16 (1855, “1853”). LECTOTYPE: E. spiculatum (Blume) Schott (Arum spiculatum Blume; see Nicolson 1967, p. 516). SYNONYM: Helicophyllum Schott, Aroideae 20 (1855, “1853”), non Bridel (1827). HABIT: medium-sized, seasonally dormant herbs, tuber subglobose. LEAVES: 3–6 (–8). PETIOLE: sheath relatively long. BLADE: oblong-elliptic (E. lehmannii, E. regelii), linear- to auriculate-hastate, or pedatifid-pedatisect with lobes of posterior divisions held ± erect in a spiral on the twisted basal ribs; primary lateral veins of lobes pinnate, forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: much shorter than petiole of subtending leaf, apex often greatly thickened. SPATHE: marcescent, tube with convolute margins, subventricose to oblong, blade oblong or ovateoblong, erect, inner surface sometimes densely rugose-puckered. SPADIX: sessile, slender, shorter than spathe, female zone short-cylindric, separated from male zone by longer zone bearing usually scattered sterile flowers, male zone ellipsoid to cylindric, shorter, equalling or longer than female, appendix usually relatively short, elongate-clavate, stoutly to narrowly cylindric, rugose or smooth. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2-androus, stamens free, anthers subsessile to sessile, connective slender, thecae oblong-ellipsoid, dehiscing by apical slit. POLLEN: presented in a loose, powdery mass, not extruded in strands, grains inaperturate, spherical or subspheroidal, medium-sized

89. Eminium

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K C

F

B

J E G H D AA

Y

X

A U Z

V

L

M

N

P

Q

R

S

T

W

Plate 88. Arum. A, habit × 1/5; B, tuber × 2/3; C, tuber × 2/3; D, leaf × 1; E, detail of leaf venation next to midrib × 6; F, detail of leaf venation near margin × 6; G, leaf × 2/3; H, inflorescence, front of lower spathe removed × 2/3; J, spathe, front view × 2/3; K, spathe, side view × 2/3; L, spadix × 2/3; M, detail of fertile portion of spadix × 2; N, spadix × 2/3; P, detail of fertile portion of spadix × 2; Q, spadix × 2/3; R, detail of fertile portion of spadix × 2; S, spadix × 2/3; T, detail of fertile portion of spadix × 2; U, stamen × 10; V, stamens, lower one longitudinally sectioned × 10; W, gynoecia, upper one longitudinally sectioned × 10; X, pistillode × 15; Y, staminode × 15; Z, infructescence × 2/3; AA, seed × 2. Arum maculatum: A, Boyce s.n. (Kew slide collection); A. alpinum: B, Cook & Keesing 56 (Kew spirit collection 51396); A. concinnatum: C, L–M, U, X–Y, Boyce 10 (Kew spirit collection 53952); A. rupicola var. rupicola: D–F, Sintenis 5575 (K); A. italicum subsp. italicum: G, Herb. Churchill s.n. (K); A. orientale subsp. orientale: H, Cult. Tucker (Kew spirit collection 49773); A. rupicola subsp. rupicola: J, Boyce 87 (Kew spirit collection 51615); A. creticum: K, N–P, Cult. Kew (Kew spirit collection 51368); A. orientale subsp. orientale: Q–R, Cult. Tucker (Kew spirit collection 49773); A. pictum: S–T, V–W, Ferguson & Ferguson 4126, Cult. Kew 1985–01888 (Kew spirit collection 51818); A. italicum subsp. italicum: Z, photo 1336 (Kew slide collection); A. alpinum: AA, Cook & Keesing 56 (Kew spirit collection 51898).

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B

C

L

D

A F

E

G

H

M

N

K

J

Plate 89. Eminium. A, habit × 1/3; B, seedling showing contractile root × 2/3; C, detail of fertile portion of spadix × 1 1/2; D, habit × 2/3; E, spadix × 1 1/2; F, stamen × 15; G, gynoecium × 15; H, gynoecium, longitudinal section × 15; J, inflorescence × 2/3; K, spadix × 1 1/2; L, stamen × 15; M, gynoecium × 15; N, gynoecium, longitudinal section × 15. Eminium albertii: A, Furse 7390 (K); Gibbons & Gibbons 47 (K); B, Gibbons & Gibbons 47 (K); C, Grey-Wilson & Hewer 765 (Kew spirit collection 29047.120); E. koenenianum: D, Willdenowia 20: 47, f.4 (1991); E–H, Koenen 32*89 (Kew spirit collection 55074 & 59044); E. spiculatum subsp. spiculatum: J, Boyce 197 (Kew spirit collection 51645 & Kew slide collection); K–N, Boyce 197 (Kew spirit collection 51645).

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(mean 44 µm.; 15–20 µm in E. koenenianum), exine spinose. STERILE FLOWERS: composed of subulate, sometimes flattened, patent, straight to slightly curved processes. FEMALE FLOWER: ovary ellipsoid-obovoid, 1-locular, ovules 2, orthotropous, funicle short, placenta basal to subbasal, stylar region short to inconspicuous, stigma hemispheric. BERRY: subglobose, 1(–2) seeded. SEED: obnapiform to subglobose, testa leathery, rugose, with large strophiole, embryo small, elongate, endosperm copious. See Plates 89, 127B. CHROMOSOMES: 2n = 24, 28. DISTRIBUTION: 8 spp.; Middle East, south central Asia: Afghanistan, Egypt, Iran (N), Iraq (N), Israel, Jordan, Kazakhstan, Kirghizia, Lebanon, Syria, Tajikistan, Turkey, Turkmenestan, Uzbekistan. ECOLOGY: warm temperate and subtropical habitats in fields, savannas, semideserts, deserts; geophytes, stony or sandy ground, consolidated sand. ETYMOLOGY: ancient name eminion, mentioned by Dodoens (1574). TAXONOMIC ACCOUNTS: Engler (1920a), Riedl (1963, 1969, 1985), Lobin & Boyce (1991).

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90. Dracunculus Dracunculus P. Miller, Gard. Dict., Abr. ed., [455] (1754). LECTOTYPE: D. vulgaris Schott (see Nicolson 1975, p. 467). SYNONYMS: Anarmodium Schott in Bonplandia 9: 368 (1861); Dracontium J. Hill, Brit. Herb. 336 (1756, non L. 1753). HABIT: large, seasonally dormant herbs, tuber subglobose, stoloniferous as well in D. canariensis. LEAVES: few. PETIOLE: sheath very long, spotted or not, tightly convolute around peduncle forming substantial erect pseudostem. BLADE: deeply pedatifid; primary lateral veins of each lobe pinnate, forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: shorter than petiole. SPATHE: marcescent, constricted, tube with convolute margins, erect, thick-walled, oblong-ellipsoid, shorter than blade, blade ovate-lanceolate, acuminate, expanded, deep purple or white, eventually bent backwards. SPADIX: subequal to spathe, female zone ± sessile, cylindric to conic, contiguous

with male zone, male zone ellipsoid-cylindric, separated from appendix by a short zone of subulate staminodes, appendix long, long-stipitate, then elongate-conoid. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4androus, stamens quadrate, filaments distinct, connate at base, connective slender and prominulent, thecae shortly ellipsoid, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, ellipsoid to spheroidal, medium-sized (mean 45 µm.), exine verrucate. FEMALE FLOWER: ovary ellipsoid, 1-locular, ovules few, orthotropous, placentae apical and basal, stylar region shortly conic, stigma hemispheric. BERRY: obovoid, few-seeded, red-orange. SEED: globoseovoid, testa rugose, embryo axile, short, endosperm copious. See Plates 90, 127C. CHROMOSOMES: 2n = 28. DISTRIBUTION: 2 spp.; Mediterranean Europe, Macaronesia:– Albania, Bosnia–Hercegovina, Bulgaria, Croatia, Greece (incl. Crete and islands), Italy (incl. Sardinia, Sicily), Macedonia, Portugal (incl. Azores, Madeira), ? Serbia, Slovenia, Spain (Canary Is.), Turkey. ECOLOGY: temperate and warm temperate scrub, woodland, macchie, olive groves, chestnut forest; geophytes, under trees and shrubs, also stony open ground. NOTES: The verrucate pollen exine of Dracunculus is unique in the tribe. The other genera have a spinose, spinulose or rarely smooth exine. The seed of Dracunculus lacks the characteristic strophiole found in other genera of the tribe. ETYMOLOGY: ancient name, Latin draco (dragon, snake) and suffix -unculus (diminutive). TAXONOMIC ACCOUNTS: Engler (1920a), Amaral Franco, Webb & Prime (1980), Boyce (1994).

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91. Helicodiceros Helicodiceros Schott ex K. Koch in Index Sem. Hort. Berol. 1855, Appendix, p. 2 (1856), nom. cons. TYPE: H. crinitus K. Koch, nom. illeg. (Dracunculus crinitus Schott, nom. illeg., Arum muscivorum L. f., H. muscivorus (L.f.) Engler). SYNONYM: Megotigea Rafinesque, Fl. Tell. 3: 64 (1837, “1836”), nom. rej. HABIT: seasonally dormant herb, tuber depressed-globose. LEAVES: several. PETIOLE: variously spotted, sheath long,

90. Dracunculus

AREAE : HELICODICEROS

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B

G J

L

K

H

D

A

E

F

C

Plate 90. Dracunculus. A, habit × 1/6; B, leaf × 1/3; C, detail of spadix fertile portion and appendix base × 1; D, stamen × 10; E, gynoecium × 10; F, gynoecium, longitudinal section × 10; G, infructescence × 2/3; H, spadix × 1; J, stamen × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10. Dracunculus vulgaris: A, Fleming s.n. (Kew slide collection); B, Cult. Cheston (K); C–F, Boyce 19 (Kew spirit collection 51381); G, Cult. Kew (Kew spirit collection 25760); Zabeau 11/65 (Kew slide collection); D. canariensis: H–L, Cult. Boyce (Kew spirit collection 29047.679).

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B

E

A

F

G

C

D

Plate 91. Helicodiceros. A, habit × 1/3; B, detail of hairs on spathe limb interior × 1; C, spadix × 2/3; D, detail of spadix fertile portion and appendix base × 1; E, stamen × 10; F, gynoecium × 10; G, gynoecium, longitudinal section × 10. Helicodiceros muscivorus: A, Boyce 36 (Kew slide collection); B–G, Boyce 36 (Kew spirit collection 29047.638).

91. Helicodiceros

forming loose pseudostem. BLADE: deeply pedatifid, anterior division oblong, lobes of posterior divisions tightly spiralled on basal ribs and thus held ± erect; primary lateral veins of each lobe pinnate, weakly differentiated, forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: much shorter than petiole. SPATHE: spotted on outer surface, constricted between tube and blade, tube with convolute margins, erect, green, thick, subcylindric, blade sharply bent backwards at constriction, large, oblong-ovate, widely spreading, pale purple, inner surface covered with purple hairs. SPADIX: shorter than spathe, bent backwards at spathe constriction, female zone sessile to shortly stipitate, subcylindric, sterile zone separating male and female zones short, covered with subulate pistillodes or naked, male zone subcylindric to ellipsoid, shorter than or subequal to female, appendix vermiform, much longer than rest of spadix, weakly stipitate, thicker at middle, attenuate towards apex and base, ± lying on spathe blade, entirely covered with upwardly curved, setiform staminodial processes becoming gradually larger and subulate towards appendix base. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–3-androus,

AREAE : HELICODICEROS

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stamens free, anthers sessile, connective slender, inconspicuous, thecae ± ellipsoid, dehiscing by broad apical slit. POLLEN: extruded in strands, inaperturate, ± spherical, exine spinose. FEMALE FLOWER: ovary 1-locular, ovules up to 6, orthotropous, placentae 2, apical placenta with 3–4 ovules and basal placenta with 1–2 ovules, stylar region as broad or broader than ovary, stigma subhemispheric. BERRY: obovoid to ellipsoid, bearing small stigma remnant, (1–)2–3-seeded, orange. SEED: borne on either one or both placentae, broadly ovoid, with yellowish strophiole, testa leathery, strongly reticulate, pale brown when fresh, embryo axile, elongate, endosperm copious. See Plates 91, 127D. CHROMOSOMES: 2n = 56. DISTRIBUTION: 1 sp.; western Mediterranean Europe:– France (Corsica), Italy (Sardinia), Spain (Balearic Islands). ECOLOGY: warm temperate habitats; geophytes, in limestone and granite rock crevices beside the sea, rarely on stony ground. ETYMOLOGY: Greek “helix, helicos” (spiral), “dis” (twice) and “keras” (horn), referring to the two basal lobes of the leaf that are twisted and erect like horns. TAXONOMIC ACCOUNTS: Arcangeli (1884), Engler (1920a); Arrigoni et al. (1982, as Dracunculus muscivorus), Boyce (1994).

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92. Theriophonum Theriophonum Blume, Rumphia 1: 127 (1837, “1835”). TYPE: T. crenatum (R. Wight) Blume (Arum crenatum R. Wight). SYNONYMS: Tapinocarpus Dalzell in Hooker’s J. Bot. Kew Gard. Misc. 3: 345 (1851); Calyptrocoryne Schott in Oesterr. bot. Wochenbl. 7: 262 (1857); Pauella Ramamurthy & Sebastine in Bull. Bot. Surv. India 8: 348 (1967). HABIT: small, seasonally dormant herbs, tuber small, depressed-subglobose. LEAVES: several. PETIOLE: sheath relatively long. BLADE: sagittate-hastate or hastate, sometimes linear-lanceolate or oblong-ovate; primary lateral veins pinnate, forming submarginal collective vein, 1–2 distinct marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with the leaves. PEDUNCLE: usually short, erect in flower, deflexing in fruit. SPATHE: marcescent, constricted between tube and blade, tube convolute, blade much longer than tube, erect and gaping to spreading, or sometimes ± reflexed. SPADIX: sessile, shorter or subequal to spathe, female zone few-flowered, very short, separated from male by axis partly or completely covered with sterile flowers, sterile flowers either basal, central or at apex of sterile zone, male zone cylindric, sometimes with zone of sterile flowers above, appendix stipitate or not, erect, shorter or much longer than rest of spadix, cylindric, elongate-conoid to very long-subulate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–2-androus, stamens free, anthers subsessile, connective slender, sometimes elongated and prominent, thecae ovoid, dehiscing by apical pore or broad slit. POLLEN: inaperturate, spherical, medium-sized (mean 25 µm.), exine spinose. STERILE FLOWERS: spreading, somewhat decurved or suberect, long, slender and filiform to shorter and subulate. FEMALE FLOWER: gynoecia oriented vertically, ovary ellipsoid to obovoid, 1-locular, ovules 3–9 (or more), orthotropous, funicle short, placentae 2, apical and basal, stigma ± sessile, discoid-hemispheric, sometimes rather massive with central depression. BERRY:

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92. Theriophonum

ovoid to ellipsoid, few-seeded. SEED: broadly ovoid to ellipsoid, apiculate, strophiolate, testa smooth to rugose, embryo axile, elongate, endosperm copious. See Plate 92. CHROMOSOMES: 2n = 16 (14, 18). DISTRIBUTION: 5 spp.; India, Sri Lanka. ECOLOGY: tropical forest; geophytes, forest floor, grassy places, ditches, damp sites on rocky, lateritic soil. ETYMOLOGY: Greek name theriophonon from therion (wild beast) and phenô (I slay); means “beast-slayer”. TAXONOMIC ACCOUNTS: Engler (1920a), Sivadasan & Nicolson (1982).

93. Typhonium Typhonium Schott in Wiener Z. Kunst 1829 (3): 732 (1829). LECTOTYPE: T. trilobatum (L.) Schott (Arum trilobatum L.; see Nicolson in Taxon 16: 519. 1967). SYNONYMS: Desmesia Rafinesque, Fl. Tell. 3: 63 (1837, “1836”); Heterostalis (Schott) Schott in Oesterr. bot. Wochenbl. 7: 261 (1857). HABIT: very small to medium-sized, seasonally dormant or evergreen, rarely pubescent (T. hirsutum) or glandularpubescent herbs, tuber globose, subglobose or irregular, sometimes rhizomatous or stoloniferous, rarely an epigeal stem (T. fultum). LEAVES: few to several. PETIOLE: apex and middle portion rarely tuberculate, sheath rather short. BLADE: usually cordate-sagittate, sagittate to hastate, trifid, pedatifid or pedatisect, rarely linear, narrowly lanceolate, elliptic-oblong or cordate, apex rarely tuberculate; primary lateral veins of blade or lobes pinnate, forming submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with or without (T. hayatae) or after the leaves. PEDUNCLE: shorter than petiole. SPATHE: constricted between tube and blade, tube with convolute or rarely basally connate (T. hirsutum) margins, persistent or rarely evanescent (T. nudibaccatum), blade eventually bending backwards from constriction, broadly ovate to lanceolate, ± acuminate, usually purple, rarely white within, tube persistent, blade marcescent. SPADIX: sessile, shorter, subequal or much longer than spathe, female zone cylindric, subconic to subglobose, separated from male zone by rather long axis

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Plate 92. Theriophonum. A, habit × 2/3; B, detail of fertile portion of spadix and appendix base × 3; C, stamen with prominent connective × 15; D, gynoecium × 15; E, gynoecium, longitudinal section × 15; F, infructescence × 1; G, habit × 1; H, leaf × 1; J, leaf × 1; K, detail of fertile portion of spadix and appendix base × 4; L, detail of fertile portion of spadix and appendix base × 3; M, stamen × 15. Theriophonum minutum: A, Barnes 699 (K); Barnes 700 (K); Balasubramanian 2047 (K); Bogner s.n. (Kew slide collection); B–E, Barnes 698 (K & Kew spirit collection 58090); F, Nicolson 4266 (K); T. sivaganganum: G–K, Rajagopalan RHT 30294 (K & Kew spirit collection 58131); T. infaustum: L–M, Barnes 1333 (K & Kew spirit collection 58071).

AREAE : THERIOPHONUM

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Plate 93. Typhonium. A, spadix × 1 1/2; B, habit × 1/2; C, leaf × 1/2; D, spadix × 1 1/2; E, infructescence × 2/3; F, habit, part of petiole removed × 1/2; G, detail of fertile portion of spadix × 1 1/2; H, stamen × 10; J, gynoecium, side view × 10; K, gynoecium, longitudinal section × 10. Typhonium brownii: A, Thomson 2360 (Kew spirit collection 57319); T. hirsutum: B, comm. Mayo ‘Arisaema 1’ (Kew spirit collection 29047.710); T. trilobatum: C, D, Hay 2045 (Kew spirit collection 55956); E, Walker s.n. (K); T. giganteum: F, Cult. Cambridge Bot. Gard. (K), Henry 6537 (K); Hetterscheid H.AR. 31 (Kew spirit collection 57592); G–K, Hetterscheid H.AR. 31 (Kew spirit collection 57592).

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NOTES: Some species are weedy and naturalized in various parts of the world outside Asia in tropical, subtropical and warm temperate regions. Sriboonma, Murata & Iwatsuki (1994) recognized 5 sections:– sect. Hirsuta, sect. Diversifolia, sect. Pedata, sect. Gigantea, and sect. Typhonium. ETYMOLOGY: cited by Dodoens (1574) as Greek Typhonion, an ancient plant name; Typhon, the youngest child of Gaia, the Greek Earth goddess, was a beast, half man, half reptile, later slain by Zeus (A. Hay pers. comm.) TAXONOMIC ACCOUNTS: Engler (1920a), Nicolson & Sivadasan (1981), Sivadasan (1982), Bogner (1988b), Murata (1990b), Murata & Mayo (1991), Hay (1993a), Sriboonma, Murata & Iwatsuki (1994).

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94. Sauromatum

93. Typhonium

covered either entirely or only in basal part with sterile flowers of various shapes, rarely with sterile flowers above male flowers, male zone cylindric to ellipsoid, usually densely but rarely sparsely flowered (T. albispathum), appendix usually shortly stipitate, rarely with disc-like extension at the base or sessile (T. hirsutum), smooth, conoid to extremely slender and filiform-subulate, usually long-exserted. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–3-androus but usually 1-androus, stamens free or sometimes ± connate, anthers subsessile, connective slender, sometimes prominulent, thecae ovoid to ellipsoid, dehiscing by pore or lateral slit extending to the middle or nearly to the base and confluent apically. STERILE FLOWERS: either all similar or diverse in the same spadix, capitate, clavate to spathulate, cylindric, filiform or subulate, rarely stout and flattened at the tip, or reduced to verrucae, straight to flexuose, suberect, spreading or decurved. POLLEN: extruded in amorphous mass, inaperturate, spherical to subspheroidal, medium-sized (mean 32 µm., range 28–36 µm.), exine spinose (spines very obtuse in e.g. T. trilobatum). FEMALE FLOWER: gynoecia oriented horizontally or vertically, ovary ovoid, ellipsoid or obovoid, 1-locular, ovules 1–3, orthotropous, funicle short, placenta basal, stigma sessile, discoid-hemispheric. BERRY: ovoid, 1- or rarely 2-seeded, orange-red, green or white. SEED: globose to obnapiform, testa thin, rugulose to smooth, strophiolate, embryo axile, elongate, straight, endosperm copious. See Plates 93, 128A. CHROMOSOMES: 2n = 16, 18, 20, 26, 36, 52, 54, 65, >100 (14). DISTRIBUTION: 37 spp.; tropical south, southeast and east Asia, Malay Archipelago, Australasia:– Australia, Bangladesh, Bhutan, Burma, Cambodia, Caroline Is., China (incl. Taiwan, Tibet), India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sumatra, Timor), Japan, Laos, Malaysia (Borneo, Peninsula), Mariana Is., Mongolia, Nepal, Pakistan, Papua New Guinea, Philippines, Singapore, Sri Lanka, Thailand, Vietnam. Naturalized in Brazil (Northeast, Southeast), Comores, Cuba, Ghana, Ivory Coast, Madagascar, Martinique, Mauritius, South Africa, Surinam, Tanzania (Zanzibar), Trinidad, Venezuela, ECOLOGY: warm temperate, subtropical and tropical humid and seasonal forests, secondary forest, cultivated land; geophytes, forest floor, among rocks, wet sites, streamsides, grassy places.

Sauromatum Schott in Schott & Endlicher, Melet. Bot. 17 (1832). LECTOTYPE: S. guttatum Schott (= S. venosum (Aiton) Kunth, Arum guttatum Wallich 1831, non Salisbury 1796; see Nicolson 1967). SYNONYMS: [Stauromatum Endlicher, Ench. 128 (1841), orth. var.]; Jaimenostia Guinea & Gómez-Moreno in Guinea, Ensayo Geobot. 248 (1946) & in Anales Jard. Bot. Madrid 6 (2): 465 (1946). HABIT: small to medium-sized, seasonally dormant herbs, tuber subglobose or depressed-globose, sometimes large. LEAVES: usually solitary, rarely up to 3. PETIOLE: often spotted, sheath very short. BLADE: deeply pedatifid to pedatisect; primary lateral veins of lobes pinnate, forming submarginal collective vein, 1–2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with or without leaves, borne at ground level. PEDUNCLE: very short. SPATHE: marcescent, slightly constricted between tube and blade, tube with connate margins, ± cylindric and usually somewhat ventricose basally, blade much longer, narrowly oblong-lanceolate, erect at first then reflexed and spiralled-revolute, margins undulate, conspicuously spotted on inner surface. SPADIX: subequal to spathe, free, sessile, female zone cylindric, separated from male zone by much longer axis bearing sterile flowers in lower part and naked above, male zone short, cylindric, appendix elongated, very long-exserted, relatively slender, cylindric, smooth, obtuse, suberect then somewhat forward-curving. FLOWERS: unisexual, perigone absent. MALE FLOWER: few-androus, stamens free, anthers ± sessile, somewhat compressed, connective slender, thecae oblong-obovoid, dehiscing by broad, apical slit. POLLEN: extruded in amorphous mass, inaperturate, spherical to subspheroidal, medium-sized (mean 33 µm.), exine spinose. STERILE FLOWERS: consisting of distant, patent, terete, filiform, or clavate staminodes, apex rounded to obliquely truncate-disciform. FEMALE FLOWER: gynoecium subcylindric, ovary 1-locular, ovules 1–4, orthotropous, funicle very short, placenta basal, stylar region shortly attenuate or ± as broad as ovary, stigma discoid-hemispheric. BERRY: obpyramidal, red-purple, darker and corrugated at apex, densely congested in subglobose, sometimes partly hypogeal infructescence. SEED: obnapiform, testa thin, dark-spotted, smooth to rough, strophiole present, embryo axile, elongate and curved, endosperm copious. See Plates 94, 128B. CHROMOSOMES: 2n = 26, 52, 104. DISTRIBUTION: 2 spp.; tropical Africa, Arabian peninsula, tropical and subtropical southern Asia:– Angola, Bangladesh, Bhutan, Burma, Cameroon, Central African Republic, China

AREAE : SAUROMATUM

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Plate 94. Sauromatum. A, habit in flower × 1/5; B, habit × 1/5; C, leaf × 2/3; D, inflorescence × 2/3; E, spadix × 1; F, stamen × 20; G, gynoecium × 20; H, gynoecium, longitudinal section × 20; J, tuber in section, petiole base and mature infructescence × 1/2; K, juvenile plant × 1/2; L, inflorescence × 2/3; M, spadix × 1; N, stamen × 30; P, gynoecium × 30; Q, gynoecium, longitudinal section × 30. Sauromatum venosum: A, Barnes 1617; B, Andrews 6/80 (Kew slide collection); C, Andrews 27/6/82 (Kew slide collection); D, Robertson p/8/35/5 (Kew slide collection); E–H, Cult. Maw (Kew spirit collection 29047.122); J, Jarrett & Saldanha HFP 736 (K), Kultan s.n. (Kew spirit collection 6915); K, Aitchison 268 (K); S. brevipes: L–M, Templar s.n. (K); N–Q, Templar s.n. (Kew spirit collection 47636).

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94. Sauromatum

(Tibet, Yunnan), Equatorial Guinea (Bioko) Ethiopia, India, Kenya, Malawi, Nepal, Pakistan, Saudi Arabia, Tanzania, Uganda, Yemen Republic, ?Zaire, Zambia. ECOLOGY: tropical upland forest and grassland; geophytes or occasionally epiphytes, shady, humid sites, also in humus deposits on rocks. NOTES: Differs from Typhonium only in the consistently fused margins of the spathe tube and the curved embryo. ETYMOLOGY: Greek sauros (lizard) and shortened form of chrômaton (coloured); refers to lizard-like colour of the petiole. TAXONOMIC ACCOUNTS: Engler (1920a), Sivadasan (1982), Mayo (1985a).

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95. Lazarum Lazarum A. Hay in Bot. J. Linn. Soc. 109 (3): 427–434 (1992). TYPE: L. mirabile A. Hay HABIT: very small, seasonally dormant herb; tuber subglobose, whitish. LEAVES: 1–3, appearing after flowering, dark blueish green, subtended by deliquescent cataphylls. PETIOLE: sheath relatively long. BLADE: elliptic, base obtuse, apex apiculate; midrib distinct, primary lateral veins pinnate, running ± arcuately into margin near apex, marginal vein present, higher order venation reticulate. INFLORESCENCE: solitary, subtended by long cataphylls, appearing before the leaves. PEDUNCLE: very short, elongating in fruit. SPATHE: marcescent, slightly constricted at position of an annular septum through which the spadix passes and which separates the male zone from the female zone forming two chambers, swelling again for a short distance above the constriction; tube ± broadly ellipsoid, margins connate, not persistent in fruit; blade narrowly lanceolate, sharply bent up, erect, margins inrolled. SPADIX: sessile, only slightly longer than spathe, female flowers free, 5–7 in a single whorl, zone of sterile flowers contiguous with female zone, male zone separated from zone of sterile flowers by naked axis which passes through the central aperture of the spathe septum, male flowers thus situated in the upper spathe chamber;

appendix vermiform, slightly exceeding spathe, smooth, emerging from the mouth of the spathe tube and then upturned. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2-androus, stamens free, anthers sessile, thecae opening by apical pore. POLLEN: inaperturate, ellipsoid, medium-sized (36–40µm. × 28–32µm.), exine spinose, spinulae 1.4– 1.6µm. long and apex somewhat rounded. STERILE FLOWERS: slender, cylindric, obtuse, ascending at base, then recurved and appressed against the wall of spathe tube. FEMALE FLOWER: lageniform, ovary ovoid, 1-locular, ovule 1, orthotropous, placenta basal, stylar region short, attenuate, stigma subhemispheric. BERRY: ovoid, 1-seeded, minutely beaked at apex, reddish, infructescence with long peduncle, with 5–7 berries in a whorl. SEED: ovoid to subglobose, micropyle prominent, strophiolate, testa minutely verruculose, embryo axile, endosperm copious. See Plate 94. CHROMOSOMES: 2n = c. 78. DISTRIBUTION: 1 sp.; Australia (Northern Territory, Melville Island). ECOLOGY: tropical open forest; geophytes, forest floor, in laterite on ridges and in sandy soil. ETYMOLOGY: named after biblical Lazarus, in poetic reference to the morbid colour of the partly buried inflorescence, which emerges from a leafless subterranean tuber.

95. Lazarum

AREAE : LAZARUM

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Plate 95. Lazarum. A, habit × 2; B, inflorescence, nearside half and upper part of spathe removed × 6; C, habit, fruiting × 2; D, berry × 8. Lazarum mirabile: A–B, Jones 1695 (DNA spirit collection); C–D, Gallen 3 (DNA spirit collection).

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96. Biarum Biarum Schott in Schott & Endlicher, Melet. Bot. 17 (1832), nom. cons. TYPE: B. tenuifolium (L.) Schott (Arum tenuifolium L.), typ. cons. SYNONYMS: Homaïd Adanson, Fam. 2: 470 (1763), nom. rej.; [Homaida Rafinesque, Fl. Tell. 3: 63 (1837, “1836”), orth. var.]; Ischarum (Blume) Reichenbach, Deutsches Bot. Herbarienbuch, Nom. 32 (1841); Leptopetion Schott, Gen. Aroid. t. 8 (1858); Cyllenium Schott, Gen. Aroid. t. 9 (1858); Stenurus Salisb., Gen. Pl. 5 (1866). HABIT: seasonally dormant herbs, tuber subglobose, smooth. LEAVES: few to many. PETIOLE: often only weakly differentiated from blade, sheath long to short. BLADE: linear,

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oblong-lanceolate, oblong-ovate or -obovate, or ellipticoblong, margins smooth to strongly crispate; primary lateral veins pinnate, ± forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: 1(–2) in each floral sympodium, usually appearing before leaves, rarely with or directly after the leaves at the end of the vegetative period. PEDUNCLE: short, usually hypogeal. SPATHE: constricted between tube and blade, erect, eventually evanescent, tube cylindric to ventricose, hypogeal or held just above ground surface, margins connate near base or for most of length, rarely margins free (B. aleppicum), blade usually much longer, rarely shorter and reduced, expanded to gaping, linear to oblong-lanceolate or oblong, rarely campanulate or fornicate or subtriangular, usually dark brown-purple within, rarely white, pale yellowish or red-purple. SPADIX: sessile, shorter

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Plate 96. Biarum. A, leaf × 2/3; B, detail of fertile portion of spadix and appendix base × 2; C, stamen × 10; D, gynoecium × 10; E, gynoecium, longitudinal section × 10; F, berry, longitudinal section × 2; G, habit × 2/3; H, habit × 2/3; J, detail of fertile portion of spadix and appendix base × 2; K, habit × 2/3; L, inflorescence × 1; M, inflorescence × 1; N, detail of fertile portion of spadix and appendix base × 2; P, stamen, side view × 10; Q, stamen, view from below × 10; R, gynoecium × 10; S, gynoecium, longitudinal section × 10. Biarum pyrami var. pyrami: A, Davis 3844 (K); B–F, Mathew 11056 (Kew spirit collection 55694); B. tenuifolium subsp. tenuifolium: G, du Pavillon 189 (K); H, Davis 1138 (K); J, Archibald & Archibald 5904, Cult. Kew 1985–05490 (Kew spirit collection 49785); B. straussii: K, Hewer 1951 (K); B. davisii subsp. marmarisense: L, Baytop s.n. (Kew spirit collection 55936); B. ditschianum: M, Bonn accessions 22592 (Kew spirit collection 55925 & 59042); B. spruneri: N–S, Cult. Mathew (Kew spirit collection 51563).

AREAE : BIARUM

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96. Biarum

to much longer than spathe, female zone free, short, hemispheric to subhemispheric, subglobose, separated from male zone by rather long axis bearing sterile flowers at base or throughout or rarely naked, male zone 2–3 times longer than female, rarely with a short zone of sterile flowers above, appendix smooth, either ± slender, elongate-vermiform and not or hardly stipitate or thicker, cylindric to conoid and stipitate, rarely procurved and filiform, rarely with a short, basal zone of filiform processes. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1–2-androus, stamens free, anthers sessile, connective slender, apiculate and slightly prominent or not, thecae ellipsoid, with posterior microsporangia ± overtopping anterior ones, dehiscing by pore or lateral slit extending to the middle or nearly to the base and confluent apically. POLLEN: presented free or extruded in strands, grains inaperturate, spherical to subspheroidal, medium-sized (mean 30 µm., range 26–56 µm.), exine spinose or smooth. STERILE FLOWERS: consisting of either a single filiform, acute or subulate-conic projection, or rarely composed of 2–3 uncinate, recurved projections from a single receptacle. FEMALE FLOWER: ovary ovoid or oblongovoid, 1-locular, ovule 1, orthotropous, funicle very short, placenta basal, stylar region very shortly conic to elongated, stigma subhemispheric. BERRY: obovoid, white to dull purple, pericarp carnose, not juicy, infructescence globose to hemispheric, borne at or slightly below soil level. SEED: obovoid to subglobose, with obconic strophiole, testa smooth to irregularly rugose, thickish, embryo axile, more than half as long as endosperm, endosperm copious. See Plates 96, 128C. CHROMOSOMES: 2n = 16, 20, 22, 24, 26, 32, 36, 74, 96, 98. DISTRIBUTION: 22 spp.; Mediterranean Europe, Middle East, North Africa:– Albania, Algeria, Croatia, Egypt, Greece (incl. Crete and islands), Iran, Iraq, Israel, Italy (incl. Sardinia, Sicily), Jordan, Lebanon, Libya, Macedonia, Morocco, Portugal, Spain, Syria, Tunisia, Turkey. ECOLOGY: temperate and warm temperate scrub, woodland, open ground in fields, vineyards, pastures; geophytes, stony ground, under trees and shrubs, sandy areas, on limestone, rock crevices. NOTES: Boyce (in press 1) recognized 3 subgenera:– subgen. Biarum; subgen. Cyllenium; subgen. Ischarum (with 3 sections). ETYMOLOGY: Greek biaron, a name mentioned by Dodoens (1574). TAXONOMIC ACCOUNTS: Engler (1920a), Bogner & Boyce (1989), Boyce & Athanasiou (1991), Boyce (in prep.).

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Tribe Arisaemateae Tribe Arisaemateae Nakai, Ord. Fam. Trib. Nov. 221 (1943, “Arisaematieae”). Laticifers simple, articulated; tuber depressed-globose, rarely rhizomatous; leaf blade usually compound, trisect, pedatisect or radiatisect, rarely cordate or trifid, primary lateral veins of lobes or divisions pinnate, higher order venation reticulate; spathe tube usually cylindric, blade usually fornicate; spadix with terminal appendix, appendix often cylindric, clavate or elongate-tapering; flowers unisexual, perigone absent; ovary 1-locular, ovules orthotropous, placenta basal, style ± attenuate, stigma small; seed testa rough, endosperm copious.

97. Pinellia Pinellia Tenore in Atti Accad. Sci. Napoli 4: 69 (1839), nom. cons. TYPE: P. tuberifera Tenore, nom. illeg. (Arum subulatum Desfontaines, Cat. Pl. Horti Paris p. 385. 1829). SYNONYMS: Atherurus Blume, Rumphia 1: 135 (1837, “1835”); Hemicarpurus Nees, Delect. Sem. Horto Bot. Vratisl. 1839: [3] (1839) & in Linnaea 14 Litt.-Ber. 167 (1840). HABIT: seasonally dormant herbs, tuber subglobose. LEAVES: few. PETIOLE: sometimes bearing tubercles near base and at apex, sheath fairly long. BLADE: usually deeply trifid or trisect or pedatisect, or simply cordate, ovate or oblong; primary lateral veins of each lobe or division pinnate, forming submarginal collective vein, 1–2 distinct marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: shorter or longer than petiole. SPATHE: slightly constricted between tube and blade, tube convolute, narrowly ellipsoid to ovate, persistent, almost closed within by a transverse septum, gaping at base (P. tripartita), blade oblong-elliptic, boat-shaped, gaping, fornicate, green to purple. SPADIX: much longer than spathe, female zone adnate to spathe, separated from male zone by spathe septum and by short, free, naked part of spadix axis, male zone cylindric, short, terminal sterile appendix smooth, elongate-subulate, often sigmoid, long-exserted from spathe. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1androus, stamens sometimes united congenitally in pairs or groups of four, short, compressed laterally, anthers sessile,

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Plate 97. Pinellia. A, habit × 2/3; B, stamen × 20; C, gynoecium × 20; D, gynoecium, longitudinal section × 20; E, petiole bulbil × 4; F, infructescence × 2/3; G, leaf × 2/3; H, infructescence × 2/3; J, habit × 2/3; K, inflorescence, nearside half of spathe removed × 3; L, stamen × 20; M, gynoecium × 20; N, gynoecium, longitudinal section × 20. Pinellia ternata: A–D, Kirkham, Flanagan & Boyce 146, (Kew spirit collection 57610); Wilson 4100 (K); E, Hickin s.n. (K); P. tripartita: F, Oldham 821 (K); P. pedatisecta: G, Wang x–333 (K); H, Cult. Kew 1962–24001 (Kew spirit collection 37614); P. cordata: J, Faber 82 (K); K–N, Cult. Boyce (Kew spirit collection 51852 & 58114).

ARISAEMATEAE : PINELLIA

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97. Pinellia

connective slender, thecae ellipsoid, dehiscing by apical slit. POLLEN: extruded in amorphous mass, inaperturate, spherical or subspheroidal, small to medium-sized (mean 25 µm., range 21–29 µm.), exine spinulose. FEMALE FLOWER: ovary ovoid to ovoid-oblong, 1-locular, ovule 1, orthotropous, funicle very short, placenta basal, stylar region attenuate, stigma small, hemispheric. BERRY: oblong-ovoid, green. SEED: obnapiform to ellipsoid, testa irregularly verrucose-rugulose or smooth, embryo axile, elongate or very small and subglobose, endosperm copious. See Plates 97, 128D. CHROMOSOMES: 2n = 26, 52. DISTRIBUTION: 6 spp.; temperate east Asia:– China (incl. Taiwan), Japan (incl. Ryukyu Is.), Korea N. and S.; P. ternata is occasionally naturalized in Europe and North America. ECOLOGY: temperate woodland and forest; geophytes, on ground or on rocks, also on grassy banks and in fields as a weed (P. ternata). NOTES: Ulharz (1985) reports that P. tripartita is self-pollinating, probably anemophilous. ETYMOLOGY: named after Giovanni V. Pinelli (1535–1601). TAXONOMIC ACCOUNTS: Engler (1920a), Benzing (1969), Li (1979), Ulharz (1985, 1986), Boyce (1988), Rugh (1990).

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98. Arisaema Arisaema Martius in Flora 14: 459 (1831). TYPE: A. speciosum (Wallich) Martius ex Schott (Arum speciosum Wallich), lectotype selected by Pfeiffer, Nom. Bot. 1: 265. (1873). SYNONYMS: Dochafa Schott, Syn. Aroid. 24 (1856); Muricauda J.K. Small, Fl. Southeast U.S. 227 (1903); Flagellarisaema Nakai in J. Jap. Bot. 25: 5 (1950); Pleuriarum Nakai in J. Jap. Bot. 25: 5 (1950); Heteroarisaema Nakai in J. Jap. Bot. 25: 6 (1950); Ringentiarum Nakai in J. Jap. Bot. 25: 6 (1950). HABIT: seasonally dormant or evergreen herbs, small to fairly large, stem usually a depressed-globose tuber, producing tubercles or stolons, more rarely a branching,

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horizontal rhizome. LEAVES: 1–2, rarely 3, cataphylls often attractively mottled and spotted. PETIOLE: sheaths usually rather long and imbricate to form a long, usually mottled pseudostem, margins either free or fused from base to apex and then with a fringed, ligulate mouth. BLADE: usually compound, trisect, radiatisect, pedatisect, very rarely simple and ovate, lobes 3–19 or sometimes more, usually lanceolate-elliptic, varying from linear to broadly ovate, elliptic or obovate, sometimes rhomboid, sessile or central leaflet often stalked, margin entire, serrate, erose or coarsely dentate; primary lateral veins of each lobe or division pinnate, forming submarginal collective vein, 1–2 conspicuous outer marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, usually appearing with the leaves, sometimes before them, often subtended by conspicuous cataphylls, many species producing male and female inflorescences successively from the same plant at different seasons (paradioecy). PEDUNCLE: very short to longer than petiole, rarely tuberculate in upper portion (A. scortechinii), sometimes decurved in fruit. SPATHE: marcescent, usually unconstricted, rarely slightly constricted (e.g. A. flavum), lower part erect, convolute into cylindric, often longitudinally striped tube, tube mouth often with revolute to broadly or even grotesquely auriculate margins, blade usually strongly fornicate, sometimes erect, widely expanded to galeate, apex acute to long-acuminate, sometimes drawn into very long filiform, erect or drooping thread. SPADIX: usually sessile, sometimes shortly stipitate, free, unisexual or monoecious, female or monoecious spadices more robust and differing from male in appendix shape and presence of sterile organs, female zone densely flowered, usually conoid, male zone usually laxly flowered and contiguous with female in monoecious inflorescences, sterile terminal appendix erect, procurved or pendent, entirely hidden within spathe to very long-exserted, usually somewhat longer than spathe tube, stipitate or not, cylindric, clavate, rounded, rugose or apically echinate or drawn out into a sometimes very long thread, rarely lacking altogether (A. exappendiculatum), often with a few subulate to filiform projections in basal part, more rarely entirely composed of long filiform projections. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–5-androus, filaments connate forming synandria, synandria ± distant from one another, sessile to long-stipitate, connective slender, usually inconspicuous, thecae shortly ovoid, dehiscing by short to long slit or pore, sometimes confluent into ± lunate or even circular compound thecae dehiscing by a single slit. POLLEN: very dry and powdery, inaperturate, spherical or subspheroidal, small (mean 22 µm., range 17–39 µm.), spinose. FEMALE FLOWER: ovary 1-locular, ovoid or oblong-ovoid, ovules 3–10, orthotropous, erect, funicle short, placenta basal, stylar region short to attenuate, always narrower than ovary, stigma usually rather small, subhemispheric. BERRY: obovoid to obconic, rounded apically, rarely conical, usually few-seeded, bright red, rarely yellow, glossy. SEED: ovoid to globose, bearing a strophiole, testa hard, rough, light brown, embryo axile, endosperm copious. See Plates 98i–iv, 129A. CHROMOSOMES: 2n = 20, 24, 26, 28, 39, 42, 48, 52, 56, 70, 72, 112, 140, 168 (64). DISTRIBUTION: ca. 170 spp.; temperate and warm temperate eastern and southeastern North America, northern Mexico, tropical east and northeast Africa, Arabian Peninsula, temperate east Asia, tropical south, southeast and east Asia, Malay Archipelago:– Afghanistan, Bhutan, Brunei, Burma,

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Plate 98 (i). Arisaema. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2. Arisaema candidissimum: A, Cult. Kew 1986–3895 (Kew slide collection); A. grapsospadix: B, Cult. Kew 1987–2421 (Kew slide collection), Murata s.n. (Kew slide collection); A. consanguineum: C, Cult. Kew 1985–854 (Kew slide collection); A. heterophyllum: D, Cult. Kew 1987–2226 (Kew slide collection); A. abei: E, Cult. Kew 1987–3585 (Kew slide collection).

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Plate 98 (ii). Arisaema. A, habit × 1/4; B, detail of inflorescence emergence from pseudostem ape× × 2/3; C, habit × 1/4; D, habit × 1/4; E, habit in flower × 1/4. Arisaema abei: A–B, Cult. Kew 1987–3585 (Kew slide collection); A. grapsospadix: C, Cult. Kew 1987–2421 (Kew slide collection); Murata s.n. (Kew slide collection); A. kiushianum: D, Murata s.n. (Kew slide collection); A. candidissimum: E, Cult. Kew 1986–3895 (Kew slide collection).

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Plate 98 (iii). Arisaema. A, male inflorescence × 2/4; B, male spadix × 2; C, synandrium × 10; D, female spadix × 2; E, gynoecium × 10; F, gynoecium, longitudinal section × 10; G, male inflorescence × 2/3; H, male spadix, most of appendi× removed × 2; J, synandrium × 10; K, male inflorescence × 2/3; L, male spadix × 2; M, synandrium × 10; N, female inflorescence × 2/3; P, female spadix × 2; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, female inflorescence × 2/3. Arisaema nikoense var. nikoense: A–F, Cult. Kew 1987–2861 (Kew spirit collection 55951 & Kew slide collection); A. griffithii: G–J, Cult. Kew 1988–466 (Kew spirit collection 55695 & Kew slide collection); A. taiwanense: K–M, Cult. Kew 1987–3583 (Kew spirit collection 49912 & Kew slide collection); A. ringens: N–R, Cult. Kew 1987–3583 (Kew spirit collection 49912 & Kew slide collection); A. tortuosum: S, Cult. Kew 1984–4797 (Kew slide collection).

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Plate 98 (iv). Arisaema. A, male inflorescence × 2/3; B, male spadix × 1; C, stamen × 10; D, female spadix × 2; E, gynoecium × 10; F, gynoecium, longitudinal section × 10; G, monoecious inflorescence × 2/3; H, monoecious spadix × 2; J, stamen × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10; M, female inflorescence × 2/3. Arisaema exappendiculatum: A–F, Bown 184/26 (Kew slide collection); B–F, Cult. Kew 1969–9202 (Kew spirit collection 49914); A. flavum subsp. flavum: G–L, Cult. Kew 1968–65 (Kew spirit collection 54924 & Kew slide collection); A. fimbriatum: M, Rubeli s.n. (Kew slide collection).

98. Arisaema

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Burundi, Cambodia, Canada, China (incl. Taiwan), Ethiopia, India, Indonesia (Borneo, Flores, Java, Sulawesi, Sumatra, Timor), Japan, Kenya, N. and S. Korea, Laos, Malaysia (Borneo, Peninsula), Mexico, Nepal, Oman, Pakistan, Philippines, Russia (Far East), Rwanda, Saudi Arabia, Somalia, Sri Lanka, Sudan, Tanzania, Thailand, Uganda, USA, Vietnam, Yemen Republic, Zaïre. ECOLOGY: temperate, subtropical and upland tropical forest, more rarely savanna, lowland tropical forest (Sarawak), subdesert or montane grassland (up to 4500m alt.); geophytes, forest floor, rocky slopes, rarely in wet places, very rarely epiphytic. NOTES: Murata (1990a) recognized 11 sections:– Fimbriata, Decipientia, Trisecta, Franchetiana, Pedatisecta, Clavata, Tortuosa, Dochafa, Tenuipistillata, Sinarisaema and Arisaema. ETYMOLOGY: Greek aris, aridos (name of a small herb mentioned by Pliny) and haima, haimatos (blood). TAXONOMIC ACCOUNTS: Engler (1920a), Hara (1971), Li (1979, 1980), Ohashi & Murata (1980), Sivadasan (1982), Murata (1984, 1987, 1990a), Mayo & Gilbert (1986), Pradhan (1990).

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Tribe Colocasieae Tribe Colocasieae Engler in Nova Acta Acad. Leopold.Carol. 39: 149 (1876). Laticifers anastomosing (except Ariopsis); primary lateral veins of leaf blade or leaflets (Protarum) pinnate, forming submarginal collective vein, at least 1–2 other marginal veins also present, higher order venation intermediate between parallel-pinnate and reticulate; spathe usually constricted centrally (except Ariopsis) with lower, convolute, persistent tube and upper, gaping, ± boat-shaped, marcescent to caducous blade; flowers unisexual, perigone absent; male flower a synandrium of several connate stamens, common connectives thick, fleshy, thecae lateral, pollen inaperturate; endosperm copious.

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99. Ariopsis Ariopsis Nimmo in J. Graham, Cat. Pl. Bombay 252 (1839). TYPE: A. peltata Nimmo Laticifers simple, articulated. HABIT: small, seasonally dormant herbs, tuber ± subglobose, hypogeal. LEAVES: usually solitary, rarely few. PETIOLE: slender, sheath fairly short. BLADE: peltate, cordate-ovate or only emarginate basally, thin, glaucous below, posterior lobes very short; primary lateral veins pinnate and also radiating from petiole insertion, forming submarginal collective vein, marginal vein also present, higher order venation reticulate. INFLORESCENCE: 1–3 in each floral sympodium, appearing with or without leaves. PEDUNCLE: very slender, much longer than spathe, erect. SPATHE: ovate, boat-shaped, fornicate, not constricted, gaping widely, not convolute at base, marcescent. SPADIX: shorter than spathe, female zone adnate to spathe, very short and few-flowered, sometimes separated from male zone by short, free, naked axis, male zone fertile to apex, relatively thick, cylindric-conoid, many-flowered. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium peltate, connate filaments forming a stipe longer and narrower than dilated common connective, thecae subglobose to ellipsoid,

99. Ariopsis

dehiscing by oval pore, synandria all connate apically, forming continuous surface punctured by cavities with somewhat prominent margins into which pollen is shed from the 6(–8) surrounding thecae (each pair of thecae belonging to a different synandrium). POLLEN: inaperturate, spherical or subspheroidal, small (mean 20 µm.), exine spinose. FEMALE FLOWER: ovary ovoid to ovoid-oblong, 1-locular, ovules many, orthotropous, placentae 4–6, parietal, extending from base to apex of locule, stylar region absent, stigma stellate with 4–6laciniate lobes, lobes initially erect, later spreading and reflexed. BERRY: 4–6-angled, stigma persistent, many-seeded. SEED: oblong, apically narrowed and obtuse, with indistinct strophiole, testa thickish, longitudinally costate, embryo axile, small, endosperm copious. See Plates 99, 129B. CHROMOSOMES: 2n = 28, 84. DISTRIBUTION: 2 spp.; tropical south Asia:– Bhutan, Burma, India (Assam, Sikkim, Western Ghats), Nepal. ECOLOGY: tropical evergreen forest; geophytes on forest floor or in rock crevices. NOTES: The correct morphological interpretation of the male flowers remains unclear. ETYMOLOGY: Greek aron (Arum) and -opsis (appearance). TAXONOMIC ACCOUNTS: Engler (1920a), Nicolson (1976), Sivadasan (1982), Bogner & Boyce (in prep.).

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100. Protarum Protarum Engler in Bot. Jahrb. 30, Beibl. 67: 42 (1901). TYPE: P. sechellarum Engler HABIT: seasonally (?irregularly) dormant herb, tuber subglobose, with numerous, very densely arranged, annular leaf scars. LEAF: solitary. PETIOLE: long, sheath short. BLADE: pedatisect to radiatisect, almost radiate, leaflets shortly stalked, elliptic to narrowly elliptic, acuminate; primary lateral veins of lobes pinnate, numerous, forming submarginal collective vein, 2 marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, appearing with or without leaves. PEDUNCLE: short, much shorter than petiole. SPATHE: slightly constricted between tube and blade, tube convolute, shorter than blade, blade erect, boat-shaped, marcescent. SPADIX: sessile, much shorter than spathe, female zone conoid, sep-

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Plate 99. Ariopsis. A, habit × 1; B, spadix × 6; C, gynoecium × 15; D, gynoecium, longitudinal section × 15; E, section through male portion of spadix to show synandrium arrangement × 10; F, infructescence × 6; G, habit in flower, showing branching of tubers × 1; H, leaf × 1; J, spadix × 6; K, berry × 6; L, seed × 12. Ariopsis peltata: A, Talbot 496 (K); B–E, Bogner 1922 (Kew spirit collection 56425); F, Barnes 1087 (K); A. protanthera: G–H, Cult. Kew.1851 (K); J, Kurz s.n. (K); K–L, Cult. Kew. 1851 (Kew spirit collection 58040).

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Plate 100. Protarum. A, leaf × 1/6; B, leaflet × 1/3; C, tuber × 1/2; D, inflorescence × 2/3; E, spadix × 2; F, detail of gynoecia, top view × 8; G, synandrium, top view × 15; H, gynoecium with associated staminode, longitudinal section × 15; J, infructescence, nearside half of spathe removed × 2/3; K, seed × 4. Protarum sechellarum: A, 5495 (Kew slide collection); B, Milne 14 (K); C, Jeffrey 485 (Kew spirit collection 25047); D, Thomasset s.n. (K); E–H, Whitehead 35 (Kew spirit collection 37323); J–K, Gardiner 110 (K & Kew spirit collection 58062).

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100. Protarum

101. Steudnera

arated from male zone by more slender sterile zone covered with narrowly elongated synandrodes or partly naked, male zone ± cylindric to obconic, terminal sterile appendix thicker, digitiform, obtuse, ± smooth. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 3–6-androus, sessile, subprismatic, ± excavated centrally, fused connectives broad, margins lobed, thecae broadly ellipsoid to subglobose, dehiscing by broad apical slit. POLLEN: inaperturate, ellipsoid, small (21 µm., range 20–22 µm.), exine striate. FEMALE FLOWER: gynoecium surrounded by whorl of 4–6, free, ± imbricate, thickish, oblong, subprismatic staminodes, ovary ovoid to ellipsoid, 1-locular, ovules 4, hemiorthotropous, micropyle elongate, funicle short, placenta basal, stylar region shortly attenuate or nearly absent, stigma 2–4-lobed, much broader than style. BERRY: ± ellipsoid to obovoid, with persistent stigma remnant, usually 1-seeded, orange, infructescence enclosed by persistent and much enlarged spathe tube. SEED: ovoid to ellipsoid-oblong, testa costate, dark brown, embryo small, flattened and disc-like, endosperm copious. See Plate 100. CHROMOSOMES: 2n = 28. DISTRIBUTION: 1 sp.; Seychelles Islands (Mahé, Praslin, Silhouette). ECOLOGY: tropical (palm) forest, moss forest; geophytes, forest floor, leaf litter in rock crevices, often on large granitic rocks. ETYMOLOGY: Greek protos (first) and Arum. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1973b), Robertson (1989).

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101. Steudnera Steudnera K. Koch in Wochenschr. Gärtnerei Pflanzenk. 5: 114 (1862). TYPE: S. colocasiifolia K. Koch (“colocasiaefolia”). HABIT: medium sized to robust herbs, usually evergreen, sometimes seasonally dormant, stem epigeal, stout, erect or creeping, covered with fibrous remains of leaves and cata-

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phylls. LEAVES: solitary or few to several. PETIOLE: sheath very short. BLADE: peltate, ovate or ovate-oblong, acuminate, often emarginate at base; basal ribs short, well-developed, primary lateral veins pinnate, forming submarginal collective vein very near margin, marginal vein also present, secondary and tertiary laterals arising from the primaries at a wide angle, then arching towards leaf margin and forming ± conspicuous interprimary collecting vein, higher order venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: shorter than petiole. SPATHE: yellow or ± dark purple within, ovate or ovate-lanceolate, acuminate, not constricted, barely convolute at base, ± fully expanded at anthesis, lower part persistent to fruiting stage, upper part becoming reflexed and revolute, marcescent. SPADIX: much shorter than spathe, densely flowered, female zone cylindric, often longer than male, mostly adnate to spathe, male zone contiguous with female, cylindric, ellipsoid or subglobose, fertile to apex, obtuse. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–6-androus, stamens connate, synandrium strongly lobed, apically truncate, common connective relatively small, impressed at apex, thecae oblong, contiguous, dehiscing by apical pore. POLLEN: inaperturate, ellipsoid-oblong, small (mean 22 µm.), exine striate. FEMALE FLOWER: gynoecium surrounded by whorl of 2–5, short, claviform staminodes, more rarely staminodes absent, ovary subglobose to ovoid, 1-locular, ovules numerous, hemiorthotropous, funicle distinct, placentae 2–5, parietal, in basal part only or extending from base to apex, stylar region ± lacking, stigma strongly 2–5-lobed. BERRY: ovoid, manyseeded. SEED: ovoid to ellipsoid, testa costate, embryo axile, short, conoid, endosperm copious. See Plates 101, 129C. CHROMOSOMES: 2n = 28, 56. DISTRIBUTION: 8 spp.; tropical south and southeast Asia:– Bangladesh, Burma, ?Cambodia, China (Guangxi, Yunnan), India (Assam), Laos, Thailand, Vietnam. ECOLOGY: tropical humid forest; geophytes on forest floor. ETYMOLOGY: named after H. Steudner (1832–1863). TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920), Boyce (1995c).

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Plate 101. Steudnera. A, habit × 2/3; B, spadix × 2; C, synandrium, top view × 15; D, synandrium, side view × 15; E, gynoecium, top view × 15; F, gynoecium, longitudinal section × 15; G, inflorescence, nearside half of spathe removed × 2/3; H, synandrium, side view × 15; J, gynoecium with associated staminode × 15. Steudnera henryana: A, Henry 11986 (K); B–F, S. discolor: Bogner 1814 (Kew spirit collection 57576); S. colocasioides: G–J, Cult. Kew (K & Kew spirit collection 58060).

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102. Remusatia Remusatia Schott in Schott & Endlicher, Melet. Bot. 18 (1832). TYPE: R. vivipara (Roxburgh) Schott (Arum viviparum Roxburgh, “viviparium”). SYNONYM: Gonatanthus Klotzsch in Link, Klotzsch & Otto, Icon. Pl. Rar. Hort. Berol. 1: 33 (1841). TYPE: G. sarmentosus Klotzsch HABIT: small to medium-sized, seasonally dormant herbs, tuber subglobose, producing erect to spreading, unbranched or branching stolons from axils of scarious, deciduous cataphylls, stolons producing small, ovoid tubercles at nodes, each invested by numerous, apically revolute or flexuose scales (minute cataphylls). LEAVES: 1–2. PETIOLE: sometimes slender, sheath relatively short. BLADE: peltate, cordate-lanceolate to cordate-ovate, acuminate; basal ribs well-developed, primary lateral veins pinnate, forming submarginal collective vein very close to margin, marginal vein also present, secondary and tertiary laterals arising from the primaries at a wide angle, then arching towards leaf margin and forming inconspicuous interprimary collective veins, higher order venation reticulate. INFLORESCENCE: solitary, appearing with or without leaf. PEDUNCLE: shorter than petiole. SPATHE: strongly constricted between tube and blade, sometimes with secondary constriction above the spadix, tube with convolute margins, persistent, enclosing female zone and sterile zone of spadix, blade yellow or red, longer than tube, fully expanded or remaining convolute and opening only at base, sometimes becoming reflexed (R. vivipara, R. yunnanensis), later deciduous. SPADIX: sessile or subsessile, much shorter than spathe, female zone subcylindric, about half as long as spathe tube, separated from male zone by much narrower zone of sterile male flowers, male zone ellipsoid or subclavate, fertile to apex, obtuse. FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–3-androus, stamens connate into cuneate-clavate, 4–6-sulcate synandrium, fused filaments distinct, common connectives somewhat excavated at apex, thecae 4–6, oblong to ellipsoid, dehiscing by api-

102. Remusatia

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cal pore-like slit. POLLEN: extruded in strands, inaperturate, spherical to subspheroidal, medium-sized (mean 32–33 µm.), spinose. STERILE MALE FLOWERS: each a ± elongated synandrode. FEMALE FLOWER: staminodes absent, ovary subcylindric to subglobose, 1-locular or partially 2–4-locular at apex, ovules many, hemiorthotropous, funicle short to long, placentae 2–4 and parietal or placenta 1 and basal, stylar region very shortly attenuate or lacking, stigma discoid-subcapitate or slightly 3–4-lobed. BERRY: obovoid to globose, many-seeded, infructescence ellipsoid, borne within persistent spathe tube. SEED: ellipsoid to subglobose, covered by thick, succulent sarcotesta or testa verruculose to irregularly costate, embryo axile, short, ovoid to subglobose, endosperm copious. See Plates 102, 129D. CHROMOSOMES: 2n = 28, 42, 56. DISTRIBUTION: 4 spp.; tropical Africa, tropical Asia, Malay Archipelago, Australasia: Australia (N.), Bangladesh, Bhutan, Cameroon, China (Taiwan, Yunnan, Tibet), Christmas Is., Ethiopia, Guinea, India, Indonesia (Java), Ivory Coast, Liberia, Madagascar, Nepal, Nigeria, Oman, ?Saudi Arabia, Sierra Leone, Sri Lanka, Tanzania, Thailand, Vietnam, Yemen Republic (Socotra), Zaïre, Zambia. ECOLOGY: tropical seasonal forest; epiphytes, lithophytes or geophytes, moss-laden boughs, rock crevices, damp banks, forest floor. NOTES: Li & Hay (1992a, b) recognize 2 sections, sect. Remusatia and sect. Gonatanthus. ETYMOLOGY: named after A. Rémusat (1788–1832). TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920), Sivadasan (1982), Mayo (1985a), Li (1987a, b, 1991, 1992), Li & Hay (1992a, 1992b).

103. Colocasia Colocasia Schott in Schott & Endlicher, Melet. Bot. 18 (1832), nom. cons. TYPE: C. antiquorum Schott (Arum colocasia L.), typ. cons. SYNONYM: Leucocasia Schott in Oesterr. bot. Wochenbl. 7: 34 (1857).

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Plate 102. Remusatia. A, habit in flower with stolons × 1/4; B, habit with stolons × 1/4, C, leaf × 2/3; D, tubercle × 6; E, plantlet emerging from tubercle × 2/3; F, inflorescence × 2/3; G, spadix × 2; H, synandrium, top view × 12; J, synandrium, side view × 12; K, synandrode × 15; L, gynoecium, longitudinal section × 12; M, gynoecium, transverse section × 12; N, habit × 1/3; P, inflorescence × 2/3; Q, gynoecium, longitudinal section × 12. Remusatia vivipara: A, 299 (Kew slide collection); B, 300 (Kew slide collection); C–D, Kerr 1438 (K); E, Cult. Kew 1969–5188 (Kew spirit collection 59039); F–M, Bogner 2121 (Kew spirit collection 57570); Cult. Kew 1947–85 (Kew spirit collection 19155); R. pumila: N, Grey-Wilson & Phillips 128 (K); P, C.K. 227 (K); Q, Grierson & Long 2215 (K).

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Plate 103. Colocasia. A, habit × 1/2; B, inflorescence × 2/3; C, synandria, top view × 5; D, gynoecium, longitudinal section × 11; E, floral sympodium with associated petiole sheath × 1/3; F, spadix × 1; G, synandria, top view × 5; H, synandrium, side view × 10; J, gynoecium, longitudinal section; K, gynoecium, transverse section; L, habit × 1/5. Colocasia fallax: A, Gamble 27041 (K); B–D Chantaranothai et al. 90/464 (Kew spirit collection 59076); C. gigantea: E, Bown 197/26 (Kew slide collection); F–K, Bogner 427 (Kew spirit collection 34491); C. esculenta: L, Townsend s.n. (Kew slide collection).

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HABIT: small, medium-sized or gigantic, seasonally dormant or evergreen herbs, stem either a hypogeal, subglobose or subcylindric tuber or mostly epigeal, massive. LEAVES: several, rosulate in acaulescent plants, forming terminal crown in arborescent species. PETIOLE: sheath rather long. BLADE: peltate, ovate-cordate to sagittate-cordate, posterior lobes rounded, shortly to almost entirely connate; basal ribs welldeveloped, primary lateral veins pinnate, forming submarginal collective vein, 1–2 marginal veins also present, secondary and tertiary laterals arising from the primaries at a wide angle, then arching strongly towards leaf margin and forming inconspicuous interprimary collective veins, higher order venation reticulate. INFLORESCENCE: 1–many in each floral sympodium, appearing with the leaves. PEDUNCLE: much shorter than petiole. SPATHE: constricted between tube and blade, sometimes with a second constriction above male zone of spadix, tube with convolute margins, usually much shorter than blade, ovoid or oblong, persistent, enlarging in fruit and then splitting open irregularly, blade white to yellow, oblong and boat-shaped to narrowly lanceolate, reflexing at anthesis, later deciduous. SPADIX: sessile, shorter than spathe, female zone short, separated from male by narrower zone of sterile male flowers, male zone cylindric to fusiform, terminal appendix erect, ± smooth, elongate-conoid to fusiform or subulate, sometimes reduced to small, mucronate stub, rarely absent. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–6-androus, stamens connate into ± truncate synandrium, thecae lateral, oblong-linear, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, ellipsoid-oblong or spherical to subspheroidal (C. esculenta), small to medium-sized (mean 25 µm., range 25–26 µm), exine finely striate, coarsely swirling-fossulate or fossulatespinose. STERILE MALE FLOWERS: synandrodes depressed-obpyramidal, truncate, laterally compressed. FEMALE FLOWER: ovary ovoid or oblong, 1-locular, ovules many, hemiorthotropous, funicles relatively long, placentae 2–5, parietal, stylar region short, narrowed or not, sometimes ± absent, stigma discoid-capitate or weakly lobed. BERRY: greenish to whitish or red(?), obconoid or oblong, stigma remnants persistent, many-seeded. SEED: ovoid to ellipsoid,

103. Colocasia

testa thickish, costate, embryo axile, cylindric, endosperm copious. See Plates 103, 130A. CHROMOSOMES: 2n = 28, 42, 56. DISTRIBUTION: 8 spp.; tropical Asia, Malay Archipelago:– Bangladesh, Bhutan, Burma, Cambodia, China (Guandong, Guangxi, Guizhou, Hainan, Hunan, Sichuan, Taiwan, Yunnan), India, Indonesia (Java, Lesser Sunda Is.), Laos, Malaysia (Peninsula), Nepal, Sri Lanka, Thailand, Vietnam. The country list given here does not include C. esculenta. This species is cultivated and naturalized throughout the humid tropics and subtropics. Its natural area of distribution remains somewhat obscure but is certainly tropical Asian, extending into the Malay Archipelago and perhaps as far as Papuasia and Australia. ECOLOGY: tropical humid forest habitats; geophytes, terrestrial or helophytes, wet places, along streams and ponds, forest floor in leaf litter, between rocks, sometimes on limestone (C. gigantea). NOTES: Krause in Engler & Krause (1920) recognized 2 sections:– sect. Colocasia (“Tuberosae”), sect. Caulescentes. VERNACULAR NAMES AND USES: taro, eddoe; C. esculenta is an important food plant in tropical regions, especially the Pacific. ETYMOLOGY: classical name, Greek kolokasia, from an old Middle Eastern name qolqas. TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920); Sivadasan (1982), Plucknett (1983), Shaw (1984), Sreekumari & Mathew (1991a, b), Li & Wei (1993).

C

104. Alocasia Alocasia (Schott) G. Don in Sweet, Hort. Brit., ed. 3: 631 (1839), nom. cons. TYPE: A. cucullata (Loureiro) G. Don (Arum cucullatum Loureiro), typ. cons. SYNONYMS: Colocasia sect. Alocasia Schott in Schott & Endlicher, Melet. Bot. 18 (1832); Ensolenanthe Schott in Bonplandia 9: 368 (1861); Xenophya Schott in Ann. Mus. Bot. Lugduno–Batavum 1: 124 (1863); Schizocasia Schott ex Engler in Bot. Jahrb. 1: 185 (1880, “1881”); Panzhuyuia Z.Y. Zhu in J. Sichuan Chinese Medicinal School 4 (5): 49 (1985).

104. Alocasia

COLOCASIEAE : ALOCASIA

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C

A

F

D E

B

K J

G

M

H

L

Plate 104 (i). Alocasia. A, leaf × 1/3; B, inflorescence × 1; C, synandrium, side view × 8; D, gynoecium, longitudinal section × 8; E, leaf × 1/3; F, habit × 1/6; G, infructescence with associated petiole and plant base × 1/8; H, spadix × 2; J, synandrium × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10; M, habit × 2/3. Alocasia brancifolia: A, Cult. Veitch 10 June 1884 (K); B–D, Bogner 1736 (Kew spirit collection 58916); A. guttata: E, Forman 931 (K); A. lowii: F–G, Boyce 384 (Kew slide collection); H–L, Boyce 384 (Kew spirit collection 49923); A. beccarii : M, Simpson 2271 (K); Wood 798 (Kew spirit collection 52879).

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D

E

C

F

B

A

Plate 104 (ii). Alocasia. A, habit × 2/3; B, spadix × 2; C, synandrium, side view × 15; D, gynoecium × 15; E, gynoecium, longitudinal section × 15; F, infructescence. Alocasia beccarii: Alston 14176 (BM).

HABIT: medium-sized to rarely arborescent and gigantic, seasonally dormant to evergreen herbs, stem thick, often hypogeal, sometimes stoloniferous and bulbiferous, epigeal stem usually erect, rarely elongated and creeping, with clear to milky latex. LEAVES: few to several in terminal crown, sometimes each subtended by a cataphyll. PETIOLE: long, sometimes asperate or glandular, sheath relatively long, sometimes deciduous. BLADE: sometimes pubescent, juvenile blade peltate, at maturity usually sagittate, less often ± hastate or cordate, peltate in some species, margin entire, sinuate or slightly to deeply pinnatifid, posterior divisions ovate or triangular; basal ribs well-developed, glands present in axils of primary lateral veins and midrib, primary lateral veins pinnate, forming submarginal collective vein,1–2 closely adjacent marginal veins also present, secondary and tertiary lateral veins arising from the primaries at a wide angle, then arching strongly

towards leaf margin, sometimes forming interprimary veins, higher order venation reticulate. INFLORESCENCE: 2–many in each floral sympodium, appearing with the leaves. PEDUNCLE: usually shorter than petiole. SPATHE: strongly constricted between tube and blade, tube with convolute margins, shorter than blade, ovoid or oblong, persistent and then splitting irregularly in fruit, blade oblong, usually boat-shaped, rarely fornicate, at anthesis at first erect, then reflexing and later usually deciduous; in Xenophya group blade is persistent, erect, convolute, gaping only basally. SPADIX: sessile, sometimes shortly stipitate, rarely obliquely adnate, shorter than spathe, female zone short, conoidcylindric, separated from male by a much narrower zone of sterile flowers, male zone usually cylindric, appendix conoid to cylindric, surface with irregular, labyrinthine network of fissures. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–12(–36)-androus, stamens connate into

COLOCASIEAE : ALOCASIA

285

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an obpyramidal, subhexagonal, truncate, rarely linear (A. brisbanensis) synandrium, thecae oblong to linear-oblong, lateral, dehiscing by apical pore. POLLEN: extruded in strands, inaperturate, spherical to subspheroidal, mediumsized (mean 35 µm., range 31–39 µm.), exine spinose. STERILE MALE FLOWERS: synandrodes shallow, obpyramidal, compressed, truncate. FEMALE FLOWER: ovary ovoid or oblong, 1-locular or partially 3–4-locular at apex, ovules 6–10, orthotropous, hemiorthotropous, hemianatropous or anatropous, funicle short, placenta basal, stylar region short, stigma depressed-capitate, ± distinctly 3–4-lobed. BERRY: generally reddish, ellipsoid or obconic-ellipsoid or subglobose, 1–5-seeded, stigma remnants persistent. SEED: subglobose to ellipsoid, testa thickish, smooth or scabrous, embryo broadly conoid, shortly cylindric or elongate, endosperm copious. See Plates 104i–ii, 130B. CHROMOSOMES: 2n = 28, 42, 56, 70, 84. DISTRIBUTION: 60–70 spp.; tropical Asia, Australasia, Malay Archipelago, Melanesia:– Australia (Queensland, New South Wales), Bangladesh, Brunei, Burma, Cambodia, China (incl. Taiwan), Fiji, India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Japan (incl. Ryukyu Is.), Laos, Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, Philippines, Solomon Is., Sri Lanka, Thailand, Vietnam. A. macrorrhizos, A. plumbea and A. cucullata are cultivated and naturalized throughout the tropics. ECOLOGY: tropical and subtropical humid forest; geophytes or terrestrial, forest floor, in leaf litter, humus deposits on rocks, usually in deep shade, sometimes in exposed areas of forest regrowth. ETYMOLOGY: Greek a- (not) and Colocasia, the latter without its first syllable, implying a genus close to but different from Colocasia. TAXONOMIC ACCOUNTS: Krause & Engler in Engler & Krause (1920), Nicolson (1968a), Sivadasan (1982), Burnett (1985), Hay (1990a), Hay & Wise (1991).

C

Tribe Pistieae Tribe Pistieae Blume, Rumphia 1: 76 (1836, “Pistiaceae”). Laticifers absent; plant small, acaulescent; leaves several, rosulate, densely pubescent; petiole very short; blade obovate-cuneate to oblong, apically rounded to retuse, midrib absent, primary veins subparallel, higher order venation reticulate; inflorescence 1, very small; spathe ± constricted, lower margins connate forming tube, free margins between tube and blade folded to form a partition separating upper male partial chamber from lower female one, blade gaping; spadix:– female zone with single basal gynoecium and thin pouch-shaped flap below spathe partition, male zone with 2–8 flowers in a whorl, subtended by thin annular flap, spadix apex ± prominulent, naked, very short; flowers unisexual, perigone absent; male flower a sessile synandrium of 2 connate stamens, thecae dehiscing by single apical slit, pollen exine plicate with strongly undulate frills running length of grain; ovary obliquely adnate to spadix axis, 1locular, ovules numerous, orthotropous, placenta parietal (morphologically basal), style distinct, curved, stigma small; berry thin-walled, several-seeded, pericarp eventually decaying; seed subcylindric, testa thick, reticulate-alveolate, with micropylar operculum formed by both integuments, embryo minute, endosperm copious.

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105. Pistia Pistia L., Sp. Pl. 963 (1753). TYPE: P. stratiotes L. SYNONYMS: Kodda-Pail Adanson, Fam. 2: 75, 541 (1763); Zala Loureiro, Fl. Cochinch. 401, 405 (1790); Apiospermum Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1852: 351 (1853); Limnonesis Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1852: 352 (1853). Laticifers absent. HABIT: small, free-floating evergreen herb with pendent feathery roots, stem very short, acaulescent, stoloniferous. LEAVES: several in a rosette, densely pubescent. PETIOLE: very short, almost absent, sheath very short, ligulate, very thin, scarious at base. BLADE: somewhat spongy, obovate-cuneate to obovate-oblong, apically rounded, truncate or retuse; midrib absent, primary veins subparallel, all arising from base, diverging slightly and running into margin near apex, strongly prominent on lower surface, higher order venation reticulate. INFLORESCENCE: solitary, very small, much shorter than leaves. PEDUNCLE: very short, pubescent. SPATHE: pubescent without, glabrous within, somewhat constricted centrally, lower margins connate with each other and with ovary wall forming tube, free margins between tube and blade folded between stigma and male organs forming a partition between an upper male partial chamber and a lower female one, blade erect, ovate, expanded, acute-acuminate. SPADIX: shorter than spathe, mostly adnate to spathe, only the apical male zone free, female zone bearing single gynoecium at base and a thin, green, pouch-shaped flap just below spathe partition, male zone subtended by a thin, marginally lobed, green, annular flap, composed of a basally naked spadix axis supporting a single whorl of 2–8 flowers, naked axis sometimes extending a little beyond. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium composed of 2 connate stamens, thecae dehiscing by single apical slit. POLLEN: inaperturate, ellipsoid-elongate to -oblong, medium-sized (mean 27 µm.), exine plicate with strongly undulate frills running the length of the grain. FEMALE FLOWER: gynoecium obliquely adnate to spadix axis, ovary ovoid, 1-locular, ovules numerous, orthotropous, placenta broad, apparently parietal, probably morphologically basal, stylar region attenuate, bending inwards towards male flowers, stigma small, discoid-subcapitate. BERRY: thin-walled, utricular, several-seeded, ellipsoid, irregularly breaking up and decaying to release seeds. SEED: barrel-shaped, ± subtruncate and excavated at apex and base, testa thick, reticulate-alveolate, thicker and with operculum at micropylar end, embryo obovoid to conoid, endosperm copious. See Plates 105, 130C. CHROMOSOMES: 2n = 28. DISTRIBUTION: 1 sp.; pantropical:– Afghanistan, Angola, Argentina, Australia, Bangladesh, Belize, ?Benin, Bolivia, Botswana, Brazil, Brunei, Burkina Faso, Burundi, Cambodia, Cameroon, Central African Republic, Chad, China (incl. Taiwan), Colombia, Comores Is., Congo, Costa Rica, Cuba, Dominican Republic, Ecuador, Egypt, Equatorial Guinea (Rio Muni), Ethiopia, Gambia, Ghana, Guatemala, Guinea, Guyana, Haiti, Honduras, India, Indonesia (Borneo, Irian Jaya, Java, Moluccas), Ivory Coast, Jamaica, Kenya, Laos, Lesotho, Lesser Antilles, Liberia, Madagascar, Malawi, Malaysia (Borneo, Peninsula), Mali, Mauritius, Mexico, Mozambique, Nepal, Nicaragua, Niger, Nigeria, Pakistan,

C

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A

E

C

D

B

Plate 105. Pistia. A, habit in flower with stolons × 2/3; B, inflorescence, front view × 6; C, inflorescence, side view, longitudinal section × 6; D, fruit × 4; E, seed × 8. Pistia stratiotes: A, FTEA Araceae 67, Fig. 17 (1985); B–E, Giles & Woolliams s.n., Cult. Kew 1963–41001 (Kew spirit collection 29047.737).

PISTIEAE : PISTIA

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105. Pistia

Panama, Papua New Guinea, Paraguay, Peru, Philippines, Puerto Rico, Senegal, Sierra Leone, Somalia, South Africa, Sri Lanka, Sudan, Surinam, Swaziland, Tanzania, Thailand, Togo, Trinidad, Uganda, USA, Uruguay, Venezuela, Vietnam, Zaïre, Zambia, Zimbabwe. ECOLOGY: tropical wetlands; floating aquatic in open, tranquil, freshwater habitats. NOTE: We have followed Engler (1920a) in our interpreta-

288

THE GENERA OF ARACEAE

tion of the structure of the male flowers, which is supported by new developmental and anatomical studies (French 1986a, Buzgó 1994). ETYMOLOGY: Greek pister (hollow, trough, in the sense of a drinking trough). TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1975), Nicolson (1979), Sivadasan (1982), Mayo (1985a, 1986a), Hay (1990a).

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27 D E S C R I P T I O N O F A C O R A C E A E

Family Acoraceae Family Acoraceae C. Agardh, Aphor. Bot. 133 (1822, “Acoroideae ”). One genus, distributed throughout the northern hemisphere, temperate to tropical latitudes.

C

Acorus Acorus L., Sp. Pl. 324 (1753). TYPE: A. calamus L. Laticifers absent, raphides absent. HABIT: herbs, stem a repent, much-branched, lacunose rhizome with aromatic oil cells. LEAVES: distichous, unifacial, ensiform, not differentiated into petiole and blade, intravaginal squamules present in leaf axils; primary veins parallel, higher order venation parallel. INFLORESCENCE: solitary, terminal, borne laterally on leaf-like scape, continuation shoot arising in axil of leaf preceding spathe. SPATHE: much longer than spadix, erect, persistent, appearing merely as a vertical extension of the leaf-like peduncle. SPADIX: jutting out at an angle from peduncle, conoid and finger-like or slender and tail-like. FLOWERS: bisexual, perigoniate, densely arranged, bractless, 3-merous; tepals 6, in 2 whorls of 3, thin, fornicate. STAMENS: 6, in 2 whorls of 3, free, filaments linear-oblong, anthers introrse, thecae rounded-elliptic, subopposite, dehisc-

ing by longitudinal slit, connective inconspicuous. POLLEN: monosulcate to subulcerate, ellipsoid, small (mean 18 µm; range 15–20 µm), exine shallowly and remotely or more densely foveolate, otherwise psilate, apertural exine subpsilate. GYNOECIUM: obconic-cylindric, slightly exceeding tepals, 2–3-locular, ovules several per locule, pendent, orthotropous, both integuments bearing trichomes, inner integument longer than outer and forming micropyle, placenta apical, stigma minute, sessile. BERRY: oblong-obovoid with thin, leathery pericarp, enclosed by tepals, 1–5(–9)seeded, whitish with brownish stigma remnant when fresh, soon drying to straw-brown. SEED: oblong to ellipsoid, testa light brown, foveolate (A. calamus) or smooth (A. gramineus), long integumentary trichomes present (A. gramineus) or absent, embryo axile, cylindric or conoid (A. gramineus), perisperm present, endosperm copious. See Plates 106, 130D. CHROMOSOMES: 2n = 22, 24, 36, 44, 48. DISTRIBUTION: 2(–4) spp.; temperate to subtropical Asia and North America; introduced and naturalized in Europe. ECOLOGY: temperate to tropical wetlands, up to 1100m alt. in central Europe and up to 2600m alt. in China; helophyte, marshes, streams, ponds, swampy sites, pastures, meadows. ETYMOLOGY: akoron, an ancient Greek plant name. TAXONOMIC ACCOUNTS: Engler (1905), Huttleston (1953), Röst (1978, 1979a, b), Röst & Bos (1979), Riedl (1977–1979), Sivadasan (1982), Grayum (1987).

ACORACEAE : ACORUS

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C M

K

L

F

A

H

J

G

E D

B

Plate 106. Acorus. A, habit × 1/6; B, base of plant and rhizome × 2/3; C, detail of leaf venation × 3; D, inflorescence × 2/3; E, detail of culm × 3; F, detail of spadix × 5; G, flower with perigone × 8; H, flower, perigone removed × 8; J, gynoecium, longitudinal section × 8; K, berry × 8; L, berry, transverse section × 8; M, immature seed × 100. Acorus calamus: A, Birch Wolfe Herb. (K); B, Turrill s.n. (K); C–D, Mitchell 1143 (K); E–J, M, Barnes 1629 (Kew spirit collection 6913); K–L, Licent 1473 (K).

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Wakabayashi, S. (1957a). Studies on the structure and formation of mucilage cells in corms of Amorphophallus konjac. J. Jap. Bot. 32: 337–346. Wakabayashi, S. (1957b). On the form and formative stage of raphidian cells in some araceous and orchidaceous plants. J. Jap. Bot. 32: 268–274. Wall, M.E., Krider, M.M., Krewson, C.F., Eddy, C.R., Willaman, J.J., Correll, D.S. & Gentry, H.S. (1954a). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents I. [“VII.”]. J. Amer. Pharm. Assoc. 43: 1–7. Wall, M.E., Krider, M.M., Krewson, C.F., Eddy, C.R., Willaman, J.J., Correll, D.S. & Gentry, H.S. (1954b). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents II [“XIII.”]. Supplementary table of data. US Dept. Agric., Agric. Res. Service, Eastern Utilization Res. Branch, Philadelphia (February 1954). Wall, M.E., Eddy, C.R., Willaman, J.J., Correll, D.S., Schubert, B.G. & Gentry, H.S. (1954c). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents III. [“XII.”]. J. Amer. Pharm. Assoc. 43: 503–505. Wall, M.E., Eddy, C.R., Willaman, J.J., Correll, D.S., Schubert, B.G. & Gentry, H.S. (1954d). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents IV. [“XV.”]. Supplementary table of data. US Dept. Agric., Agric. Res. Service, Eastern Utilization Res. Branch, Philadelphia (June 1954). Wall, M.E., Fenske, C.S., Willaman, J.J., Correll, D.S., Schubert, B.G. & Gentry, H.S. (1955a). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents V. [“XXV.”]. J. Amer. Pharm. Assoc. 44: 438–440. Wall, M.E., Fenske, C.S., Willaman, J.J., Correll, D.S., Schubert, B.G. & Gentry, H.S. (1955b). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents VI. [“XXVI.”]. Supplementary table of data. US Dept. Agric., Agric. Res. Service, Eastern Utilization Res. Branch, Philadelphia (September 1955). Wall, M.E., Fenske, C.S., Kenney, H.E., Willaman, J.J., Correll, D.S., Schubert, B.G. & Gentry, H.S. (1957). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents VII. [“XLIII.”]. J. Amer. Pharm. Assoc. 46: 653–684. Wall, M.E., Fenske, C.S., Garvin, J.W., Willaman, J.J., Jones, Q., Schubert, B.G. & Gentry, H.S. (1959). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents VIII. [“LV.”]. J. Amer. Pharm. Assoc. 48: 695–722. Wall, M.E., Garvin, J.W., Willaman, J.J., Jones, Q. & Schubert, B.G. (1961). Steroidal sapogenins. Survey of plants for steroidal sapogenins and other constituents IX. [“LX.”]. J. Pharm. Sci. 50: 1001–1034. Wang, J.C. (1996). The systematic study of Taiwanese Arisaema (Araceae). Bot. Bull. Acad. Sin. 37: 61–87. Wang, J.-K. (1983). Taro: a review of Colocasia esculenta and its potentials. 400 pp. University of Hawaii Press, Honolulu. Warming, E. (1867). Nogle Iagttagelser over Varmeudviklingen hos en Aroidee, Philodendron lundii. Vidensk. Meddel. naturhistor. Forening Kjöbenhavn 1867 (8–11): 127–144, t. IV. Warming, E. (1883). Tropische Fragmente. I. Die Bestäubung von Philodendron bipinnatifidum Schott. Bot. Jahrb. 4: 328–340. Wattendorf, J. (1974). Ultrahistochemical reactions of the suberized cell walls in Acorus and Larix. Z. Pflanzenphysiol. 73: 214–225. Wealth of India, The. (1959). Raw Materials, Vol. V. (H–K), pp. 112–113. C.S.I.R. India, New Delhi. Wealth of India, The. (1985). Revised edition. Vol. I: A, pp. 230–235. New Delhi. Webber, E.E. (1960). Observations on the epidermal structure and stomatal apparatus of some members of the Araceae. Rhodora 62: 251–258.

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REFERENCES AND SELECTED TAXONOMIC LITERATURE

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29 G L O S S A RY

N.B. Terms describing pollen exine sculpture and ornamentation are not included (see Erdtman 1996). For terms in general botanical use we have been guided especially by the following reference works:– Bell (1991), Jackson (1928) and Stearn (1992).

abscission layer, zone – the region where one organ, such as a leaf or spathe, becomes detached from another, such as its node.

blade – the expanded, normally dorsiventrally flattened portion of the leaf borne at the apex of the petiole.

aculeate – prickly, armed with prickles.

boat-shaped spathe – a spathe which has the shape of the hull of a simple boat standing on end, thus the margins held apart and without any central contriction; the margins often overlap somewhat towards the base.

acuminate – the shape of e.g. a leaf or spathe apex which narrows gradually in such a way that the apex margins are concave. adnate – the fusion of two different structures, e.g. spathe and spadix; see connate. aerial – refers to a plant or stem which is situated above the ground or water. agravitropic – growing without responding to the direction of gravity, neither negatively nor positively; e.g. anchor roots.

bract – a modified leaf associated with a flower or inflorescence, usually of simple, undifferentiated structure. bulbil – small organs of vegetative propagation covered with minute scale-leaves, e.g. Remusatia; also used to refer to the tubercles found in the axils of the major leaf veins of Amorphophallus bulbifer and in Pinellia species.

amphitropous ovule – ovule with its axis strongly curved like a horse-shoe so that the two ends are situated near to each other. anastomosing laticifers – laticifers which branch and fuse with others forming a network.

caducous – cataphyll or spathe; falling when tissues are still fresh and alive.

anatropous ovule – ovule with a more-or-less straight axis, with the micropyle situated near the funicle.

campanulate – bell-shaped, in Araceae used in the sense of an inverted bell to describe spathe shapes.

anchor root – roots which anchor a hemiepiphyte or epiphyte to its substrate, generally a tree or rock.

campylotropous ovule – ovule with its axis slightly curved; see amphitropous.

annular insertion – the insertion of a leaf or spathe so that it encircles the node.

capitate stigma – head-shaped, i.e. approximately globose.

anterior division of leaf – that part of the leaf blade which lies above a horizontal line drawn through the petiole insertion (i.e. perpendicular to the midrib); that part of the leaf blade which surrounds the midrib; see posterior division.

cataphyll – a modified leaf which lacks a blade and in appearance corresponds to a petiole sheath; may be used to describe other leaf types whose technical names are defined by position rather than form, e.g. prophylls are usually of cataphyll shape in Araceae; see prophyll.

apical placentation – placenta situated at the apex of the ovary locule.

claviform – club-shaped.

aquatic – refers to a plant which lives in water; see helophyte. article – sympodial unit; a determinate unit of a sympodium derived from a single meristem. auriculate – with ears, especially of the apices of leaf sheaths and certain spathe blade types. axile placentation – placenta or placentae situated along the central angle formed by the septa of a multilocular ovary. axillary – lying in a leaf axil, i.e. the angle formed by a leaf and its adjacent internode.

collar rhizoid – epidermal trichomes occurring at the level of the root collar (Wurzelhals) on the hypocotyl. colocasioid venation – a type of higher order leaf venation found in tribes Colocasieae and Caladieae in which the finer veins branch almost at right angles from the primary lateral veins and then arch strongly towards the leaf margin, often fusing along the way to form a more-or-less sinuose interprimary collective vein between the primary lateral veins, and finally joining within the margin to form a submarginal collective vein. conchiform – shaped like the shell of a mollusc. connate – of two organs of the same type which are fused together, e.g. stamens; see adnate.

basal placentation – placenta situated at the base of the ovary locule.

connective – the tissue which connects the thecae of an anther

basifixed – of a stamen in which the base of the anther is attached to the apex of the filament.

constricted, constriction of spathe – of spathes which are narrowed somewhere along their length, usually near the middle.

bifacial leaf – a typical leaf with the blade flattened in a horizontal plane when viewed in cross section, and with upper (adaxial) and lower (abaxial) surfaces; = dorsiventral leaf; see ensiform, unifacial.

continuation shoot – the succeeding sympodial unit (= article) at any point in a sympodium.

bifid – divided into two, refers especially to the ultimate leaf lobes of certain genera, e.g. Anchomanes, Pseudohydrosme. biforine – a larger type of raphide idioblast (specialized cell containing raphides) with thickened side walls and thinner, nipple-like end walls. bipinnatifid leaf – a leaf blade divided pinnately, with each primary pinna or lobe itself divided pinnately; see -fid. bisexual flower – equivalent to hermaphrodite or monoclinous, i.e. fertile gynoecium and androecium present in each flower.

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contractile root – a type of root common in e.g. tribes Areae and Arisareae that contracts following initial extension and serves to prevent the tuber from growing above the soil surface, or even pulls the tuber further into the soil. convolute – see supervolute cordate, cordiform – heart-shaped. coriaceous – of leathery texture. costate – seed; ribbed or finely ribbed. cotyledonar sheath – sheath (bifacial base) of the cotyledon. cucullate – spathe, tepal; hooded or hood-shaped; see fornicate.

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C

E

F

A

B

G

D

Venation types: A, a type of reticulate venation and sagittate leaf typical of subfamily Lasioideae (Cyrtosperma cuspidispathum); B, reticulate on pinnate leaf segment (Amydrium zippelianum); C, reticulate on entire leaf (Arum maculatum); D, reticulate with collective vein (Pothos hosei); E, colocasioid (Colocasia esculenta); F, parallel-pinnate (Philodendron craspedodromum); G, reticulate venation in leaf segment of tribe Spathicarpeae (Gorgonidium vargasii).

GLOSSARY

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culm – a stalk, refers to the peduncle of Gymnostachys and Acorus (Acoraceae). cuneate – narrowed in a wedge-shape, usually used to describe the base of a leaf blade. cuspidate – of leaves and spathe blades, narrowed suddenly at the apex; an exaggerated form of acuminate.

female zone – that part of the spadix covered by female flowers; always situated in the basal portion of the spadix except in Spathicarpa; = pistillate zone. fenestrate leaf – equivalent to perforate; a leaf blade with holes between the major veins that have arisen by necrosis at an early stage in ontogeny. -fid – division of a leaf in which the sinus between each lobe extends over halfway, but not completely, to the midrib.

deciduous – cataphyll or spathe; falling after complete or partial death of tissues. dehiscent – anther thecae, fruits; splitting open. deliquescent – cataphyll or spathe; dying and immediately decomposing while still attached.

flagelliform branch/shoot – fast-growing branches in which the internodes are more elongated and the leaves somewhat to highly reduced; a typical adaptation of hemiepiphytes for searching out and colonizing new host trees.

depressed-globose – tuber; a sphere flattened somewhat by pressing in the two opposite poles.

flexuose – zigzag, as in the spadix axis of Pedicellarum.

diclinous flower – equivalent to unisexual; stamens and gynoecia borne in separate flowers.

floral sympodium – (in Araceae) that part of the sympodial flowering shoot which is composed of two or more successive inflorescences and their associated cataphyllary prophylls, e.g. the pseudoaxillary spadix clusters in Homalomena, many Philodendron species, etc.

dimorphic roots – roots specialized into two different functional types, anchor roots and feeder roots; typical of hemiepiphytic aroids. diporate pollen – pollen grain type having two apertures. discoid stigma – a thickly disc-shaped stigma resembling in miniature a round, flattened loaf of bread. distichous leaves – a form of phyllotaxis when alternate leaves are arranged in two rows when seen from above, usually 180° apart. dorsiventral leaf – equivalent to bifacial leaf. dracontioid leaf – elaborated forms of sagittate, hastate or trisect leaves in which the anterior and posterior divisions are highly dissected and subdivided; see Amorphophallus, Dracontium, Pseudohydrosme, Pycnospatha. druse – a type of microscopic crystalline structure found in specialized cells (idioblasts) with the form of a medieval mace, i.e. ± spherical with crystal apices pointing in all directions.

elliptic – of e.g. a leaf having the shape of an ellipse. emarginate – of e.g. a leaf having a rounded apex with a shallow central notch. endemic – a geographic term meaning restricted entirely to the area referred to, e.g. endemic to Madagascar. endothecial (cell wall) thickening – conspicuous thickenings in the cell walls of the endothecium; various patterns are characteristic of particular taxa. endothecium – a specialized cell layer surrounding the tapetal layer of the microsporangia (pollen sacs) in an anther. ensiform leaf – a type of unifacial leaf which is flattened in a vertical plane when viewed in cross section, as in Acorus, Iris. entire – of leaves, without lobing or division and with an even margin. epigeal – of an organ or process located above ground. epigeal germination – a type of germination in which the cotyledon is carried above the ground and normally becomes green and photosynthetic, e.g. Philodendron. epiphyte – a non-parasitic plant which grows on another plant, its host, and which is not connected to the ground during its life cycle. Eurasia – continental Europe and continental Asia together.

312

filiform – thread-shaped.

floral bract – bract subtending a single flower.

foliage leaf – a leaf of mature form, in most genera thus differentiated into petiole and blade. forate – a type of pollen grain with several apertures having a more-or-less equidistant distribution over the grain surface. fornicate – of spathes; arched over, hooded; = cucullate. funicle – usually slender organ connecting the chalaza of the ovule (or seed) to the placenta; in the anatropous ovule the funicle is partly adnate laterally to the outer integument fusiform seed – thick but tapering towards each end, spindleshaped.

geniculum, geniculate, (pulvinus, pulvinate) – swelling or joint located usually at the apex of the petiole that permits independent movement of the leaf blade. geophyte – plants that have subterranean stems, implies a tuberous or rhizomatous habit. gynoecium – equivalent to pistil; the female organ consisting of ovary, stylar region or style and stigma.

hastate – halberd-shaped, i.e. a sagittate leaf with the posterior divisions turned outwards. helophyte – marsh or swamp plants, i.e. growing in ground flooded for at least part of the year and with the foliage above the water level. hemianatropous ovule – similar to anatropous but with the axis of the ovule more-or-less horizontal. hemiepiphyte – plants which grow on hosts and are detached from the ground at some stage of their life cycle, later becoming reconnected with the ground by sending down feeder roots. hemiorthotropous ovule – an ovule in which the micropyle points away from the funicle but in which the funicle is attached sublaterally near the chalazal end and not terminally; see orthotropous. hermaphrodite flower – equivalent to bisexual; a flower with both androecium and gynoecium.

extrorse anther – anther in which the thecae dehisce facing away from the centre of the flower.

heteroblasty – the production of leaves of differing shape and size during the development from juvenile to mature form; typical of hemiepiphytes.

feeder roots – specialized roots of hemiepiphytes which extend down to the soil and provide nutrients to the plant.

higher order venation – fine venation, normally refers to tertiary and quaternary, etc. degrees of venation in the leaf.

female flower – a flower composed only of a gynoecium, sometimes associated with one or several staminodes, a perigone maybe present or absent; = pistillate flower.

hilum – scar left on seed surface after abscission of funicle.

THE GENERA OF ARACEAE

hyperphyll (of cotyledon) – portion of cotyledon distal to sheath (“Oberblatt”), always unifacial in araceous seedlings.

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B

A

F

E C

D

G

H L

K

J

P M

Q

N

Leaf shapes: A, linear; B, elliptic; C, ovate; D, ovate and peltate; E, cordate, cordiform; F, sagittate; G, hastate; H, trifid; J, trisect; K, pedatifid; L, pedatisect; M, pinnately lobed; N, pinnatifid; P, pinnatisect; Q, dracontioid.

GLOSSARY

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hypocotyl – that part of the stem axis which lies between the cotyledonar node and the primary root. hypogeal – of an organ or process located below ground.

microsporangium, microsporangia – the individual pollen sacs making up the thecae and anthers; in Araceae each theca is normally composed of 2 microsporangia, and each anther of 2 thecae.

imbricate – overlapping.

midrib – the large, central, axial vein of the anterior division of the leaf.

inaperturate – a type of pollen grain which lacks any obvious aperture.

monad – of pollen grains shed as single grains; see tetrads.

infructescence – the inflorescence at fruiting stage, i.e. the mass of fruits considered as one composite structure.

monoclinous – of flowers; equivalent to bisexual, hermaphrodite; stamens and gynoecia borne in the same flower; see diclinous.

internode – the portion of stem between each pair of nodes.

monopodium, monopodial – a shoot axis which is formed by the vegetative extension of a single apical meristem.

interprimary vein – a vein, thicker than the fine veins but thinner than the primary lateral veins, lying approximately parallel to and between them.

monosulcate – of a pollen grain with a single narrowly elliptical aperture.

introrse anther – anther in which the thecae dehisce towards the centre of the flower.

naked axis, axial zone, region, portion – a portion of the spadix which lacks any kind of floral organ or specialized tissues, and consists of a simple, smooth axis.

intrusive (intrusive-parietal) placenta – a placenta borne on a septum that extends from the ovary wall almost to the centre of the locule.

neotropical – the tropical regions of the New World, i.e. tropical Mexico, Central America, the West Indies and tropical South America.

involute – a form of folding of a single leaf in which the two leaf margins are each inrolled without either clasping the other, as in Lagenandra.

node – the point on the stem where a leaf is inserted.

obconic – in the shape of an inverted cone. juvenile shoot – shoots which have yet to acquire their mature size or bear leaves of mature form.

kettle – a basal tubular portion of the spathe in which the margins are connate, forming a chamber; characteristic of Lagenandra and Cryptocoryne.

laciniate leaf blade – a form of leaf perforation in which the major perforations between the primary lateral veins are very narrow and elongated, resembling slashes, and often reach the margin, e.g. Cercestis mirabilis. laticifers – longitudinal rows of slender, tubular cells usually associated with vascular bundles and containing clear or milky fluid called “latex”. latrorse anther – anther with the thecae dehiscing laterally, as seen in cross section; see extrorse, introrse. lenticular seed – shaped like a double convex lens. ligule, ligulate – a free extension of the apex of the petiole sheath; very elongated in most genera of the Schismatoglottideae. linear – narrow and elongated with more-or-less parallel margins. lithophytes – plants that grow on rocks. lobe – as used here referring in a non-specific way to any subdivision of the leaf blade within the anterior or posterior divisions.

Malay Archipelago – as used in this book, refers to the geographical region which includes Malaysia, Indonesia, the Philippines and Papuasia; also known as Malesia. male flower – a flower composed only of an androecium, sometimes associated with a pistillode, a perigone maybe present or absent; = staminate flower. male zone – that part of the spadix covered by male flowers; = staminate zone. marcescent – cataphyll or spathe; remaining attached to the plant after death and partial decomposition of tissues. membranaceous – of thin texture.

obpyramidal – in the shape of an inverted pyramid. operculate – furnished with a lid, e.g. seeds of Pistia. orbicular – of a flat body circular in shape. orthotropous ovule – ovule in which the body of the ovule is straight, the micropyle faces directly away from the funicle and the funicle is attached terminally (not laterally) to the chalazal end of the ovule. osmophoric – having the function of an osmophore, i.e. an organ or area of tissue specialized for the production of odours. ovate – of a flat body having the shape of an egg, with the broader blunter half at the base and narrower more acute half at the top.

palaeotropical – the tropical regions of the Old World; i.e. Africa, Asia, Malay Archipelago, Melanesia, Oceania, tropical Australasia. paradioecy – the production, from the same stem, of entirely male or entirely female inflorescences in different seasons; in Araceae observed only in Arisaema. parallel venation – a leaf venation pattern in which the primary, secondary and often tertiary veins run longitudinally and parallel to one another, from the base to the apex of the leaf. parallel-pinnate venation – a leaf venation pattern in which the primary and secondary lateral veins arise pinnately from the midrib and then run parallel to one another towards the leaf margin; see e.g. Philodendron. parietal – borne on a wall, of ovules in which the placenta and hence the funicle attachment lie on the side walls of the ovary. pedati- (pedatifid, pedatisect) – literally, foot-shaped; of a leaf divided in such a way that the midribs of the lateral segments, lobes or pinnae are inserted successively on two basal ribs rather than all together at the petiole insertion; cf. radiati-; see e.g. Sauromatum. pedicellate – of a flower borne on a pedicel or stalk. peduncle – the internode between the spathe and the preceding leaf.

-mery, -merous – as in 2-merous, 3-merous; refers to the number of parts in the flower.

pellucid – wholly or partially transparent.

mesocarp – the middle layer of the fruit wall.

perianth – the floral envelope consisting of calyx and corolla; in Araceae referred to as a perigone.

micropyle – apical aperture of the ovule formed by the inner and outer integuments.

314

obovate – of a flat body in the shape of an egg but with the broader, blunter half at the top and the narrower, more acute half at the base.

THE GENERA OF ARACEAE

perforated – of a leaf blade punctured by holes; see e.g. Monstera.

pericarp – a fruit wall derived directly from ovary wall tissue.

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perigone – the floral envelope of a flower in which there is no differentiation of calyx from corolla, it may be a single structure (connate tepals) or composed of individual, similar tepals.

pseudolateral – an inflorescence or shoot which appears to be axillary to a leaf, i.e. derived from an axillary bud, but which is really a terminal axis displaced to one side.

perigoniate – of a flower which possesses a perigone.

pseudomonomerous ovary – a unilocular ovary which is presumed to be phylogenetically derived from an ancestor with a multilocular (multicarpellary) ovary.

persistent – of a leaf or spathe which remains attached, with its tissues alive and functioning. petiole – the stalk of a leaf.

pseudostem – an erect, stem-like structure formed by tightly imbricate leaf sheaths, as in bananas; see Typhonodorum, Arisaema.

petiole sheath – the basal, sheathing part of the petiole which normally has an annular insertion at the node; the sheath may be conspicuous or not, and persistent or not.

psilate – smooth (pollen surface).

phyllotaxis, phyllotaxy – the pattern of arrangement of leaves on the stem.

ptyxis – the manner in which a single leaf is folded while still in bud; see involute, supervolute.

pinna – a single lateral subdivision of the anterior or posterior leaf divisions; may also be referred to as a segment or even lobe.

pubescent – hairy.

pinnate – literally, in the form of a feather; a structural pattern, e.g. primary leaf venation, in which there in a central longitudinal axis bearing lateral subaxes. pinnati- (pinnatifid, pinnatisect) – of a leaf divided pinnately; see -fid and -sect. pistil – equivalent to gynoecium. pistillate – equivalent to female, refers to female flowers or the zone of the spadix bearing female flowers. pistillode – floral structures which are considered to be homologous (in position or form or both) to gynoecia (pistils) but which lack ovules. placenta (plural: placentae) – specialized area of tissue within the ovary to which the ovules are attached by their funicles; in Araceae the placentae are almost always covered with a dense epithelium of glandular hairs which secrete a clear, mucilaginous substance. placentation – the position of the placenta or placentae within the ovary; e.g. apical, axile, basal, parietal. plesiomorphy, plesiomorphic – a character, occurring in a monophyletic group, that evolved, not in the immediate common ancestor, but in a more distant ancestor which also gave rise to other extant monophyletic groups; this term is strictly defined conceptually, but it may be regarded as loosely equivalent to “generalized” or “primitive” character. pollenkitt – material derived from the microsporangial inner cell layers which coats the surface of the pollen grain; in Araceae it often acts to glue the pollen grains together into strands on extrusion from the thecae, e.g. Philodendron, Zantedeschia. posterior divisions – the two portions of the leaf blade, one on each side of the leaf axis, which lie below a horizontal line drawn through the petiole insertion perpendicular to the midrib. primary lateral veins – the major veins which compose the midrib and basal ribs of the leaf blade and which branch laterally from them. primary root – radicle, i.e. the root which develops exogenously from the root pole of the embryo; in Araceae always short-lived or absent and replaced by adventitious roots. primary veins – the veins which extend into the leaf blade from the stem via the petiole. primary venation – the overall pattern of the major veins of the leaf.

pulvinate, pulvinus – bearing a pulvinus, i.e. a swollen portion of e.g. a petiole which acts as an articulation, permitting movements of the petiole and blade relative to one another; see geniculum, geniculate. punctiform – in the form of a point or dot.

quadripinnatifid – a leaf blade which is divided in such a way that there are four hierarchical levels of pinnate leaf lobe division.

radiati- (radiatisect) – a leaf blade in which the midribs of the segments, pinnae, lobes or subdivisions all converge on and unite at the petiole insertion. raphe – the part of the funicle when it has become adnate laterally to the body of the ovule, forming a ridge or mark on the surface; may be conspicuous in the seed. raphides – microscopic needle-like crystals of calcium oxalate. repent – creeping on the ground and rooting. resin canal – tubular structures within vegetative and floral tissues of certain genera which contain as yet inexactly determined substances, probably of terpenoid type; resin canals are often conspicuous as translucent lines or dots in leaf blades, e.g. Cercestis, Culcasia, or, in Philodendron, as yellow to brown lines on the inner surface of spathes. reticulate leaf venation – fine leaf venation of a net-like pattern. rheophytes – plants which grow in fast-flowing streams between the high and low seasonal water levels and usually submerged during the flood season; usually attached to rocks and adapted to this habitat. rhizome – a subterranean stem type of cylindrical form, may be horizontal or vertical in orientation. root collar – boundary region between the hypocotyl and the primary root (Wurzelhals); usually bearing collar rhizoids. rosulate – rosette-like. rugose – of a surface which is rough but without sharp projections or scales.

sagittate – arrow-shaped; of a leaf blade shape with somewhat acutely tipped, backwardly-directed posterior divisions; cf. hastate.

prophyll – the first leaf of a branch (or sympodial unit); in Araceae almost always a 2-keeled cataphyll, often confused with cataphyll:– cataphyll refers to a particular type of morphology (reduced leaf), prophyll refers to the position of the leaf along a branch.

scabrid – of a rough surface.

prostrate – lying flat.

-sect – division of a leaf blade in which the sinus between each lobe extends completely to the midrib.

protogynous – a plant or inflorescence or flower in which the stigmas become receptive before the anthers release their pollen. pseudoaxile placentation – a form of parietal placentation in which the placentae are borne on very deeply intrusive partial septa which may be partially fused (e.g. basally).

scape – a more-or-less leafless vertical axis bearing an inflorescence, e.g. grass, Gymnostachys. secondary and tertiary veins – leaf veins of successively higher order (finer) than the primary lateral veins.

segment, leaf – equivalent to lobe; as used here referring in a non-specific way to any subdivision of the leaf blade within the anterior or posterior divisions. septate – divided by one or more partitions or septa.

GLOSSARY

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sessile – lacking a stalk, pedicel, peduncle, style or stipe. shingle (leaves, plant) – a type of juvenile morphology, found in some hemiepiphytic species, in which the petiole is very short and the leaf blade relatively broad and more-or-less overlapping with its neighbours to resemble the tiles (or shingles) of a roof; such plants are found climbing up larger tree trunks; e.g. Monstera dubia, Philodendron scandens. simple leaf blade – a leaf blade which is neither lobed nor subdivided. spadix – a spike (racemose inflorescence with sessile flowers) with a thickened axis; in Araceae the axis is almost always rather fleshy and the flowers are not subtended by floral bracts; see spathe. spathe – a simple, bract-like foliar organ associated with the spadix; morphologically the last “leaf” of the flowering shoot (sympodial unit) in Araceae; the spathe is usually coloured and may have a complex shape; the combined unit of spathe and spadix is widely regarded as the “inflorescence” in Araceae, since there is such an intimate morphological and functional relationship between them; however, in strictly morphological terms it is the spadix which is the inflorescence, while the spathe is a modified leaf; see spadix, sympodial leaf. spathe blade – the upper, expanded (sometimes only temporarily) part of the spathe. spathe constriction – a constricted portion of the spathe occurring in many genera of Araceae; generally occurring at the point corresponding to the junction between the male and female zones of the spadix (e.g. Caladium, Xanthosoma), sometimes lying above the fertile zones (e.g. Arum). spathe tube – the lower, tubular part of the spathe formed by the supervolute (= convolute) or sometimes connate margins. spathulate – oblong, with the basal (proximal) end narrowed so that the whole resembles a chemist’s spatula. spike – a racemose (monopodial) inflorescence with sessile flowers. staminate – equivalent to male, refers to male flowers or the zone of the spadix bearing male flowers. staminode, staminodes, staminodia – floral structures which are considered to be homologous (in position or form or both) to stamens but which lack microsporangia; the term is often applied in the Araceae to structures which are quite different in form to fertile stamens; see e.g. Arum.

stylar region – tissue lying between ovary locules and stigmatic epidermis, distinguished from “style” because it is often as thick as the ovary in Araceae. style – a stylar region which is narrower than the ovary and somewhat elongated. subdracontioid – a leaf blade shape which corresponds to dracontioid form but in which the posterior divisions are weakly elaborated; see e.g. Taccarum. submarginal collective vein – leaf; a vein running parallel to and near the leaf margin into which the primary lateral veins run, diagnostic of brochidodromous leaf venation; see Anthurium. subpalmatifid – describes a leaf shape in which the segments are only partly divided from one another and their midribs converge on the petiole insertion (see radiati-). subulate – awl-shaped (i.e. narrowly cylindric with a sharp point). supervolute – a type of ptyxis in which one side of the leaf blade is wrapped around the other; also called convolute. sympodial leaf – the leaf or cataphyll which subtends the spathe and spadix, i.e. the last leaf of a sympodial unit. sympodium, sympodial – a shoot axis built up by a linear series of units (sympodial units), each new distal unit developing from a single apical meristem arising from an axillary bud situated on the previous unit; the axis is thus constructed successively by the activity of several different apical meristems; see monopodium. synandrium, synandria – a male flower composed of connate stamens, or the congenital fusion of several separate stamen primordia. synandrodium, synandrodes, synandrodia – a sterile synandrium, i.e. shaped like a synandrium but lacking microsporangia. synapomorphy, synapomorphic – a character, occurring in a monophyletic group of taxa, that evolved in their immediate common ancestor; this term is strictly defined conceptually, but it may be regarded as loosely equivalent to “specialized”, “derived” or “advanced”. syncarp – an infructescence in which the component fruits are congenitally or postgenitally fused together. synflorescence – (in Araceae) complex inflorescence composed of several spadices; equivalent to floral sympodium.

stellate – in the shape of a star. sterile appendix – a terminal portion of a spadix which is covered with sterile flowers, staminodia or may be merely rough to smooth; probably always osmophoric; e.g. Arum, Amorphophallus. sterile flower – this term is often applied in Araceae literature to infertile floral structures occurring on the spadix that bear little or no resemblance to fertile flowers, but which are probably derived ontogenetically from primordia which are homologous to floral primordia. sterile zone – portion of the spadix which bears sterile flowers or lacks fertile floral organs or tissues. stigmatoid – a sterile portion of a male flower which is thought to be derived from a primordium homologous with that of the stigma of a fertile gynoecium; see e.g. Spathantheum, Taccarum. stipe – a stalk; in Araceae used especially to refer to the axial region commonly present between the spathe insertion and the base of the floral zone of the spadix proper; see e.g. Aglaonema. stipitate – borne on a stipe or stalk. stolon – a stem branch specialized for vegetative reproduction by long internodes and, usually, reduced leaves. stomial pore, stomium – the opening in the microsporangial wall through which pollen is emitted. striate – marked with fine, longitudinal parallel lines. strophiole – an aril-like (fleshy) outgrowth of the raphe of a seed, usually appearing as an appendage to the hilum ; thought to be a structure that plays a role in seed dispersal (?food attractant to vectors).

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tannin cells – specialized cells found scattered throughout the plant tissues (in the Araceae) with contents that stain brown in alcoholic preservatives and when the tissues are dried; these contents are thought to be tanniniferous but this has rarely been rigorously established chemically. tepals – the individual component parts of the floral envelope or perigone; distinguished from sepals and petals in that the tepals of a single flower are all similar in shape and colour. terminal appendix – see sterile appendix. terrestrial – of a plant which grows on the ground (in contrast to e.g. an epiphyte) or on dry land (as distinct from an aquatic or helophyte). testa – the seed coat. tetrad – of pollen shed when the four pollen grains which are the product of a single meiosis remain united; see monads. theca, thecae – an adjoining pair of microsporangia; see microsporangium. trapezoid – having the shape of a trapezium, i.e. a plane figure with two parallel sides and the other opposite pair of sides not parallel to one another. trichosclereid – literally a microscopic hair-like sclereid; fibre cells (cells with thick, lignified walls) which are very slender and elongated so as to be visible to the naked eye as hair-like structures (e.g. on tearing the leaf blade they can be seen emerging from the torn edge, as in e.g. Spathiphyllum); either T- or Hshaped.

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tri- (trifid, trisect) – of a leaf blade which is subdivided, partially or completely, into three parts:– a central anterior division and two posterior divisions. tripinnatifid – of a leaf blade which is divided in such a way that there are three hierarchical levels of pinnate leaf lobe division. tropical southeast Asia – strictly speaking, the southeastern part of the Asian continent which lies within the tropics. Sometimes loosely used to include the islands of the Malay Archipelago. truncate – of the apex of a structure which appears as if cut off at the end in a single snip. tuber – in Araceae used to describe the swollen, subterranean stems characteristic of many genera, e.g. Arisaema, Arum, Amorphophallus, Dracontium. tuberculate – beset with knobbly projections or excrescences.

umbonate – bearing a boss in the centre, or shaped like a nipple. unifacial leaf – a leaf type in which the normal lateral extension during ontogeny is suppressed, resulting in a more or less cylindrical leaf; see bifacial, ensiform. unisexual – equivalent to diclinous; stamens and gynoecia borne in separate flowers. urceolate – cup-shaped. velamen – a covering of dead cells which protects the roots of some epiphytic plants, usually white when dry; e.g. Anthurium gracile. ventricose – literally, having the shape of a belly; in Araceae used for stoutly swollen spathe tubes; e.g. Stylochaeton. verrucose – warty, covered with wart-like projections.

tubercule – small tubers that develop from axillary buds or accessory buds on leaves, e.g. Amorphophallus bulbifer, Pinellia.

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GLOSSARY

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30 A P P E N D I X

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Table 9. Fungal parasites (including some bacteria) of Araceae

(data mainly from Brandenburger 1985 and Farr et al. 1989). Acorus calamus (Acoraceae)

Asteromella acorella, Cylindrosporium acori (leaf spot), Darluca filum, Leptosphaeria acorella, Physoderma calami, Ramularia aromatica (leaf spot), Septocylindrium sp., Sphaerulina acori (on dying leaves), Stagonospora calami (leaf spot), Uromyces sparganii (syn. U. pyriformis) (rust).

Aglaonema

Ascochyta minima, Botrytis cinerea, Colletotrichum sp. (leaf spot), Fusarium oxysporum, Glocosporium graffii (leaf spot), Mycosphaerella anthurii, Myrothecium roridum (leaf spot), Phyllosticta cavarae, Ph. microspora (leaf spot), Phytophthora sp., Pythium splendens (root rot), Rhizoctonia solani (aerial blight), Sclerotium rolfsii (southern blight).

Anthurium scherzerianum, A. andraeanum

Colletotrichum anthurii, C. gloeosporoides (leaf spot), Gloeosporium anthurii (leaf spot), G. minimum (leaf or burn spot), Mycosphaerella anthurii (leaf spot), Pestalotia briosiana (leaf spot), Phoma anthurii, Phyllosticta cavarae (leaf spot), Phytophthora parasitica (shoot base rot), Pythium splendens (root rot), Rhizoctonia solani (root rot), Septoria anthurii (leaf spot), Uredo anthurii (rust).

Arisaema dracontium

Uromyces ari-triphylli (rust).

Arisaema triphyllum

Cladosporium sp. (sooty mould), Ramularia arisaemae (leaf spot), Septotinia arisaemae, Uromyces ari-triphylli (rust) (Uromyces caladii of most authors), Volutella sp. (leaf spot).

Arisarum

Melanotaenium ari, Phyllosticta arisari (leaf spot).

Arum

Ascochyta arigena, A. arophila, A. pellucida, Melanotaenium ari, Phyllosticta aricola, P. tuszonii, Puccinia sessilis var. sessilis (rust), Ramularia ari, Septoria ari (leaf spot).

Biarum

Melanotaenium ari

Caladium bicolor (syn. C. x hortulanum)

Fusarium solanii (root rot), Pythium myriotylum (root rot), Rhizoctonia solani (blight), Sclerotium rolfsii (southern blight).

Calla palustris

Cercospora callae, Marssonia callae, Septoria callae (leaf spot).

Colocasia esculenta

Botryodiplodia theobromae (spongy black rot), Ceratocystis fimbriata (black rot), Cladosporium colocasiae (leaf spot), Erwinia carotovora, E. chrysanthemi (bacterial soft rot), Fusarium solani (fusarium dry rot), Leptosphaeria colocasiae (leaf spot), Phyllosticta colocasiophila (leaf spot), Phytophthora colocasiae (leaf blight), Pythium aphanidermatum, P. carolinianum, P. graminicolum, P. irregulare, P. myriotylum, P. splendens, P. ultimum, Pythium sp. (soft rot in tubers), Rhizoctonia bataticola (on leaves), Rhizopus stolonifer (rhizopus rot), Sclerotium rolfsii (southern blight).

Dieffenbachia seguine (and cultivars) (syn. D. maculata, D. picta)

Acremonium crotocinigenum (leaf spot and stem rot), Colletotrichum sp. (leaf spot), Cephalosporium cinnamomeum (leaf spot), Erwinia chrysanthemi (withering), Fusarium oxysporum, F. solani (stem rot), F. sp. (root rot), Glomerella cingulata (leaf spot), Leptosphaeria sp. (brown leaf spot), Myrothecium roridum (leaf spot), Phaeosphaeria eustoma (leaf spot), Phytophthora palmivora (stem rot), P. parasitica (leaf spot), Plectosphaerella sp. (leaf spot), Pythium sylvaticum (stem rot), Pythium sp. (root rot), Rhizoctonia solani (blight), Sclerotium rolfsii (southern blight)

Epipremnum pinnatum ‘Aureum’

Erwinia ssp., Pseudomonas cichorii (bacterial leaf spots), Pythium splendens (root rot), Rhizoctonia solani (leaf spot), Sclerotium rolfsii (southern blight).

THE GENERA OF ARACEAE

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Monstera

Colletotrichum sp. (leaf spot), Phyllosticta fragosoana (leaf spot), Rhizoctonia solani (aerial blight), Sclerotium rolfsii (southern blight).

Orontium aquaticum

Botrytis streptothrix (leaf blight), Epicoccum duriaeanum, Mycosphaerella sp. (leaf spot), Phyllosticta orontii (leaf spot), Physalospora orontii, Ramularia orontii (leaf spot), Stilbum aciculosum, Volutella diaphana (leaf spot).

Peltandra sagittifolia

Cercospora callae (leaf spot), Colletotrichum sp. (leaf spot), Uromyces ari-triphylli (rust).

Peltandra virginica

Cercospora callae (leaf spot), Gloeosporium paludosum (leaf spot), Pestalota aquatica (leaf spot), Ramularia sp. (leaf spot), Sclerotium caladii, Uromyces aritriphylli (rust).

Philodendron

Botrytis cinerea (blight), Cercospora sp. (leaf spot), Colletotrichum sp. (leaf spot), Dactylaria humicola (leaf spot), Myrothecium roridum (leaf spot), Phyllosticta philodendrina (leaf spot), Phytophthora parasitica (leaf spot), Pythium sp. (root rot), Rhizoctonia solani (aerial blight), Sclerotium rolfsii (southern blight).

Pistia

Phyllosticta stratiotis (leaf spot).

Spathiphyllum

Cylindrocladium spathiphylli (root and petiole rot), Myrothecium roridum (leaf spot), Phytophthora parasitica (root and shoot base rot), Pythium sp. (root rot), Rhizoctonia solani (aerial blight), Sclerotium rolfsii (southern blight).

Symplocarpus foetidus

Botrytis sp. (leaf blight), Cercospora symplocarpi (leaf spot), Gloeosporium foetidophilum, Nectria semenicola, Septoria spiculosa (leaf spot).

Syngonium podophyllum

Acremonium crotocinigenum (leaf spot), Ceratocystis fimbriata (black cane rot), Colletotrichum sp. (leaf spot), Myrothecium roridum (leaf spot), Pythium sp. (root rot), Rhizoctonia solani (aerial blight), Sclerotium rolfsii (southern blight).

Zantedeschia aethiopica

Cercospora callae, Cercospora richardiaecola (leaf spot), Colletotrichum montemartinii (leaf spot), Gloeosporium callae (leaf spot), Lycopersicon virus 3 (deformed leaves), Pecobacterium carotovorum var. aroideae (soft rot), Phyllosticta richardiae (leaf spot), Phytophthora erythroseptica, P. richardiae (tuber and leaf rot).

Zantedeschia hybrids

Ascochyta sp. (leaf spot), Cercospora richardiaecola, Gloeosporium callae (leaf spot), Pestalota richardiae (leaf spot), Phyllosticta richardiae (leaf spot), Phytophthora richardiae, Septoria sp. (leaf spot).

APPENDIX

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Table 10. Schott’s 1860 classification of Aroideae

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Names and taxon numbers are as in the original publication.

Aroideae I. Diclines A. Efilamentatae a. Stenozeugmaticae α. Orthotropooae Tribus Alleluchieae Subtr. Cryptocoryninae 1. Cryptocoryne 2. Lagenandra Subtr. Pinellinae 3. Pinellia Tribus Arisareae 4. Arisarum 5. Arisaema Tribus Dracunculeae Subtr. Biarinae 6. Biarum 7. Leptopetion 8. Cyllenium 9. Ischarum 10. Sauromatum Subtr. Arinae 11. Gymnomesium 12. Arum Subtr. Helicophyllinae 13. Theriophonum 14. Tapinocarpus 15. Calyptrocoryne 16. Typhonium 17. Heterostalis 18. Eminium 19. Helicophyllum 20. Helicodiceros Subtr. Dracunculinae 21. Dracunculus

ß. Anatropooae Tribus Zomicarpeae 22. Zomicarpa Tribus Pythonieae Subtr. Amorphophallinae 23. Allopythion 24. Pythonium 25. Plesmonium 26. Rhaphiophallus 27. Synantherias 28. Brachyspatha 29. Conophallus 30. Amorphophallus Subtr. Hydrosminae 31. Corynophallus 32. Hydrosme 33. Hansalia 34. Anchomanes

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b. Pachyzeugmaticae α. Gymnogoneae Tribus Caladieae Subtr. Colocasinae 35. Ariopsis 36. Remusatia 37. Colocasia 38. Leucocasia Subtr. Alocasinae 39. Gonatanthus 40. Alocasia 41. Peltandra Subtr. Anubiadinae 42. Anubias Subtr. Syngoninae 43. Typhonodorum 44. Hapaline 45. Caladium 46. Xanthosoma 47. Acontias 48. Syngonium Subtr. Problematicae 49. Zamioculcas Tribus Philodendreae Subtr. Aninginae 50. Montrichardia Subtr. Culcasinae 51. Culcasia 52. Nephthytis 53. Cercestis Subtr. Philodendrinae 54. Philodendron Subtr. Anaporinae 55. Aglaonema 56. Aglaodorum Subtr. Homalomeninae 57. Zantedeschia 58. Homalomena 59. Chamaecladon Subtr. Adeloneminae 60. Adelonema 61. Philonotium Subtr. Schismatoglottidinae 62. Apatemone 63. Bucephalandra 64. Apoballis

ß. Peristatogoneae Tribus Richardieae 65. Richardia Tribus Asterostigmeae Subtr. Dieffenbachininae 66. Dieffenbachia Subtr. Asterostigmatinae 67. Mangonia

68. 69. 70. 71. Subtr. 72. 73.

Taccarum Asterostigma Rhopalostigmium Andromycia Spathicarpinae Spathicarpa Spathantheum

B. Filamentatae Tribus Stylochitoneae 74. Stylochiton

II. Monoclines Tribus Calleae Subtr. Callinae 75. Calla Subtr. Monsterinae 76. Stenospermation 77. Atimeta 78. Rhodospatha 79. Anepsias 80. Tornelia 81. Alloschemone 82. Monstera 83. Heteropsis 84. Rhaphidophora 85. Epipremnum 86. Anadendron 87. Scindapsus 88. Cuscuaria Tribus Orontieae Subtr. Lasinae 89. Lasia 90. Cyrtosperma 91. Anaphyllum 92. Lasimorpha 93. Urospatha 94. Arisacontis Subtr. Dracontioninae 95. Dracontium 96. Echidnium 97. Ophione 98. Symplocarpus Subtr. Orontioninae 99. Lysichiton 100. Orontium Subtr. Spathiphyllinae 101. Spathiphyllum Subtr. Anthurinae 102. Anthurium Subtr. Pothoinae 103. Pothos 104. Pothoidium Subtr. Acorinae 105. Gymnostachys 106. Acorus

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Table 11. Engler’s (1876b) classification of Araceae

Names and taxon numbers are as in the original publication. Names in brackets indicate genera accepted by Schott which Engler considered better reduced to synonymy. Araceae 1. Subfam. Pothoideae Trib. Pothoeae Subtrib. Pothoinae Pothos Pothoidium Anadendron Subtrib. Heteropsinae Heteropsis ?Amydrium Subtrib. Culcasinae Culcasia Trib. Anthurieae Anthurium Trib. Zamioculcaseae Zamioculcas (Gonatopus) Trib. Symplocarpeae Lysichitum Symplocarpus Orontium Trib. Calleae Calla Trib. Acoreae Acorus Gymnostachys 2. Subfam. Monsteroideae Trib. Anepsiadeae Subtrib. Spathiphyllinae Spathiphyllum Amomophyllum (Spathiphyllopsis) Subtrib. Anepsidinae Anepsias Rhodospatha (Atimeta) Stenospermation Trib. Raphidophoreae Raphidophora Epipremnum Trib. Monstereae Scindapsus Cuscuaria Monstera (Tornelia) Alloschemone 3. Subfam. Lasioideae Trib. Lasieae Subtrib. Lasinae Cyrtosperma (Lasimorpha) Lasia Anaphyllum Subtrib. Dracontioninae Urospatha Echidnium Ophione Dracontium (Godwinia) (Chersydrium) Trib. Montrichardieae Cercestis Nephthytis Montrichardia Syngonium

Trib. Amorphophalleae Subtrib. Pythoninae Anchomanes Plesmonium Allopythion Pythonium Subtrib. Amorphophallinae Amorphophallus (Conophallus) (Proteinophallus) (Brachyspatha) Synantherias Corynophallus Hydrosme (Hansalia) Raphiophallus 4. Subfam. Philodendroideae Trib. Richardieae Richardia Trib. Peltandreae Peltandra ?Trib. Typhonodoreae Typhonodorum Trib. Philodendreae Subtrib. Homalomeninae Homalomena (Curmeria) Chamaecladon ?Adelonema Subtrib. Schismatoglottidinae Bucephalandra Schismatoglottis (Apoballis) (Colobogynium) Apatemone Subtrib. Philodendrinae Philodendron Philonotion Trib. Anubiadeae Anubias 5. Subfam. Aglaonemoideae Trib. Aglaonemeae Aglaonema ??Aglaodorum Trib. Dieffenbachieae Dieffenbachia 6. Subfam. Colocasioideae Trib. Steudnereae Steudnera Trib. Caladieae Caladium ?Xanthosoma (Acontias) (Phyllotaenium) (Andromycia) Trib. Colocasieae Subtrib. Colocasinae Colocasia (Leucocasia) Schizocasia Remusatia Subtrib. Alocasinae Alocasia Gonatanthus

7. Subfam. Staurostigmoideae Mangonia Staurostigma ?Gamochlamys Taccarum Lysistigma (Endera) 8. Subfam. Aroideae Trib. Stylochitoneae Stylochiton Trib. Zomicarpeae Zomicarpa Xenophya Trib. Ariopsideae Ariopsis Trib. Spathicarpeae Spathantheum Spathicarpa ?Gorgonidium Trib. Areae Subtrib. Arisarinae Arisarum Arisaema Pinellia Subtrib. Sauromatinae Sauromatum Subtrib. Biarinae Biarum Leptopetion (Ischarum) (Cyllenium) Subtrib. Arinae Arum (Gymnomesium) Helicodiceros Helicophyllum (Eminium) Dracunculus Theriophonum (Tapinocarpus) (Calyptrocoryne) Typhonium (Heterostalis) Trib. Ambrosinieae Ambrosinia Trib. Cryptocoryneae Lagenandra Cryptocoryne 9. Subfam. Pistioideae Pistia 10. Subfam. Lemnoideae Trib. Lemneae Spirodela Lemna Trib. Wolffieae Wolffia

APPENDIX

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Table 12. Engler’s (1920b) classification of Araceae

Names and taxon numbers are as in the original publication. Araceae Subfam. I. Pothoideae Trib. Pothoeae 1. Pothos 2. Pothoidium 3. Anadendron 3a. Epipremnopsis Trib. Heteropsideae 4. Heteropsis Trib. Anthurieae 5. Anthurium Trib. Culcasieae 6. Culcasia Trib. Zamioculcaseae 7. Zamioculcas 8. Gonatopus Trib. Acoreae 9. Acorus 10. Gymnostachys Subfam. II. Monsteroideae Trib. Monstereae 11. Raphidophora 12. Afroraphidophora 13. Epipremnum 14. Scindapsus 15. Stenospermation 16. Rhodospatha 17. Anepsias 18. Monstera 19. Alloschemone 20. Amydrium Trib. Spathiphylleae 21. Spathiphyllum 22. Holochlamys Subfam. III. Calloideae Trib. Symplocarpeae 23. Lysichitum 24. Symplocarpus 25. Orontium Trib. Calleae 26. Calla Subfam. IV. Lasioideae Trib. Lasieae 27. Cyrtosperma 28. Lasia 29. Anaphyllum 30. Podolasia 31. Urospatha 32. Dracontioides 33. Echidnium 34. Dracontium Trib. Amorphophalleae 35. Pseudohydrosme 36. Plesmonium 37. Anchomanes 38. Thomsonia 39. Pseudodracontium 40. Amorphophallus Trib. Nephthytideae 41. Nephthytis 42. Cercestis 43. Rhektophyllum Trib. Montrichardieae 44. Montrichardia

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THE GENERA OF ARACEAE

Subfam. V. Philodendroideae Trib. Philodendreae Subtrib. Homalomeninae 45. Homalomena 46. Diandriella Subtrib. Schismatoglottidinae 47. Schismatoglottis 48. Bucephalandra 49. Aridarum 50. Piptospatha 51. Microcasia Subtrib. Philodendrinae 52. Philodendron 53. Philonotion Trib. Anubiadeae 54a. Amauriella 54b. Anubias Trib. Aglaonemateae 55. Aglaonema 56. Aglaodorum Trib. Dieffenbachieae 57. Dieffenbachia Trib. Zantedeschieae 58. Zantedeschia Trib. Typhonodoreae 59. Typhonodorum Trib. Peltandreae 60. Peltandra Subfam. VI. Colocasioideae Trib. Colocasieae Subtrib. Steudnerinae 61. Steudnera 62. Remusatia 63. Gonatanthus Subtrib. Hapalininae 64. Hapaline Subtrib. Caladiinae 65. Caladiopsis 66. Caladium 67. Aphyllarum 68. Chlorospatha 69. Xanthosoma Subtrib. Colocasiinae 70. Colocasia Subtrib. Alocasiinae 71. Alocasia 72. Schizocasia Trib. Syngonieae 73. Porphyrospatha 74. Syngonium Trib. Ariopsideae 75. Ariopsis Subfam. VII. Aroideae Trib. Stylochitoneae 76. Stylochiton Trib. Asterostigmateae 77. Mangonia 78. Andromycia 79. Taccarum 80. Asterostigma 81. Synandrospadix 82. Spathantheum 83. Gorgonidium 84. Gearum 85. Spathicarpa

Trib. Protareae 86. Protarum Trib. Callopsideae 87. Callopsis Trib. Zomicarpeae 88. Scaphispatha 89. Xenophya 90. Zomicarpa 91. Zomicarpella 92. Ulearum Trib. Areae Subtrib. Arinae 93. Arum 94. Dracunculus 95. Helicodiceros 96. Theriophonum 97. Typhonium 98. Sauromatum 99. Eminium 100. Biarum Subtrib. Arisarinae 101. Arisarum Subtrib. Arisaematinae 102. Arisaema Subtrib. Pinelliinae 103. Pinellia Subtrib. Ambrosiniinae 104. Ambrosinia Subtrib. Cryptocoryninae 105. Lagenandra 106. Cryptocoryne Subfam. IX. Pistioideae 107. Pistia

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Table 13. Grayum’s (1990) classification of Araceae

Names and taxon numbers are as in the original publication. Araceae I. Subfam. Pothoideae Trib. Gymnostachydeae Gymnostachys Trib. Spathiphylleae Spathiphyllum Holochlamys Trib. Anthurieae Anthurium Trib. Potheae Pothos Pedicellarum Pothoidium Trib. Anadendreae Anadendrum Trib. Monstereae Subtrib. Heteropsideae Heteropsis Subtrib. Monsterinae Rhaphidophora Monstera Amydrium Epipremnum Scindapsus Alloschemone Stenospermation Rhodospatha Trib. Zamioculcadeae Zamioculcas Gonatopus II. Subfam. Calloideae Calla Alliance

Trib. Calleae Calla Nephthytis Alliance

Trib. Nephthytideae Nephthytis Anchomanes Pseudohydrosme Trib. Callopsideae Callopsis Ulearum Filarum Zomicarpella Trib. Montrichardieae Montrichardia Aglaonema Alliance

Trib. Anubiadeae Anubias Trib. Zantedeschieae Zantedeschia Trib. Aglaonemateae Aglaonema Aglaodorum Trib. Spathicarpeae Mangonia Asterostigma Synandrospadix Taccarum Gorgonidium Gearum Spathantheum Spathicarpa Trib. Dieffenbachieae Dieffenbachia

Trib. Bognereae Bognera Peltandra Alliance

Trib. Peltandreae Peltandra Typhonodorum Trib. Arophyteae Arophyton Carlephyton Colletogyne Trib. Schismatoglottideae Schismatoglottis Piptospatha Bucephalandra Phymatarum Aridarum Heteroaridarum Hottarum Philodendron Alliance

Trib. Culcasieae Culcasia Trib. Cercestideae Cercestis Trib. Homalomeneae Furtadoa Homalomena Trib. Philodendreae Philodendron III. Subfam. Colocasioideae Trib. Zomicarpeae Zomicarpa Trib. Colocasieae Subtrib. Protarinae Protarum Subtrib. Steudnerinae Steudnera Subtrib. Remusatiinae Remusatia Gonatanthus Subtrib. Colocasiinae Colocasia Alocasia Trib. Caladieae Subtrib. Jasarinae Jasarum Subtrib. Scaphispathinae Scaphispatha Subtrib. Caladiinae Caladium Xanthosoma Chlorospatha Aphyllarum Subtrib. Syngoniinae Syngonium Subtrib. Hapalininae Hapaline

Cyrtosperma Lasia Anaphyllum Podolasia Urospatha Dracontioides Dracontium Subtrib. Pycnospathinae Pycnospatha Trib. Stylochaetoneae Stylochaeton V. Subfam. Aroideae Trib. Thomsonieae Pseudodracontium Amorphophallus Trib. Arisareae Arisarum Trib. Pinellieae Pinellia Trib. Pistieae Pistia Trib. Cryptocoryneae Cryptocoryne Lagenandra Trib. Ambrosineae Ambrosina Trib. Ariopsideae Ariopsis Trib. Arisaemateae Arisaema Trib. Areae Arum Dracunculus Helicodiceros Theriophonum Typhonium Sauromatum Eminium Biarum

IV. Subfam. Lasioideae Trib. Symplocarpeae Lysichitum Symplocarpus Trib. Orontieae Orontium Trib. Lasieae Subtrib. Dracontiinae

APPENDIX

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Table 14. Bogner & Nicolson’s (1991) classification of Araceae

Names and taxon numbers are as in the original publication. Araceae Subfam. 1. Gymnostachydoideae 1. Gymnostachys Subfam. 2. Pothoideae 2. Pothos 3. Pedicellarum 4. Pothoidium Subfam. 3. Monsteroideae Trib. Anadendreae 5. Anadendrum Trib. Monstereae 6. Amydrium 7. Rhaphidophora 8. Epipremnum 9. Scindapsus 10. Alloschemone 11. Stenospermation 12. Rhodospatha 13. Monstera Trib. Heteropsideae 14. Heteropsis Trib. Spathiphylleae 15. Spathiphyllum 16. Holochlamys Subfam. 4. Calloideae 17. Calla Subfam. 5. Lasioideae Trib. Orontieae 18. Lysichiton 19. Symplocarpus 20. Orontium Trib. Anthurieae 21. Anthurium Trib. Lasieae Subtrib. Dracontiinae 22. Cyrtosperma 23. Lasimorpha 24. Lasia 25. Anaphyllum 26. Anaphyllopsis 27. Podolasia 28. Urospatha 29. Dracontioides 30. Dracontium Subtrib. Pycnospathinae 31. Pycnospatha Trib. Zamioculcadeae 32. Zamioculcas 33. Gonatopus Trib. Callopsideae 34. Callopsis Trib. Nephthytideae 35. Pseudohydrosme 36. Anchomanes 37. Nephthytis 38. Cercestis Trib. Culcasieae 39. Culcasia Trib. Montrichardieae 40. Montrichardia Subfam. 6. Philodendroideae Trib. Philodendreae Subtrib. Homalomeninae

324

THE GENERA OF ARACEAE

41. Furtadoa 42. Homalomena Subtrib. Schismatoglottidinae 43. Schismatoglottis 44. Piptospatha 45. Hottarum 46. Bucephalandra 47. Phymatarum 48. Aridarum 49. Heteroaridarum Subtrib. Philodendrinae 50. Philodendron Trib. Anubiadeae 51. Anubias 52. Bognera Trib. Aglaonemateae 53. Aglaonema 54. Aglaodorum Trib. Dieffenbachieae 55. Dieffenbachia Trib. Zantedeschieae 56. Zantedeschia Trib. Typhonodoreae 57. Typhonodorum Trib. Peltandreae 58. Peltandra Subfam. 7. Colocasioideae Trib. Caladieae 59. Xanthosoma 60. Chlorospatha 61. Caladium 62. Scaphispatha 63. Jasarum Trib. Steudnereae Subtrib. Steudnerinae 64. Steudnera 65. Remusatia 66. Gonatanthus Subtrib. Hapalininae 67. Hapaline Trib. Protareae 68. Protarum Trib. Colocasieae 69. Colocasia 70. Alocasia Trib. Syngonieae 71. Syngonium Trib. Ariopsideae 72. Ariopsis Subfam. 8. Aroideae Trib. Stylochaetoneae 73. Stylochaeton Trib. Arophyteae 74. Carlephyton 75. Colletogyne 76. Arophyton Trib. Spathicarpeae 77. Mangonia 78. Taccarum 79. Asterostigma 80. Gorgonidium 81. Synandrospadix 82. Gearum 83. Spathantheum 84. Spathicarpa

Trib. Zomicarpeae 85. Zomicarpa 86. Filarum 87. Zomicarpella 88. Ulearum Trib. Thomsonieae 89. Amorphophallus 90 Pseudodracontium Trib. Areae Subtrib. Arinae 91. Arum 92. Dracunculus 93. Helicodiceros 94. Theriophonum 95. Typhonium 96. Sauromatum 97. Eminium 98. Biarum Subtrib. Arisarinae 99. Arisarum Subtrib. Arisaematinae 100. Arisaema Subtrib. Atherurinae 101. Pinellia Subtrib. Ambrosininae 102. Ambrosina Subtrib. Cryptocoryninae 103. Lagenandra 104. Cryptocoryne Subfam. 9. Pistioideae 105. Pistia

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Table 15. Generic country lists

Only indigenous records are given. We have not cited genera which are exotic or naturalized in countries where they are not native (the Lesser Antilles are treated as one unit). Our citations are based on existing records in herbaria or literature; we would appreciate receiving any new country records. Acorus is excluded from this list.

Afghanistan: Arisaema, Arum, Eminium, Pistia Albania: Arisarum, Arum, Biarum, Dracunculus Algeria: Ambrosina, Arisarum, Arum, Biarum Andaman Is.: Epipremnum, Scindapsus Andorra: Arum Angola: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Pistia, Sauromatum, Stylochaeton, Zantedeschia Argentina: Anthurium, Caladium, Dieffenbachia, Gorgonidium, Philodendron, Pistia, Spathantheum, Spathicarpa, Synandrospadix, Taccarum, Xanthosoma Armenia: Arum Australia: Alocasia, Amorphophallus, Epipremnum, Gymnostachys, Lazarum, Pistia, Pothos, Remusatia, Rhaphidophora, Scindapsus, Typhonium Austria: Arum, Calla Azerbaijan: Arum

Bangladesh: Aglaonema, Alocasia, Amorphophallus, Colocasia, Cryptocoryne, Homalomena, Lagenandra, Lasia, Pistia, Pothos, Remusatia, Rhaphidophora, Sauromatum, Scindapsus, Steudnera, Typhonium Belgium: Arum, Calla Belize: Anthurium, ?Dieffenbachia, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Syngonium Belorussia: Arum Benin: Amorphophallus, ?Anubias, Cercestis, Culcasia, Lasimorpha, ?Nephthytis, ?Pistia, Stylochaeton Bhutan: Amorphophallus, Ariopsis, Arisaema, Colocasia, Remusatia, Rhaphidophora, Sauromatum, Scindapsus, Typhonium Bolivia: Anthurium, Asterostigma, Caladium, ?Dieffenbachia, Dracontium, Gorgonidium, Heteropsis, Homalomena, Monstera, Philodendron, Pistia, Rhodospatha, Scaphispatha, Spathantheum, Spathicarpa, Stenospermation, Synandrospadix, Syngonium, Taccarum, Xanthosoma Bosnia-Hercegovina: Arisarum, Arum, Dracunculus Botswana: ?Amorphophallus, Pistia, Zantedeschia Brazil: Alloschemone, Anaphyllopsis, Anthurium, Asterostigma, Bognera, Caladium, Dieffenbachia, Dracontioides, Dracontium, Gearum, Heteropsis, Homalomena, Mangonia, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Scaphispatha, Schismatoglottis, Spathicarpa, Spathiphyllum, Stenospermation, Syngonium, Taccarum, Ulearum, Urospatha, Xanthosoma, Zomicarpa, Zomicarpella Brunei: Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Aridarum, Arisaema, Bucephalandra, Cryptocoryne, Cyrtosperma, Epipremnum, Hapaline, Homalomena, Hottarum, Lasia, Phymatarum, Piptospatha, Pistia, Pothos, Rhaphidophora, Schismatoglottis, Scindapsus Bulgaria: Arum, Dracunculus Burkina Faso: Amorphophallus, Culcasia, Pistia, Stylochaeton Burma: Aglaonema, Alocasia, Amorphophallus, Amydrium, Ariopsis, Arisaema, Colocasia, Cryptocoryne, Epipremnum, Hapaline, Homalomena, Lasia, Pothos, Rhaphidophora, Sauromatum, Schismatoglottis, Scindapsus, Steudnera, Typhonium Burundi: ?Amorphophallus, Arisaema, ?Cercestis, ?Culcasia, Pistia

Cabinda: ?Amorphophallus, ?Anchomanes, Anubias, ?Cercestis, Culcasia, Stylochaeton Cambodia: Aglaonema, Alocasia, Amorphophallus, Anadendrum, Arisaema, Colocasia, Cryptocoryne, Epipremnum, ?Hapaline, Homalomena, Lasia, Pistia, Pothos, Pseudodracontium, Rhaphidophora, Schismatoglottis, Scindapsus, ?Steudnera, Typhonium Cameroon: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Remusatia, Rhaphidophora, Sauromatum, Stylochaeton Canada: Arisaema, Calla, Lysichiton, Peltandra, Symplocarpus Caroline Is.: Cyrtosperma, Epipremnum, Rhaphidophora, Scindapsus, Typhonium Central African Republic: Amorphophallus, Anchomanes, ?Anubias, Cercestis, Culcasia, Lasimorpha, Pistia, Sauromatum, Stylochaeton Chad: Amorphophallus, Anchomanes, Lasimorpha, Pistia, ?Stylochaeton China: Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Arisaema, Arum, Calla, Colocasia, Cryptocoryne, Cyrtosperma, Epipremnum, Hapaline, Homalomena, Lasia, Pinellia, Pistia, Pothoidium, Pothos, Remusatia, Rhaphidophora, Sauromatum, Schismatoglottis, Scindapsus, Steudnera, Symplocarpus, Typhonium Christmas Is.: Remusatia Colombia: Anthurium, Caladium, Chlorospatha, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Schismatoglottis, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma, ?Zomicarpella Comores Is.: Pistia, Pothos, Typhonodorum Congo: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia Cook Is.: Cyrtosperma Costa Rica: Anthurium, Caladium, Chlorospatha, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Croatia: Arisarum, Arum, Biarum, Calla, Dracunculus Cuba: Anthurium, Philodendron, Pistia, Xanthosoma Cyprus: Arisarum, Arum Czech Republic: Arum, Calla Denmark: Arum, Calla Dominican Republic: Anthurium, Dieffenbachia, Philodendron, Pistia, Xanthosoma Ecuador: Anthurium, Asterostigma, Caladium, Chlorospatha, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Stenospermation, Syngonium, Xanthosoma Egypt: Arisarum, Biarum, Eminium, Pistia El Salvador: Anthurium, Dieffenbachia, Monstera, Philodendron, Spathiphyllum, Syngonium, Xanthosoma Estonia: Calla Equatorial Guinea (Bioko): Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Rhaphidophora, Sauromatum, Stylochaeton

APPENDIX

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Equatorial Guinea (Rio Muni): Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Rhaphidophora, Stylochaeton Ethiopia: Amorphophallus, Arisaema, Pistia, Remusatia, Sauromatum, Stylochaeton Europe: Ambrosina, Arisarum, Arum, Biarum, Calla, Dracunculus, Helicodiceros Fiji: Alocasia, Cyrtosperma, Epipremnum, Rhaphidophora, Scindapsus Finland: Calla France: Arisarum, Arum, Calla, Helicodiceros French Guiana: Anaphyllopsis, Anthurium, Caladium, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Rhodospatha, Schismatoglottis, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Gabon: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pseudohydrosme, Rhaphidophora, Stylochaeton Gambia: Amorphopallus, ?Anubias, Anchomanes, Cercestis, ?Culcasia, Lasimorpha, Pistia Georgia: Arum Germany: Arum, Calla Ghana: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Rhaphidophora, Stylochaeton Gilbert & Ellice Is.: Cyrtosperma Greece: Arisarum, Arum, Biarum, Dracunculus Guam: Cyrtosperma Guatemala: Anthurium, Dieffenbachia, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Guinea: Amorphophallus, ?Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Pistia, Remusatia, Stylochaeton Guinea-Bissau: Amorphophallus, ?Anchomanes, ?Anubias, Cercestis, ?Culcasia, Lasimorpha Guyana: Anthurium, Caladium, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Jasarum, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, ?Schismatoglottis, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Haiti: Anthurium, Dieffenbachia, Pistia, Syngonium, Xanthosoma Honduras: Anthurium, Dieffenbachia, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Syngonium, Xanthosoma Hungary: Arum, ?Calla India: Aglaonema, Alocasia, Amorphophallus, Anaphyllum, Ariopsis, Arisaema, Arum, Colocasia, Cryptocoryne, Homalomena, Lagenandra, Lasia, Pistia, Pothos, Remusatia, Rhaphidophora, Sauromatum, Scindapsus, Steudnera, Theriophonum, Typhonium Indonesia: Aglaodorum, Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Arisaema, Bucephalandra, Colocasia, Cryptocoryne, Cyrtosperma, Epipremnum, Furtadoa, Holochlamys, Homalomena, Hottarum, Lasia, Pedicellarum, Phymatarum, Piptospatha, Pistia, Podolasia, Pothoidium, Pothos, Remusatia, Rhaphidophora, Schismatoglottis, Scindapsus, Spathiphyllum, Typhonium Iran: Arum, Biarum, Eminium Iraq: Arum, Biarum, Eminium Ireland: Arum

326

THE GENERA OF ARACEAE

Israel: Arisarum, Arum, Biarum, Eminium Italy: Ambrosina, Arisarum, Arum, Biarum, Dracunculus, Helicodiceros Ivory Coast: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Remusatia, Rhaphidophora, Stylochaeton Jamaica: Anthurium, Dieffenbachia, Philodendron, Pistia, Syngonium, Xanthosoma Japan: Alocasia, Amorphophallus, Arisaema, Epipremnum, Lysichiton, Pinellia, Pothos, Rhaphidophora, Symplocarpus, Typhonium Jordan: Arum, Biarum, Eminium Kazakhstan: Arum, Eminium Kenya: Amorphophallus, Anchomanes, Arisaema, Callopsis, Culcasia, Gonatopus, Pistia, Sauromatum, Stylochaeton, Zamioculcas Kirghizia: Arum, Eminium Korea N.: Arisaema, Calla, Pinellia, Symplocarpus, Korea S.: Arisaema, Pinellia, Symplocarpus Laos: Aglaonema, Alocasia, Amorphophallus, Anadendrum, Arisaema, Colocasia, Cryptocoryne, Hapaline, Homalomena, Lasia, Pistia, Pothos, Pseudodracontium, Pycnospatha, Rhaphidophora, Schismatoglottis, ?Scindapsus, Steudnera, Typhonium Lebanon: Arisarum, Arum, Biarum, Eminium Lesotho: Pistia, Zantedeschia Lesser Antilles: Anthurium, Caladium, Dieffenbachia, Monstera, Montrichardia, Philodendron, Pistia, ?Syngonium, Xanthosoma Liberia: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Remusatia, Rhaphidophora, ?Stylochaeton Libya: Arisarum, Arum, Biarum Liechtenstein: Arum Luxembourg: Arum Macedonia: Arisarum, Arum, Biarum, Dracunculus Madagascar: Amorphophallus, Arophyton, Carlephyton, Colletogyne, Pistia, Pothos, Remusatia, Typhonodorum Malawi: Amorphophallus, ?Anchomanes, Gonatopus, Pistia, Sauromatum, Stylochaeton, Zamioculcas, Zantedeschia Malaysia: Aglaodorum, Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Aridarum, Arisaema, Bucephalandra, Colocasia, Cryptocoryne, Cyrtosperma, Epipremnum, Furtadoa, Hapaline, Heteroaridarum, Homalomena, Hottarum, Lasia, Nephthytis, Pedicellarum, Phymatarum, Piptospatha, Pistia, Podolasia, Pothos, Rhaphidophora, Schismatoglottis, Scindapsus, Typhonium Mali: Amorphophallus, Anubias, Culcasia, Pistia, Stylochaeton Mariana Is.: Cyrtosperma, Typhonium Marquesas Is.: Cyrtosperma Marshall Is.: Cyrtosperma, Epipremnum Mauritius: Pistia, Typhonodorum Mexico: Anthurium, Arisaema, Dieffenbachia, Dracontium, Monstera, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Syngonium, Xanthosoma Moldavia: Arum Mongolia: Typhonium Morocco: Arisarum, Arum, Biarum Mozambique: Amorphophallus, Anchomanes, Gonatopus, Pistia, Stylochaeton, Zamioculcas, ?Zantedeschia

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Namibia: ?Amorphophallus, Zantedeschia Nepal: Alocasia, Amorphophallus, Ariopsis, Arisaema, Arum, Colocasia, Lasia, Pistia, Pothos, Remusatia, Rhaphidophora, Sauromatum, Scindapsus, Typhonium Netherlands: Arum, Calla New Caledonia: Rhaphidophora Nicaragua: Anthurium, Dieffenbachia, Dracontium, Heteropsis, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Niger: Culcasia, Pistia Nigeria: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Remusatia, Rhaphidophora, Stylochaeton Norway: Calla Oman: Arisaema, Remusatia Pakistan: Arisaema, Arum, Pistia, Sauromatum, Typhonium Panama: Anthurium, Caladium, Chlorospatha, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Papua New Guinea: Aglaonema, Alocasia, Amorphophallus, Amydrium, Cryptocoryne, Cyrtosperma, Epipremnum, Holochlamys, Homalomena, Lasia, Pistia, Pothos, Rhaphidophora, Schismatoglottis, Scindapsus, Spathiphyllum, Typhonium Paraguay: Anthurium, Dieffenbachia, Dracontium, Philodendron, Pistia, Spathicarpa, Synandrospadix, Taccarum, Xanthosoma Peru: Anthurium, Asterostigma, ?Bognera, Caladium, Chlorospatha, Dieffenbachia, Dracontium, Filarum, Gorgonidium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Schismatoglottis, Spathantheum, Spathiphyllum, Stenospermation, Synandrospadix, Syngonium, Taccarum, Ulearum, Urospatha, Xanthosoma Philippines: Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Arisaema, Cryptocoryne, Cyrtosperma, Epipremnum, Homalomena, Pistia, Pothoidium, Pothos, Rhaphidophora, Schismatoglottis, Scindapsus, Spathiphyllum, Typhonium Poland: Arum, Calla Portugal: Arisarum, Arum, Biarum, Dracunculus Puerto Rico: Anthurium, Caladium, Dieffenbachia, Dracontium, Montrichardia, Philodendron, Pistia, Xanthosoma Romania: Arum, Calla Russia: Arisaema, Arum, Calla, Lysichiton, Symplocarpus Rwanda: ?Amorphophallus, Arisaema, ?Cercestis, ?Culcasia Samoa: Cyrtosperma, Rhaphidophora, Scindapsus San Morino: Arum Saudi Arabia: Arisaema, ?Remusatia, Sauromatum Senegal: Amorphophallus, ?Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Pistia, Stylochaeton Serbia: Arisarum, Arum, ?Dracunculus Seychelles Is.: Protarum Sierra Leone: Amorphophallus, Anchomanes, Anubias, Cercestis, Culcasia, Lasimorpha, Nephthytis, Pistia, Remusatia, Rhaphidophora, Stylochaeton Singapore: Cryptocoryne, Cyrtosperma, Epipremnum, Homalomena, Lasia, Rhaphidophora, Typhonium

Slovak Republic: Arum, Calla Slovenia: Arisarum, Arum, Calla, Dracunculus Society Is.: Cyrtosperma Solomon Is.: Alocasia, Cyrtosperma, Epipremnum, Homalomena, Pothos, Rhaphidophora, Schismatoglottis, Scindapsus, Spathiphyllum Somalia: Amorphophallus, Arisaema, Pistia, Stylochaeton South Africa: Amorphophallus, Gonatopus, Pistia, Stylochaeton, Zamioculcas, Zantedeschia Spain: Arisarum, Arum, Biarum, Dracunculus, Helicodiceros Sri Lanka: Alocasia, Amorphophallus, Arisaema, Colocasia, Cryptocoryne, Epipremnum, Lagenandra, Lasia, Pistia, Pothos, Remusatia, Rhaphidophora, Scindapsus, Theriophonum, Typhonium Sudan: Amorphophallus, Anchomanes, Arisaema, Pistia, Stylochaeton Surinam: Anaphyllopsis, Anthurium, Caladium, Dieffenbachia, Dracontium, Heteropsis, Homalomena, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Schismatoglottis, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma Swaziland: Pistia, Stylochaeton, Zantedeschia Sweden: Arum, Calla Switzerland: Arum, Calla Syria: Arisarum, Arum, Biarum, Eminium Tahiti: Cyrtosperma Tajikistan: Arum, Eminium Tanzania: Amorphophallus, Anchomanes, Arisaema, Callopsis, Culcasia, Gonatopus, Pistia, Remusatia, Sauromatum, Stylochaeton, Typhonodorum, Zamioculcas, Zantedeschia Thailand: Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Arisaema, Colocasia, Cryptocoryne, Epipremnum, Hapaline, Homalomena, Lasia, Piptospatha, Pistia, Pothos, Pseudodracontium, Pycnospatha, Remusatia, Rhaphidophora, Schismatoglottis, Scindapsus, Steudnera, Typhonium Tibet: Typhonium Togo: Amorphophallus, Anchomanes, Anubias, ?Cercestis, Culcasia, ?Nephthytis, Pistia, Rhaphidophora, Stylochaeton Trinidad & Tobago: Anthurium, Dieffenbachia, Dracontium, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Spathiphyllum, Syngonium, Xanthosoma Tunisia: Arisarum, Arum, Biarum Turkey: Arisarum, Arum, Biarum, Calla, Dracunculus, Eminium Turkmenestan: Arum, Eminium Uganda: Amorphophallus, Anchomanes, Arisaema, Cercestis, Culcasia, Pistia, Rhaphidophora, Sauromatum, Stylochaeton Ukraine: Arum United Kingdom: Arum USA: Arisaema, Calla, Lysichiton, Orontium, Peltandra, Pistia, Symplocarpus Uruguay: Anthurium, Mangonia, Philodendron, Pistia, Spathicarpa Uzbekistan: Arum, Eminium Vanuatu: Cyrtosperma, Epipremnum, Pothos, Rhaphidophora, Schismatoglottis Venezuela: Anaphyllopsis, Anthurium, Caladium, Dieffenbachia, Dracontium, Heteropsis, ?Homalomena, Jasarum, Monstera, Montrichardia, Philodendron, Pistia, Rhodospatha, Schismatoglottis, Spathiphyllum, Stenospermation, Syngonium, Urospatha, Xanthosoma

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Vietnam: Aglaodorum, Aglaonema, Alocasia, Amorphophallus, Amydrium, Anadendrum, Arisaema, Colocasia, Cryptocoryne, Cyrtosperma, Epipremnum, Hapaline, Homalomena, Lasia, Pistia, Pothos, Pseudodracontium, Pycnospatha, Remusatia, Rhaphidophora, Schismatoglottis, Scindapsus, Steudnera, Typhonium Yemen Republic: Arisaema, Remusatia (also Socotra), Sauromatum

328

THE GENERA OF ARACEAE

Zaïre: Amorphophallus, Anchomanes, Anubias, Arisaema, Cercestis, Culcasia, Gonatopus, Lasimorpha, Pistia, Remusatia, ?Sauromatum, Stylochaeton, ?Zantedeschia Zambia: Amorphophallus, Anchomanes, Gonatopus, Pistia, Remusatia, Sauromatum, Stylochaeton, Zantedeschia Zimbabwe: Amorphophallus, Gonatopus, Pistia, Stylochaeton, Zamioculcas, Zantedeschia

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Table 16. Colour plates: photo credits and vouchers Acorus calamus L. 130D: J. Bogner. Slide from collection of J. Bogner.

Callopsis volkensii Engl. 124B: J. Bogner. Slide from collection of J. Bogner.

Aglaodorum griffithii (Schott) Schott 123A: J. Bogner. Slide from collection of J. Bogner.

Carlephyton glaucophyllum Bogner 125C: J. Bogner. Slide from collection of J. Bogner.

Aglaonema tenuipes Engl. 122D: J. Bogner. Slide from collection of J. Bogner.

Cercestis ivorensis A.Chev. 123C: J. Bogner. Slide from collection of J. Bogner.

Alocasia brisbanensis (F.M. Bailey) Domin 130B: J. Bogner. Slide from collection of J. Bogner

Chlorospatha longipoda (K. Krause) Madison 121B: J. Bogner. Slide from collection of J. Bogner.

Ambrosina bassii L. 126D: J. Bogner. Slide from collection of J. Bogner

Colletogyne perrieri Buchet 125D: J. Bogner. Slide from collection of J. Bogner.

Amorphophallus aphyllus (Hook.) Hutch. 124D: Eggers. Slide from collection of R.B.G. Kew.

Colocasia esculenta (L.) Schott 130A: J. Bogner. Slide from collection of J. Bogner.

Amorphophallus prainii Hook.f. 124C: J. Bogner. Slide from collection of J. Bogner.

Cryptocoryne longicauda Engl. 119B: P. Boyce 459. Slide from collection of P. Boyce.

Amydrium medium (Zoll. & Moritzi) Nicolson 109A: P. Boyce 403. Slide from collection of P. Boyce.

Culcasia saxatilis A.Chev. 123B: S. Mayo. Slide from collection of R.B.G. Kew.

Anaphyllopsis americana (Engl.) A. Hay 111B: F.B. 4700. Slide from collection of R.B.G. Kew.

Cyrtosperma carrii A. Hay 112A: J. Bogner. Slide from collection of J. Bogner.

Anaphyllum beddomei Schott 111D: J. Bogner. Slide from collection of J. Bogner.

Dieffenbachia maculata (Lodd.) G. Don 114B: R. Zabeau. Slide from collection of R.B.G. Kew.

Anchomanes abbreviatus Engl. 122B: R. Zabeau of Faulkner 878. Slide from collection of R.B.G. Kew.

Dracontioides desciscens (Schott) Engl. 111A: S. Mayo of R. Harley et al. 18009. Slide from collection of R.B.G. Kew.

Anthurium flavolineatum Sodiro 108C: J. Bogner. Slide from collection of J. Bogner.

Dracontium changuango G.S. Bunting 110D: J. Bogner. Slide from collection of J. Bogner.

Anubias gigantea A.Chev. 117B: J. Bogner. Slide from collection of J. Bogner.

Dracunculus vulgaris Schott 127C: S. Andrews. Slide from collection of R.B.G. Kew.

Aridarum annae Bogner 118D: J. Bogner 1400. Slide from collection of J. Bogner.

Eminium lehmannii (Regel) Kuntze 127B: P. Furze 35/3. Slide from collection of R.B.G. Kew.

Ariopsis peltata Nimmo 129B: J. Bogner. Slide from collection of J. Bogner.

Epipremnum falcifolium Engl. 109C: A. Poulsen 363. Slide from collection of R.B.G. Kew.

Arisaema ovale Nakai var. ovale 129A: A. McRobb. Slide from collection of R.B.G. Kew.

Furtadoa sumatrensis M. Hotta 116D: D. Bown 195/2. Slide from collection of R.B.G. Kew.

Arisarum simorrhinum Durieu ex Duch. 126C: P. Boyce. Slide from collection of P. Boyce.

Gonatopus marattioides (Peter) Bogner 113D: R. Zabeau of Bogner 247. Slide from collection of R.B.G. Kew.

Arophyton crassifolium (Buchet) Bogner 125B: J. Bogner. Slide from collection of J. Bogner. Arum maculatum L. 127B: P. Boyce. Slide from collection of P. Boyce. Asterostigma riedelianum (Schott) Kuntze 115B: R. Harley et al. 18565. Slide from collection of R.B.G. Kew.

Gorgonidium vermicidum (Speg.) Bogner 115C: J. Bogner. Slide from collection of J. Bogner. Gymnostachys anceps R.Br. 107A: Unknown. Slide from collection of J. Bogner. Hapaline celatrix P.C. Boyce 121D: P. Boyce 417. Slide from collection of P. Boyce.

Biarum ditschianum Bogner & P.C. Boyce 128C: W. Barthlott. Slide from collection of J. Bogner.

Helicodiceros muscivorus (L.f.) Engl. 127D: P. Boyce 36. Slide from collection of P. Boyce.

Bognera recondita (Madison) Mayo & Nicolson 114C: J. Bogner 1995. Slide from collection of J. Bogner.

Homalomena geniculata M. Hotta 117A: J. Bogner. Slide from collection of J. Bogner.

Bucephalandra motleyana Schott 118B: J. Bogner 1366. Slide from collection of J. Bogner.

Hottarum lucens Bogner 118A: J. Bogner 1439. Slide from collection of J. Bogner.

Caladium tuberosum (S. Moore) Bogner & Mayo 120C: R. Harley 10980. Slide from collection of R.B.G, Kew.

Jasarum steyermarkii G.S. Bunting 120D: H. Herkner. Slide from collection of J. Bogner.

Calla palustris L. 113B: D. Bown. Slide from collection of R.B.G, Kew

Lagenandra nairii Ramam. & Rajan 119A: J. Bogner. Slide from collection of J. Bogner.

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Lasia spinosa (L.) Thwaites 112D: J. Bogner. Slide from collection of J. Bogner.

Scaphispatha gracilis Brongn. ex Schott 120B: J. Bogner 1211. Slide from collection of J. Bogner.

Lasimorpha senegalensis Schott 112B: J. Bogner. Slide from collection of J. Bogner.

Schismatoglottis ferruginea Merr. 117C: J. Dransfield 6871. Slide from collection of R.B.G. Kew.

Lysichiton camtschatcensis Schott 107C: J. Bogner. Slide from collection of J. Bogner.

Scindapsus beccarii Engl. 109D: P. Boyce 318. Slide from collection of P. Boyce.

Mangonia tweedieana Schott 114D: J. Waechter 2347. Slide from collection of R.B.G. Kew.

Spathantheum intermedium Bogner 116A: J. Bogner. Slide from collection of J. Bogner.

Monstera adansonii Schott 110A: R. Zabeau. Slide from collection of R.B.G. Kew.

Spathicarpa gardneri Schott 116B: J. Bogner. Slide from collection of J. Bogner.

Montrichardia linifera (Arruda Câmara) Schott 123D: S. Mayo. Slide from collection of R.B.G. Kew.

Spathiphyllum floribundum N.E.Br. 108D: R. Zabeau. Slide from collection of R.B.G. Kew.

Nephthytis afzelii Schott var. graboensis Bogner 122A: J. Bogner. Slide from collection of J. Bogner.

Stenospermation multiovulatum (Engl.) N.E.Br. 110C: P. Boyce. Slide from collection of P. Boyce.

Orontium aquaticum L. 107B: J. Bogner. Slide from collection of J. Bogner.

Steudnera colocasiifolia K. Koch 129C: J. Bogner. Slide from collection of J. Bogner.

Pedicellarum paiei M. Hotta 108B: A. Church 303. Slide from collection of R.B.G. Kew.

Stylochaeton salaamicus N.E.Br. 114A: R. Zabeau of Bogner 141. Slide from collection of R.B.G. Kew.

Peltandra virginica (L.) Raf. 126A: J. Bogner. Slide from collection of J. Bogner. Philodendron rugosum Bogner & G.S. Bunting 116C: J. Bogner 1522. Slide from collection of J. Bogner. Phymatarum borneense M. Hotta 118C: J. Bogner 1476. Slide from collection of J. Bogner. Pinellia cordata N.E.Br. 128D: J. Bogner. Slide from collection of J. Bogner. Piptospatha burbidgei (N.E.Br.) M. Hotta 117D: A. Poulsen. Slide from collection ofR.B.G. Kew. Pistia stratiotes L. 130C: D. Bown 203/3. Slide from collection of R.B.G. Kew. Podolasia stipitata N.E.Br. 112C: J. Bogner. Slide from collection of J. Bogner. Pothos macrocephalus Scort. ex Hook.f. 108A: J. Bogner. Slide from collection of J. Bogner. Pseudodracontium lacourii (Lind. & André) N.E.Br. 125A: J. Bogner. Slide from collection of J. Bogner. Pseudohydrosme gabunensis Engl. 122C: J. Bogner. Slide from collection of J. Bogner. Pycnospatha arietina Thorel ex Gagnep. 111C: J. Bogner 395. Slide from collection of J. Bogner. Remusatia vivipara (Roxb.) Schott 129D: R. Zabeau of Faulkner 286. Slide from collection of R.B.G. Kew. Rhaphidophora foraminifera (Engl.) Engl. 109B: P. Boyce 235. Slide from collection of P. Boyce. Rhodospatha perezii G.S. Bunting 110B: T. Croat. Slide from collection of T. Croat. Sauromatum venosum (Aiton) Kunth 128B: Unknown. Slide from collection of R.B.G. Kew.

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Symplocarpus renifolius Schott ex Miq. 107D: J. Bogner. Slide from collection of J. Bogner. Synandrospadix vermitoxicus (Griseb.) Engl. 115D: J. Bogner. Slide from collection of J. Bogner. Syngonium steyermarkii Croat 121C: J. Bogner. Slide from collection of J. Bogner. Taccarum weddellianum Brongn. ex Schott 115A: J. Bogner. Slide from collection of R.B.G. Kew. Typhonium trilobatum (L.) Schott 128A: A. McRobb of A. Hay 2045. Slide from collection of R.B.G. Kew. Typhonodorum lindleyanum Schott 126B: P. Cribb. Slide from collection of R.B.G. Kew. Ulearum sagittatum Engl. var. viridispadix Bogner 120A: J. Bogner. Slide from collection of J. Bogner. Urospatha tonduzii Engl. 113A: D. Bown 196/6. Slide from collection of R.B.G. Kew. Xanthosoma violaceum Schott 121A: J. Bogner. Slide from collection of J. Bogner. Zamioculcas zamiifolia (Lodd.) Engl. 113C: R. Zabeau. Slide from collection of R.B.G. Kew. Zantedeschia aethiopica (L.) Spreng. 124A: P. Boyce. Slide from collection of P. Boyce. Zomicarpa riedeliana Schott 119C: J. Bogner. Slide from collection of J. Bogner. Zomicarpella amazonica Bogner 119D: J. Bogner. Slide from collection of J. Bogner.

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31 I N D E X TO S C I E N T I F I C N A M E S

References to all scientific names given in pp. v-75 are listed here; for pp. 76-290 only species binomials and major references are given; from p. 291 onwards, scientific names are not listed in this index. Generic distribution maps are not listed here but are always to be found next to their corresponding plate(s) or main description. Numbers in bold refer to main descriptions of taxa, except when preceded by “c/plate” in which case they refer to colour plate numbers; numbers in bold italic refer to line drawings. Names in bold refer to taxa which receive main treatments in the text or to species which are illustrated; plant names in italic are synonyms; all authors’ names are in italic. A Acacia Mill., 19 Acontias Schott, 209 Acoraceae C. Agardh, ix, x, 5 13, 17, 22, 23, 29, 30, 32, 36, 37, 38, 42, 47, 54, 55, 58, 61, 68, 289 Acoreae Endl., 73 Acorites Crepet, 59 Acorites heeri (E.W. Berry) Crepet, 58 Acoroideae, 72, 74 Acoropsis Conw., 59 Acoropsis eximia (Göpp. & Menge) Bogner, 59 Acoropsis minor Conw., 59 Acorus L., c/plate 130D, 5, 15, 19, 22, 23, 27, 29, 30, 32, 36, 42, 47, 54, 55, 58, 59, 60, 61, 62, 72, 73, 74, 289, 290 Acorus brachystachys Heer, 58, 59 Acorus calamus L., c/plate 130D, 17, 19, 36, 37, 38, 42, 55, 59, 289, 290 Acorus calamus L. var. americanus (Raf.) Wulff, 19 Acorus calamus L. var. calamus, 19 Acorus gramineus Sol., 19, 36, 42, 289 Acorus heeri E.W. Berry, 58 Acorus procalamus Nikitin, 59 Adelonema Schott, 177 Afrorhaphidophora Engl., 118 Aglaodorum Schott, c/plate 123A, 34, 40, 225, 226 Aglaodorum griffithii (Schott) Schott, c/plate 123A, 225, 226 Aglaonema Schott, c/plate 122D, 5, 11, 14, 21, 34, 40, 44, 55, 59, 223, 224 Aglaonema commutatum Schott, ix, 56 Aglaonema griffithii Schott, 225 Aglaonema hookerianum Schott, 224 Aglaonema integrifolium (Link) Schott, 223 Aglaonema modestum Schott ex Engl., 224 Aglaonema nitidum (Jack) Kunth, 56 Aglaonema oblongifolium Kunth, 223 Aglaonema simplex Blume, 224 Aglaonema tenuipes Engl., c/plate 122D Aglaonemateae Engl., 18, 26, 34, 40, 44, 68, 223 Aglaonemeae Engl., 223 Alismataceae Vent., 36 Alismatidae Takht., 38 Alismatiflorae Dahlgren, Clifford & Yeo, 22, 36, 37, 43, 61, 62, 63, 64, 66 Allopythion Schott, 235 Alloschemone Schott, 21, 32, 125, 126 Alloschemone occidentalis (Poepp.) Engl. & K. Krause, 125, 126 Alloschemone poeppigiana Schott, 125 Alocasia (Schott) G. Don, c/plate 130B, ix, 6, 7, 21, 22, 23, 24, 27, 28, 34, 37, 44, 47, 48, 55, 56, 57, 69, 283, 284-285 Alocasia beccarii Engl., 284, 285 Alocasia brancifolia (Schott) A. Hay, 284 Alocasia brisbanensis (F.M. Bailey) Domin, c/plate 130B, 50, 82, 286

Alocasia cucullata (Lour.) G. Don, 283, 286 Alocasia fornicata (Roxb.) Schott, 42 Alocasia guttata N.E.Br., 284 Alocasia lowii Hook., 56, 284 Alocasia macrorrhizos (L.) G. Don, 40, 47, 52, 53, 55, 56, 286 Alocasia odora (Roxb.) K. Koch, 37, 56 Alocasia plumbea Van Houtte, 286 Alocasia portei Becc. & Engl., 56 Alocasia sanderiana W. Bull, 56, 57 Alocasia zebrina K. Koch & Veitch, 56 Alocasiophyllum Engl., 228 Alpinia Roxb., 60 Amauriella Rendle, 180 Ambrosina Bassi, c/plate 126D, 7, 25, 34, 48, 71, 72, 252, 253 Ambrosina bassii L., c/plate 126D, 252, 253 Ambrosineae Schott, 18, 28, 34, 69, 252 Ambrosinia L., 252 Ambrosinieae Schott, 252 Amidena Adans., 91 Amomophyllum Engl., 110 Amorphophalleae Engl., 74 Amorphophallus Blume ex Decne., c/plate 124C–D, 7, 11, 13, 18, 21, 24, 25, 26, 27, 31, 34, 36, 37, 40, 45, 47, 48, 49, 52, 56, 73, 235, 236-239 Amorphophallus abyssinicus (A.Rich.) N.E.Br., 45 Amorphophallus albispathus Hett., 45, 238 Amorphophallus aphyllus (Hook.) Hutch., c/plate 124D Amorphophallus brooksii Alderw., 50 Amorphophallus bulbifer Blume, 6, 18, 36, 38, 49, 80, 239 Amorphophallus campanulatus Decne., 36, 37, 42, 235 Amorphophallus corrugatus N.E.Br., 239 Amorphophallus coudercii (Bogner) Bogner, 86 Amorphophallus decus-silvae Alderw., 236 Amorphophallus dracontioides N.E.Br., 236, 237, 238 Amorphophallus elliotii Hook.f., 239 Amorphophallus galbra F.M. Bailey, 47, 236 Amorphophallus gomboczianus Pic.Serm., 237, 238 Amorphophallus henryi N.E.Br., 239 Amorphophallus hildebrandtii (Engl.) Engl. & Gehrm., 45 Amorphophallus hirtus N.E.Br., 237, 238 Amorphophallus johnsonii N.E.Br., 239 Amorphophallus kerrii N.E.Br., 82 Amorphophallus kiusianus (Makino) Makino, 36 Amorphophallus konjac K. Koch, ix, 21, 23, 36, 47, 53, 55, 56, 239 Amorphophallus krausei Engl., 236, 237 Amorphophallus lambii Mayo & Widjaja, 50 Amorphophallus lewallei Malaisse & Bamps, 237 Amorphophallus longituberosus Engl. & Gehrm., 239 Amorphophallus maculatus N.E.Br., 35, 45, 236, 239 Amorphophallus manta Hett. & Ittenb., 47 Amorphophallus margaritifer Kunth, 82, 86 Amorphophallus oncophyllus Prain ex Hook.f., 36, 37 Amorphophallus paeoniifolius (Dennst.) Nicolson, 36, 37, 42, 50, 53, 55, 235, 236, 237, 238, 239

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Amorphophallus parvulus Gagnep., 239 Amorphophallus pendulus Bogner & Mayo, 237, 238 Amorphophallus prainii Hook.f. c/plate 124C, 53 Amorphophallus pusillus Hett. & Serebryanyi, 235, 237, 238 Amorphophallus pygmaeus Hett., 237, 238 Amorphophallus rivieri Durieu, 21, 23, 36, 53 Amorphophallus sumawongii (Bogner) Bogner & Mayo, 237, 238 Amorphophallus titanum Becc., ix, 45, 50 Amorphophallus variabilis Blume, 36, 37 Amydrium Schott, c/plate 109A, 6, 14, 16, 20, 32, 116, 117 Amydrium humile Schott, 116, 117, 118 Amydrium medium (Zoll. & Moritzi) Nicolson, c/plate 109A, 116, 117 Amydrium zippelianum (Schott) Nicolson, 116, 117, 311 Anadendreae Bogner & French, 32, 67, 68, 78, 110 Anadendron Schott, 113 Anadendrum Schott, 16, 19, 20, 26, 32, 35, 67, 70, 74, 75, 113, 114 Anadendrum microstachyum (de Vriese & Miq.) Backer & Alderw., 113, 114 Anadendrum montanum Schott, 113 Ananas comosus (L.) Merr., 41 Anaphyllopsis A. Hay, c/plate 111B, ix, 7, 32, 133, 134 Anaphyllopsis americana (Engl.) A. Hay, c/plate 111B, 133, 134 Anaphyllopsis cururuana A. Hay, 134 Anaphyllopsis pinnata A. Hay, 134 Anaphyllum Schott, c/plate 111D, 7, 32, 73, 135, 137 Anaphyllum beddomei Schott, c/plate 111D, 137 Anaphyllum wightii Engl., 135, 137 Anarmodium Schott, 257 Anatropooae Schott, 72 Anchomanes Schott, c/plate 122B, 7, 11, 17, 18, 33, 40, 48, 50, 56, 73, 218, 220 Anchomanes abbreviatus Engl., c/plate 122B, 220 Anchomanes boehmii Engl., 220, 221 Anchomanes difformis (Blume) Engl., 38, 220 Anchomanes hookeri Schott, 218 Anchomanes nigritianus Rendle, 220 Anchomanes petiolatus (Hook.) Hutch., 218, 220 Anchomanes welwitschii Rendle, 220, 221 Androgynanthae Schott, 72 Andromycia A.Rich., 160 Anepsias Schott, 125 Anthelia Schott, 121 Anthomyiidae, 48 Anthurieae Engl., 32, 68, 103 Anthuriophyllum Weyland, 58 Anthuriophyllum spectabile Weyland, 58 Anthurium Schott, c/plate 108C, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17, 18, 21, 22, 23, 26, 28, 31, 32, 37, 40, 44, 45, 46, 47, 48, 49, 50, 51, 54, 55, 56, 60, 67, 73, 74, 75, 103, 104-108 Anthurium sect. Anthurium, 109 Anthurium sect. Belolonchium Schott, 109 Anthurium sect. Calomystrium Schott, 109 Anthurium sect. Cardiolonchium Schott, 109 Anthurium sect. Chamaerepium Schott, 109 Anthurium sect. Dactylophyllium Schott, 109 Anthurium sect. Digitinervium Sodiro, 67, 81, 109 Anthurium sect. Episeiostenium Schott, 109 Anthurium sect. Gymnopodium Engl., 109 Anthurium sect. Leptanthurium Schott, 109 Anthurium sect. Oxycarpium Schott, 109 Anthurium sect. Pachyneurium Schott, 10, 11, 45, 80, 109 Anthurium sect. Polyneurium Engl., 109 Anthurium sect. Polyphyllium Engl., 10, 103, 109 Anthurium sect. Porphyrochitonium Schott, 109 Anthurium sect. Schizoplacium Schott, 109 Anthurium sect. Semaeophyllium Schott, 109 Anthurium sect. Tetraspermium Schott, 109 Anthurium sect. Urospadix Engl, 109 Anthurium sect. Xialophyllium Schott, 109 Anthurium acaule (Jacq.) Schott, 103 Anthurium aemulum Schott, 40

332

THE GENERA OF ARACEAE

Anthurium affine Schott, 11, 107 Anthurium andicola Liebm., 11 Anthurium andraeanum Linden, ix, 47, 56, 107 Anthurium antrophyoides Killip, 104 Anthurium bakeri Hook.f., 49 Anthurium carnosum Croat & R. Baker, 104 Anthurium clidemioides Standl., 10, 104 Anthurium coriaceum (Grah.) G. Don, 45 Anthurium crassinervium (Jacq.) Schott, 11 Anthurium cucullatum K. Koch & Sello, 11 Anthurium ellipticum K. Koch & Bouché, 10 Anthurium ernestii Engl., 42, 51 Anthurium erskinei Mayo, 45, 50 Anthurium flavolineatum Sodiro, c/plate 108C Anthurium friedrichsthalii Schott, 104 Anthurium globosum Croat, 108 Anthurium gracile (Rudge) Schott, 11, 42, 51 Anthurium hookeri Kunth, 11, 45 Anthurium huegelii Schott, 11 Anthurium interruptum Sodiro, 106 Anthurium longissimum Pittier, 105 Anthurium maximum (Desf.) Engl., 11 Anthurium melastomatis Croat, 104, 106 Anthurium oerstedianum Schott, 103, 106 Anthurium peltigerum Sodiro, 104 Anthurium polyschistum R.E. Schult. & Idrobo, 13, 15, 105, 108 Anthurium puberulinervium Croat, 104 Anthurium radicans K. Koch & Haage, 107, 108 Anthurium regale Linden, 108 Anthurium rimbachii Sodiro, 105 Anthurium salviniae Hemsl., 106 Anthurium scandens (Aubl.) Engl., 23, 40 Anthurium scandens (Aubl.) Engl. subsp. scandens, 108 Anthurium scherzerianum Schott, 23, 56 Anthurium smithii Croat, 104 Anthurium triphyllum Brongn. ex Schott, 23 Anthurium trisectum Sodiro, 105 Anthurium vallense Croat, 104 Anthurium wagenerianum K. Koch & Bouché, 12 Anthurium warocqueanum T. Moore, ix, 107 Anthurium watermaliense Bailey, 105 Anthurium wendlingeri G.M. Barroso, 80, 81, 107 Anthurium willdenowii Kunth, 11 Anubiadeae Engl., 18, 26, 33, 49, 68, 180 Anubias Schott, c/plate 117B, 11, 17, 18, 26, 33, 44, 46, 48, 56, 180, 181 Anubias afzelii Schott, 180, 181 Anubias barteri Schott, 16 Anubias barteri Schott var. barteri, 181 Anubias gigantea A. Chev., c/plate 117B, 181 Anubias gracilis A. Chev., 181 Anubias hastifolia Engl., 181 Anubias pynaertii De Wild., 180, 181 Apatemone Schott, 182 Aphyllarum S. Moore, 207 Apiospermum Klotzsch, 286 Apoballis Schott, 182 Aponogetonaceae C. Agardh, 63 Araceae Juss., 80 Araceaeites Fritel, 59 Araceaeites fritelii E.W. Berry, 59 Araceaeites parisiense Fritel, 59 Araceites hungaricus Rásky, 60 Araceophyllum Kräusel, 58 Araceophyllum engleri Kräusel, 58 Araceophyllum striatum Weyland, 58 Araceophyllum tarnocense Rásky, 58 Araceophyllum tobleri Kräusel, 58 Aracispermum Nikitin, 60 Aracispermum canaliculatum Nikitin, 60 Aracispermum hippuriformis Nikitin, 60 Aracispermum johnstrupii (Hartz) Nikitin, 60 Aracispermum jugatum Nikitin, 60

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Aracistrobus dravertii Nikitin, 60 Aracites johnstrupii (Hartz) Nikitin, 60 Arales Lindl., 37, 65 Arctiodracon A. Gray, 94 Areae, 13, 26, 28, 31, 34, 38, 40, 49, 69, 71, 73, 253, Arecaceae C.H. Schultz-Schultzenstein, 36, 38, 60, 61, 62 Areciflorae Dahlgren, Clifford & Yeo, 43 Arecites trabucci Squinab., 60 Aridarum Ridl., c/plate 118D, 14, 33, 48, 56, 192, 193 Aridarum sect. Aridarum, 192 Aridarum sect. Caulescentia M. Hotta, 192 Aridarum annae Bogner, c/plate 118D, 192 Aridarum burttii Bogner & Nicolson, 193 Aridarum caulescens M. Hotta var. caulescens, 193 Aridarum montanum Ridl., 192 Aridarum nicolsonii Bogner, 47, 193 Ariflorae Dahlgren, Clifford & Yeo, 37, 43, 62 Ariopsideae Engl., 71 Ariopsis Nimmo, c/plate 129B, 17, 20, 22, 34, 69, 74, 75, 275, 276 Ariopsis peltata Nimmo, c/plate 129B, 275, 276 Ariopsis protanthera N.E.Br., 276 Arisacontis Schott, 138 Arisaema Mart., c/plate 129A, ix, 5, 6, 7, 11, 13, 22, 24, 25, 26, 27, 29, 31, 34, 36, 44, 46, 47, 48, 50, 52, 54, 55, 69, 270, 271-274 Arisaema sect. Arisaema, 275 Arisaema sect. Clavata Engl., 275 Arisaema sect. Decipientia Engl., 275 Arisaema sect. Dochafa (Schott) Hara, 275 Arisaema sect. Fimbriata Engl., 275 Arisaema sect. Franchetiana Engl., 275 Arisaema sect. Pedatisecta Schott, 275 Arisaema sect. Sinarisaema Nakai, 275 Arisaema sect. Tenuipistillata Engl., 275 Arisaema sect. Tortuosa Engl., 275 Arisaema sect. Trisecta Schott, 275 Arisaema abei S. Seriz., 271, 272 Arisaema atrorubens (Aiton) Blume, 36 Arisaema candidissimum W.W. Sm., 271, 272 Arisaema consanguineum Schott, 271 Arisaema cretacea Lesq., 59 Arisaema dubia Hollick, 59 Arisaema exappendiculatum H. Hara, 270, 274 Arisaema fimbriatum Mast., 274 Arisaema flavum (Forssk.) Schott, 44, 270 Arisaema flavum (Forssk.) Schott subsp. flavum, 274 Arisaema grapsospadix Hayata, 271, 272 Arisaema griffithii Schott, 273 Arisaema hesperia Knowlt., 59 Arisaema heterophyllum Blume, 271 Arisaema jacquemontii Blume, 44 Arisaema japonicum Blume, 36 Arisaema kiushianum Makino, 272 Arisaema lobatum Engl., 44 Arisaema mattewanense Hollick, 59 Arisaema mooneyanum M.G. Gilbert & Mayo, 57 Arisaema nikoense Nakai var. nikoense, 273 Arisaema ovale Nakai var. ovale, c/plate 129A Arisaema ringens (Thunb.) Schott, 273 Arisaema ruwenzoricum N.E.Br., 44 Arisaema scortechinii Hook.f., 270 Arisaema serratum (Thunb.) Schott, 36 Arisaema speciosum (Wall.) Mart. ex Schott, 270 Arisaema taiwanense J. Murata, 273 Arisaema thunbergii Blume, 36 Arisaema tortuosum (Wall.) Schott, 273 Arisaema triphyllum (L.) Torr., 36, 38, 54 Arisaemateae Nakai, 13, 26, 28, 31, 34, 40, 49, 268 Arisaematieae Nakai, 268 Arisareae Dumort., 13, 28, 34, 69, 250 Arisaron Adans., 250 Arisarum Mill., c/plate 126C, 2, 6, 10, 25, 26, 34, 47, 48, 55, 71, 250, 251 Arisarum proboscideum (L.) Savi, 250, 251

Arisarum simorrhinum Durieu ex Duch., c/plate 126C, 251 Arisarum vulgare O. Targ.-Tozz., 36, 37, 38, 41, 42, 250, 251 Aristolochiaceae Juss., 62 Arodendron Werth, 247 Arodes Kuntze, 232 Aroideae, x, 6, 10, 13, 15, 18, 20, 26, 33, 40, 41, 44, 49, 64, 65, 66, 67, 70, 71, 72, 73, 74, 75, 146 Aroides Heist. ex Fabr., 146, 232 Aroides crassispatha Kutorga, 60 Aroides stutterdii Carruth., 60 Aroites tallyanus Kováts, 60 Aron Adans., 253 Aronia J. Mitch., 91 Aronites dubius Heer, 60 Aronium extinctum Ettingsh., 60 Arophyteae Bogner, 13, 26, 34, 69, 71, 74, 243 Arophyton Jum., c/plate 125B, ix, 34, 45, 242, 243 Arophyton buchetii Bogner, 45, 82, 242 Arophyton crassifolium (Buchet) Bogner, c/plate 125B Arophyton tripartitum Jum., 243 Arophyton tripartitum Jum. var. tripartitum, 242 Aropsis Rojas Acosta, 166 Arosma Raf., 169 Arum L., c/plate 127A, 2, 6, 10, 17, 24, 26, 27, 28, 29, 34, 44, 46, 47, 48, 49, 50, 54, 55, 253, 255 Arum subgen. Arum, 254 Arum subgen. Gymnomesium (Schott) Engl., 254 Arum alpinum Schott & Kotschy, 255 Arum arisarum L., 250 Arum auritum L., 214 Arum bicolor Aiton, 207 Arum colocasia L., 280 Arum concinnatum Schott, 255 Arum crenatum Wight, 260 Arum creticum Boiss. & Heldreich, 255 Arum cucullatum Lour., 283 Arum dioscoridis Sm., 46 Arum guttatum Wall., 263 Arum integrifolium Link, 223 Arum italicum Mill., 10, 37, 38, 41, 46, 254 Arum italicum Mill. subsp. italicum, 54, 255 Arum korolkowii Regel, 253, 254 Arum maculatum L., c/plate 127A, 10, 38, 40, 41, 46, 50, 54, 253, 255, 311 Arum muscivorum L.f., 257 Arum nigrum Schott, 254 Arum orientale M.Bieb. subsp. orientale, 255 Arum pictum L.f., 87, 253, 254, 255 Arum pythonium Mart., 199 Arum rupicola Boiss. var. rupicola, 255 Arum sagittaefolium L., 209 Arum seguine Jacq., 153 Arum speciosum Wall., 270 Arum spiculatum Blume, 254 Arum spirale Retz., 197 Arum subulatum Desf., 268 Arum tenuifolium L., 266 Arum trilobatum L., 260 Arum viviparum Roxb., 280 Asarum L., 24, 42 Asilidae, 48 Aspidistreae Hutch., 61 Asteraceae Dumort., 59 Asterostigma Fisch. & C.A.Mey., c/plate 115B, 14, 18, 33, 160, 161 Asterostigma sect. Asterostigma, 160 Asterostigma sect. Rhopalostigma (Schott) Engl., 160 Asterostigma cryptostylum Bogner, 161 Asterostigma integrifolium Madison, 161 Asterostigma langsdorffianum Fisch. & C.A.Mey., 160 Asterostigma riedelianum (Schott) Kuntze, c/plate 115B, 45, 161 Asterostigma vermitoxicum Griseb., 163 Asterostigmateae Schott, 74 Astragalus L., 40

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Ateles paniscus L., 51 Atherurus Blume, 268 Atimeta Schott, 125

B Balmisa Lag., 250 Batis Blanco, 98 Baursea Post & Kuntze, 169 Biarum Schott, c/plate 128C, 6,7, 31, 34, 46, 51, 55, 266, 267 Biarum subgen. Biarum, 268 Biarum subgen. Cyllenium (Schott) Engl., 268 Biarum subgen. Ischarum (Blume) Engl., 268 Biarum aleppicum J. Thiéb., 87, 266 Biarum davisii Turrill, 46 Biarum davisii Turrill subsp. marmarisense P.C. Boyce, 267 Biarum ditschianum Bogner & P.C. Boyce, c/plate 128C, 46, 253, 267 Biarum pyrami (Schott) Engl. var. pyrami, 267 Biarum spruneri Boiss., 267 Biarum straussii Engl., 267 Biarum tenuifolium (L.) Schott, 266 Biarum tenuifolium (L.) Schott subsp. tenuifolium, 267 Bognera Mayo & Nicolson, c/plate 114C, ix, 16, 18, 33, 68, 155, 156 Bognera recondita (Madison) Mayo & Nicolson, c/plate 114C, 155, 156 Bognereae Mayo & Nicolson, 117 Brachyspatha Schott, 235 Bromeliaceae Juss., 41 Bromeliiflorae Dahlgren, Clifford & Yeo, 43 Bucephalandra Schott, c/plate 118B, 14, 33, 49, 56, 189, 190 Bucephalandra catherineae P.C. Boyce, Bogner & Mayo, 190 Bucephalandra gigantea Bogner, 190 Bucephalandra motleyana Schott, c/plate 118B, 189, 190 Bucerotidae, 50

C Caladieae Schott, 8, 13, 15, 18, 20, 21, 26, 33, 44, 49, 53, 71, 204 Caladiopsis Engl., 211 Caladiosoma E.W. Berry, 58 Caladiosoma miocenica E.W. Berry, 58 Caladium Vent., c/plate 120C, 7, 11, 13, 17, 21, 24, 33, 46, 48, 51, 54, 55, 56, 58, 69, 207, 208 Caladium Alliance, 71 Caladium aculeatum G.Mey., 230 Caladium aristeguietae G.S. Bunting, 208 Caladium bicolor (Aiton) Vent., 56, 207, 208 Caladium coerulescens G.S. Bunting, 208 Caladium lindenii (André) Madison, 40, 207, 208 Caladium nymphaeifolium Vent., 36 Caladium paradoxum Bogner & Mayo, 90, 207 Caladium petiolatum Hook., 218 Caladium ternatum Madison, 207 Caladium tuberosum (S. Moore) Bogner & Mayo, c/plate 120C Caladium zamiaefolium Lodd., 146 Calamariaceae, 64 Calamites, 64 Calla L., c/plate 113B, 10, 11, 13, 14, 18, 19, 20, 22, 29, 30, 33, 44, 52, 55, 67, 74, 146, 147 Calla aethiopica L., 232 Calla calyptrata Roxb., 182 Calla palustris L., c/plate 113B, 13, 36, 40, 59, 146, 147 Callaria Raf., 146 Calloideae Endl., 11, 15, 20, 33, 67, 68, 70, 73, 74, 146 Callopsideae Engl., 34, 69, 74, 232 Callopsis Engl., c/plate 124B, 16, 34, 232, 234 Callopsis volkensii Engl., c/plate 124B, 232, 234 Calyptrocoryne Schott, 260 Camponotus femoratus, 42 Campylospermum hordwellensis M. Chandler, 60

334

THE GENERA OF ARACEAE

Candarum Rchb. ex Schott & Endl., 235 Canna L., 36 Carex eximia Göpp. & Menge, 59 Caricoidea M. Chandler, 60 Caricoidea jugata (Nikitin) Mai, 60 Carlephyton Jum., c/plate 125C, ix, 34, 45, 243, 244 Carlephyton sect. Carlephyton, 245 Carlephyton sect. Pseudocolletogyne Bogner, 82, 245 Carlephyton diegoense Bogner, 244 Carlephyton glaucophyllum Bogner, c/plate 125C, 17, 243 Carlephyton madagascariense Jum., 243, 244 Castanospermum australe A.Cunn., 40 Centropogonidae, 48 Ceratitis, 42 Cercestis Schott, c/plate 123C, 6, 7, 11, 12, 13, 14, 15, 16, 21, 34, 45, 48, 54, 228, 229 Cercestis afzelii Schott, 228, 229 Cercestis camerunensis (Ntépé) Bogner, 228 Cercestis dinklagei Engl., 229 Cercestis ivorensis A. Chev., c/plate 123C Cercestis kamerunianus (Engl.) N.E.Br., 229 Cercestis mirabilis (N.E.Br.) Bogner, 45, 81, 228, 229 Cercestis stigmaticus N.E.Br., 229 Cercestis taiensis Bogner & Knecht, 228 Cetoniidae, 48 Chamaecladon Miq., 177 Chamaedorea Willd., 15 Chersydrium Schott, 130 Chiroxiphia linearis, 50 Chlorispingus ophthalmicus, 50 Chloropidae, 48 Chlorospatha Engl., c/plate 121B, 13, 33, 35, 44, 48, 57, 69, 71, 211, 213 Chlorospatha atropurpurea (Madison) Madison, 213 Chlorospatha corrugata Bogner & Madison, 213 Chlorospatha croatiana Grayum subsp. enneaphylla Grayum, 213 Chlorospatha kolbii Engl., 211, 213, 214 Chlorospatha longipoda (K. Krause) Madison, c/plate 121B, 13, 35, 213, 214 Chlorospatha mirabilis (Mast.) Madison, 214 Chondrophyllum nordenskiöldii Heer, 58 Choridae, 48 Colletogyne Buchet, c/plate 125D, ix, 34, 45, 47, 245, 246 Colletogyne perrieri Buchet, c/plate 125D, 245, 246 Colobogynium Schott, 182 Colocasia Link, 232 Colocasia Schott, c/plate 130A, 6, 10, 11, 17, 21, 22, 24, 29, 34, 37, 38, 44, 47, 48, 51, 53, 55, 69, 280, 282 Colocasia sect. Alocasia Schott, 283 Colocasia sect. Caulescentes Engl., 283 Colocasia sect. Colocasia, 283 Colocasia sect. Tuberosae Engl., 283 Colocasia antiquorum Schott, 41, 280 Colocasia esculenta (L.) Schott, c/plate 130A, 22, 23, 36, 37, 41, 42, 50, 52, 53, 54, 55, 56, 57, 89, 90, 282, 283, 311 Colocasia fallax Schott, 282 Colocasia gigantea (Blume) Hook.f., 45, 50, 282, 283 Colocasieae Engl., 7, 8, 13, 15, 18, 20, 21, 26, 28, 34, 40, 44, 49, 53, 59, 71, 275 Colocasioideae Engl., 20, 38, 70, 71, 73, 74 Colocasioideaepites Biswas, 59 Commelinales Lindl., 65 Commelinidae Takht., 38 Commeliniflorae Dahlgren, Clifford & Yeo, 37, 43 Coniferae Juss., 58, 60 Conophallus Schott, 235 Convallaria L., 43 Corynophallus Schott, 235 Cryptocorynaceae J. Agardh, 5 Cryptocoryne Fisch. ex Wydler, c/plate 119B, 5, 6, 10, 14, 15, 20, 22, 27, 28, 31, 33, 46, 47, 48, 49, 51, 56, 57, 69, 72, 197, 198 Cryptocoryne affinis Hook.f., 46, 198

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Cryptocoryne aponogetifolia Merr., 46 Cryptocoryne ciliata (Roxb.) Fisch. ex Schott, 28, 29, 31, 46, 82, 197, 198, 225 Cryptocoryne cognata Schott, 29 Cryptocoryne consobrina Schott, 46, 80, 197 Cryptocoryne cordata Griff., 31 Cryptocoryne ferruginea Engl., 46 Cryptocoryne lingua Becc. ex Engl., 46 Cryptocoryne longicauda Engl., c/plate 119B, 198 Cryptocoryne nevillii Trimen ex Hook.f., 46, 197 Cryptocoryne pontederiifolia Schott, 46 Cryptocoryne purpurea Ridley, 46 Cryptocoryne retrospiralis (Roxb.) Fisch. ex Wydler, 197 Cryptocoryne spiralis (Retz.) Fisch. ex Wydler, 46, 57, 88, 197, 198 Cryptocoryneae Blume, 18, 20, 25, 31, 33, 44, 194 Culcasia P.Beauv., c/plate 123B, 11, 13, 17, 21, 22, 23, 34, 44, 48, 73, 225, 227 Culcasia angolensis Welw. ex Schott, 227 Culcasia liberica N.E.Br., 227, 228 Culcasia longevaginata Engl., 227 Culcasia orientalis Mayo, 227 Culcasia panduriformis Engl. & K. Krause, 227 Culcasia parviflora N.E.Br., 227 Culcasia rotundifolia Bogner, 227 Culcasia saxatilis A. Chev. c/plate 123B, 15, 38, 227 Culcasia scandens P.Beauv., 24, 225 Culcasia seretii De Wild., 227 Culcasia striolata Engl., 227 Culcasieae Engl., 13, 18, 26, 34, 44, 69, 74, 225 Curculionidae, 48 Curmeria André, 117 Cycadaceae Pers., 60 Cyclanthaceae Poit. ex A. Rich., 16, 60, 61 Cyclanthodendron, 60 Cyclanthodendron sahnii (K.P. Rode) Sahni & Surange, 60 Cyllenium Schott, 266 Cyperaceae Juss., 60 Cyrtocladon Griff., 177 Cyrtospadix K. Koch, 209 Cyrtosperma Griff., c/plate 112A, ix, 7, 17, 18, 32, 44, 48, 50, 56, 73, 138, 139 Cyrtosperma beccarianum A. Hay, 139 Cyrtosperma carrii A. Hay, c/plate 112A Cyrtosperma cuspidispathum Alderw., 138, 139, 311 Cyrtosperma lasioides Griff., 138 Cyrtosperma macrotum Engl., 139 Cyrtosperma merkusii (Hassk.) Schott, 52, 53, 55, 138, 139 Cyrtosperma senegalense (Schott) Engl., 38, 138 Cyrtospermites hordwellensis (M. Chandler) Bogner, 60

D Dacus, 42 Dacus musae, 42 Dasycladaceae, 60 Denhamia Schott, 225 Dermestidae, 48 Desmesia Raf., 260 Diandriella Engl., 177 Diclines Schott, 67, 72 Dieffenbachia Schott, c/plate 114B, 11, 13, 14, 15, 16, 17, 18, 19, 21, 23, 24, 26, 28, 33, 40, 41, 44, 46, 47, 48, 49, 50, 51, 54, 55, 56, 68, 70, 153, 154 Dieffenbachia Alliance, 70-71 Dieffenbachia cordata Engl., 38 Dieffenbachia elegans A.M.E. Jonker & Jonker, 154 Dieffenbachia humilis Poepp., 153 Dieffenbachia maculata (Lodd.) G. Don, c/plate 114B, ix, 36, 40, 55 Dieffenbachia paludicola N.E.Br. ex Gleason, 154 Dieffenbachia parlatorei Linden & André, 153 Dieffenbachia picta (Lodd.) Schott, 36, 40 Dieffenbachia seguine (L.) Schott, 55, 153, 154

Dieffenbachia seguine (L.) Schott var. viridis (Engl.) Engl., 154 Dieffenbachia seguine (L.) Schott ‘Reginae’, 154 Dieffenbachieae Engl., 18, 25, 26, 33, 40, 44, 71, 153 Dioscoreaceae R.Br., 36, 61 Dioscoreales Hook.f., 43, 61 Dochafa Schott, 270 Dracontioides Engl., c/plate 111A, 6, 7, 32, 56, 130, 132 Dracontioides desciscens (Schott) Engl., c/plate 111A, 130, 132 Dracontium J. Hill, 257 Dracontium L., c/plate 110D, 2, 6, 7, 16, 18, 26, 32, 45, 49, 54, 56, 73, 74, 130, 131 Dracontium sect. Dracontium, 130 Dracontium sect. Echidnium (Schott) G. Zhu, 130 Dracontium sect. Godwinia (Seem.) Engl., 130 Dracontium sect. Urospathopsis Engl., 130 Dracontium asperum K. Koch, 131 Dracontium camtschatcense L., 94 Dracontium changuango G.S. Bunting, c/plate 110D, 131 Dracontium cordatum Houtt., 177 Dracontium foetidum L., 96 Dracontium gigas (Seem.) Engl., 131 Dracontium lanceaefolium Jacq., 109 Dracontium margaretae Bogner, 45, 130, 131 Dracontium pertusum L., 123 Dracontium polyphyllum L., 130 Dracontium prancei G.H. Zhu, 45, 131 Dracontium soconuscum Matuda, 131 Dracontium spruceanum (Schott) G.H. Zhu, 131 Dracunculus Mill., c/plate 127C, 2, 11, 24, 26, 34, 48, 55, 257, 258 Dracunculus canariensis Kunth, 38, 257, 258 Dracunculus crinitus Schott, 257 Dracunculus vulgaris Schott, c/plate 127C, 41, 257, 258 Drosophilidae, 48

E Echidnium Schott, 130 Efilamentatae Schott, 72 Eichhornia Kunth, 10 Elachiptera formosa, 42 Elaeocarpaceae Juss. ex DC., 52 Elopium Schott, 169 Eminium (Blume) Schott, c/plate 127B, 5, 7, 34, 46, 55, 254, 256 Eminium albertii (Regel) Engl., 256 Eminium koenenianum Lobin & P.C. Boyce, 256, 257 Eminium lehmannii (Regel) Kuntze, c/plate 127B, 254 Eminium regelii Vved., 254 Eminium spiculatum (Blume) Kuntze, 36, 37, 38, 41, 254 Eminium spiculatum (Blume) Kuntze subsp. negevense Koach & Feinbrun, 46 Eminium spiculatum (Blume) Kuntze subsp. spiculatum, 256 Endera Regel, 158 Ensolenanthe Schott, 283 Ephedripites vanegensis Hammen & Garcia de Mutis, 60 Ephydridae, 48 Epipremnites Gregor & Bogner, 59 Epipremnites ornatus (Reid & Chandler) Gregor & Bogner, 59 Epipremnopsis Engl., 20 Epipremnum Schott, c/plate 109C, 9, 12, 16, 20, 21, 24, 32, 55, 56, 59, 120, 121 Epipremnum amplissimum (Schott) Engl., 121 Epipremnum aureum (Linden & André) G.S. Bunting, ix Epipremnum falcifolium Engl., c/plate 109C Epipremnum mirabile Schott, 121 Epipremnum nobile (Schott) Engl., 120 Epipremnum pinnatum (L.) Engl., 12, 21, 120, 121 Epipremnum pinnatum (L.) Engl. ‘Aureum’, ix, 52, 56 Epipremnum spadiciflorum (Renner) Tiffney & Wing, 59 Equisetum L., 14 Erithacus rubecula L., 50 Euphonia anneae, 50 Euphorbiaceae Juss., 21, 40 Eutereia Raf., 130

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F Fagus sylvatica L., 56 Felipponea Broth., 155 Felipponia Hicken, 155 Felipponiella Hicken, 155 Filarum Nicolson, ix, 33, 69, 204, 205 Filarum manserichense Nicolson, 204, 205 Flagellarisaema Nakai, 270 Fructus polyspermus Engelh., 59 Furtadoa M. Hotta, c/plate 116D, ix, 11, 21, 33, 175, 176 Furtadoa mixta (Ridl.) M. Hotta, 176, 177 Furtadoa sumatrensis M. Hotta, c/plate 116D, 175, 176, 177

G Gamochlamys Baker, 166 Gamogyne N.E.Br., 184 Gearum N.E.Br., 33, 46, 163, 165 Gearum brasiliense N.E.Br., 163, 165 Godwinia Seem., 130 Gonatanthus Klotzsch, 17, 69, 280 Gonatanthus sarmentosus Klotzsch, 280 Gonatopus Hook.f. ex Engl., c/plate 113D, 7, 11, 13, 17, 18, 28, 33, 35, 56, 149, 150 Gonatopus angustus N.E.Br., 150 Gonatopus boivinii (Decne.) Engl., 18, 19, 80, 149, 150,151 Gonatopus clavatus Mayo, 150 Gonatopus marattioides (Peter) Bogner, c/plate 113D, 150 Gonatopus petiolulatus (Peter) Bogner, 149, 150, 151 Gonioscypha Baker, 61 Goniurus C. Presl, 98 Gorgonidium Schott, c/plate 115C, 33, 44, 160, 162 Gorgonidium mirabile Schott, 160, 162 Gorgonidium vargasii Bogner & Nicolson, 160, 162, 311 Gorgonidium vermicidum (Speg.) Bogner & Nicolson, c/plate 115C, 162 Gramineae Juss., 42 Gramineae subfam. Bambusoideae Asch. & Graeb., 41 Gymnogoneae Schott, 72 Gymnomesium Schott, 253 Gymnostachydoideae Bogner & Nicolson, 32, 44, 66, 67, 68, 70, 91 Gymnostachys R.Br., c/plate 107A, 7, 8, 15, 18, 25, 29, 32, 52, 67, 72, 73, 74, 91, 92 Gymnostachys anceps R.Br., c/plate 107A, 38, 54, 91, 92

H Haemodoraceae R.Br., 38 Hansalia Schott, 235 Hapale Schott, 216 Hapaline Schott, c/plate 121D, 33, 69, 216, 217 Hapaline appendiculata Ridl., 89, 216 Hapaline benthamiana Schott, 216, 217 Hapaline celatrix P.C. Boyce, c/plate 121D, 217 Hapaline colaniae Gagnep., 217 Hapaline ellipticifolia C.Y. Wu & H. Li, 217 Helicodiceros Schott ex K. Koch, c/plate 127D, 7, 26, 34, 48, 49, 55, 257, 259 Helicodiceros crinitus K. Koch, 257 Helicodiceros muscivorus (L.f.) Engl., c/plate 127D, ix, 257, 259 Heliconia L., 52 Heliconiaceae (A.Rich.) Nakai, 52 Helicophyllum Schott, 254 Helobiae Engl., 36 Hemicarpurus Nees, 268 Hermaphroditanthae Schott, 72 Heteroaridarum M. Hotta, ix, 26, 33, 194, 194 Heteroaridarum borneense M. Hotta, 194, 194 Heteroarisaema Nakai, 270 Heterolobium Peter, 149 Heteropsideae Engl., 32, 67, 68, 113 Heteropsis Kunth, 7, 10, 13, 14, 15, 19, 20, 32, 54, 67, 74, 75, 113, 115

336

THE GENERA OF ARACEAE

Heteropsis cf. jenmanii Oliv., 115 Heteropsis melinonii (Engl.) A.M.E. Jonker & Jonker, 116 Heteropsis oblongifolia Kunth, 115 Heteropsis salicifolia Kunth, 113 Heteropsis spruceana Schott, 54, 115 Heterostalis (Schott) Schott, 260 Heterothrips arisaemae, 48 Holochlamys Engl., 26, 32, 46, 74, 110, 112 Holochlamys beccarii (Engl.) Engl., 110, 112 Homaïd Adans., 266 Homaida Raf., 266 Homalomena Schott, c/plate 117A, 7, 11, 13, 14, 17, 18, 21, 24, 27, 31, 33, 42, 44, 46, 47, 48, 52, 55, 59, 177, 178-179 Homalomena sect. Chamaecladon (Miq.) Engl., 180 Homalomena sect. Curmeria (Linden & André) Engl., 16, 85, 180 Homalomena sect. Cyrtocladon Furtado, 180 Homalomena sect. Geniculatae M. Hotta, 180 Homalomena sect. Homalomena, 180 Homalomena aromatica (Roxb.) Schott, 42 Homalomena consobrina (Schott) Engl., 178 Homalomena cordata Schott, 177 Homalomena geniculata M. Hotta, c/plate 117A, 177 Homalomena havilandii Ridl., 178 Homalomena hostifolia Engl., 179 Homalomena humilis (Jack) Hook.f., 179 Homalomena lindenii (Rodigas) Ridl., 177 Homalomena monandra M. Hotta, 177 Homalomena occulta (Lour.) Schott, 42 Homalomena pendula (Hassk.) Bakh.f., 50 Homalomena picturata Regel, 178 Homalomena propinqua Schott, 178 Homalomena rubescens (Roxb.) Kunth, 178 Homalomena rubra Hassk., 179 Homalomena sagittifolia Jungh. ex Schott, 178 Homalomena speariae Bogner & Moffler, 31, 179 Homalomena vagans P.C. Boyce, 179 Homalomena versteegii Engl., 35 Homalomeneae (Schott) M. Hotta, 18, 26, 33, 44, 49, 68, 175 Homalonema Endl., 177 Hottarum Bogner & Nicolson, c/plate 118A, ix, 14, 33, 56, 187, 188 Hottarum kinabaluense Bogner, 188 Hottarum lucens Bogner, c/plate 118A, 182, 187, 188 Hottarum sarikeense Bogner & M. Hotta, 188 Hottarum truncatum (M. Hotta) Bogner & Nicolson, 187 Houttinia Neck., 232 Humbertina Buchet, 243 Hydnostachyon Liebm., 109 Hydrocharitaceae Juss., 58, 60 Hydrosme Schott, 235 Hylocichla mustelina Gmelin, 50

I Ictodes Bigelow, 96 Ischarum (Blume) Rchb., 266 Jaimenostia Guinea & Gomez-Mor., 263

J Jasarum G.S. Bunting, c/plate 120D, ix, 7, 33, 46, 56, 57, 69, 209, 210 Jasarum steyermarkii G.S. Bunting, c/plate 120D, 46, 209, 210 Jugella Mtchedlishvili & Shakmundes, 59 Jugella sibirica Mtchedlishvili & Shakmundes, 59 Juncaceae Juss., 29 Juncaginaceae Rich., 63

K Keratosperma Cevallos-Ferriz & Stockey, 59 Keratosperma allenbyense Cevallos-Ferriz & Stockey, 59 Kodda-Pail Adans., 286 Kunda Raf., 235

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L Lagenandra Dalzell, c/plate 119A, 5, 14, 17, 20, 22, 28, 33, 49, 51, 56, 62, 195, 196 Lagenandra bogneri De Wit, 195 Lagenandra dewitii Crusio & A. de Graaf, 196 Lagenandra gomezii (Schott) Bogner & N. Jacobsen, 88, 195, 197 Lagenandra koenigii (Schott) Thwaites, 196 Lagenandra nairii Ramam. & Rajan, c/plate 119A, 88, 195, 196, 197 Lagenandra toxicaria Dalzell, 195, 196 Laportea moroides Wedd., 41 Lasia Lour., c/plate 112D, 17, 18, 22, 32, 44, 46, 73, 141, 143 Lasia aculeata Lour., 141 Lasia concinna Alderw., 141, 144 Lasia spinosa (L.) Thwaites, c/plate 112D, 40, 55, 141, 143, 144 Lasieae Engl., 67, 74, 75 Lasimorpha Schott, c/plate 112B, ix, 6, 17, 32, 138, 140 Lasimorpha senegalensis Schott, c/plate 112B, 38, 138, 140 Lasioideae Engl., 2, 5, 7, 10, 15, 18, 20, 26, 28, 32, 38, 40, 44, 47, 59, 66, 67, 68, 70, 73, 74, 75, 130 Lasiomorpha Engl., 138 Lasius Hassk., 141 Lazarum A. Hay, ix, 34, 52, 265, 266 Lazarum mirabile A. Hay, 265, 266 Lecontia W. Cooper ex Torr., 247 Leguminosae Juss., 20, 40 Lemna L., 10, 29, 37, 58, 63 Lemnaceae Gray, x, 22, 23, 29, 36, 58, 60, 61, 62, 63, 64, 65, 72, 73 Lemnoideae Engl., 72 Lemnospermum pistiforme Nikitin, 58 Leptopetion Schott, 266 Leucocasia Schott, 280 Liliaceae Juss., 61 Liliidae Takht., 38 Liliiflorae Dahlgren, Clifford & Yeo, 36, 37, 43, 61, 63, 64, 66 Lilloa Speg., 163 Limnobiophyllum Krassilov, 58, 63 Limnobiophyllum expansum (Heer) Kvacek, 58 Limnobiophyllum scutatum (Dawson) Krassilov, 58 Limnobium Rich., 58 Limnonesis Klotzsch, 286 Lysichiton Schott, c/plate 107C, 10, 28, 32, 44, 48, 52, 55, 73, 74, 94, 95 Lysichiton americanus Hultén & H. St. John, 95 Lysichiton camtschatcensis (L.) Schott, c/plate 107C, 36, 41, 94, 95 Lysichiton nevadensis MacGinitie, 59 Lysichiton washingtonensis E.W. Berry, 59 Lysichitum Schott, 96 Lysistigma Schott, 158

M Magnolia L., 42 Magnolianae, 42 Magnoliidae Takht., 41 Maguirea A.D. Hawkes, 153 Mangonia Schott, c/plate 114D, 33, 46, 155, 157 Mangonia tweedieana Schott, c/plate 114D, 155, 157, 158 Mangonia uruguaya (Hicken) Bogner, 157 Manucodia atra Lesson, 50 Maranta L., 36 Massovia Benth. & Hook.f., 110 Massowia K. Koch, 109 Megotigea Raf., 257 Melanthiaceae Batsch, 29, 61, 66 Meliphaga lewinii, 50 Microcasia Becc., 189 Microcasia sect. Truncatae M. Hotta, 187 Microcasia truncata M. Hotta, 187 Microculcas Peter, 149 Monoclines Schott, 72

Monocolpopollenites, 60 Monocolpopollenites tranquillus (R. Potonié) Thomson & Pflug, 60 Monstera Adans., c/plate 110A, 7, 8, 10, 11, 12, 14, 16, 18, 20, 21, 23, 32, 45, 47, 48, 52, 55, 56, 123, 124 Monstera sect. Echinospadix Madison, 125 Monstera sect. Marcgraviopsis Madison, 125 Monstera sect. Monstera, 125 Monstera sect. Tornelia (Guttierez) Madison, 125 Monstera adansonii Schott, c/plate 110A, 123 Monstera adansonii Schott var. laniata (Schott) Madison, 124 Monstera deliciosa Liebm., ix, 6, 10, 11, 12, 17, 19, 22, 23, 24, 36, 38, 41, 54, 55 Monstera dubia (H.B.K.) Engl. & K. Krause, 123 Monstera lechleriana Schott, 124 Monstera obliqua Miq., 123 Monstera oreophila Madison, 124 Monstera punctulata (Schott) Engl., 20 Monstera subpinnata (Schott) Engl., 124 Monstera tuberculata Lundell var. tuberculata, 124 Monstereae Engl., 12, 13, 16, 20, 26, 28, 32, 47, 58, 59, 67, 68, 75, 116 Monsteroideae Engl., 7, 10, 11, 12, 13, 14, 18, 19, 20, 23, 26, 32, 38, 44, 51, 67, 70, 74, 75, 109 Monsteroideaepites eospathiphyllum Biswas, 59 Montrichardia Crueg., c/plate 123D, 11, 13, 14, 15, 16, 17, 18, 34, 37, 46, 51, 230, 231 Montrichardia aculeata (G.Mey.) Schott, 230 Montrichardia arborescens (L.) Schott, 37, 38, 55, 231 Montrichardia linifera (Arruda Camara) Schott, c/plate 123D, 54, 55, 231 Montrichardieae Engl., 26, 34, 69, 74, 230 Moraceae Link, 40 Muricauda Small, 270 Musa L., 36 Mycetophilidae, 48 Myrica L., 60 Myricaceae Blume, 60

N Nebrownia Kuntze, 182 Nephthytideae Engl., 13, 26, 33, 40, 44, 69, 74, 216 Nephthytis Schott, c/plate 122A, 17, 28, 30, 33, 40, 44, 47, 48, 51, 216, 219 Nephthytis afzelii Schott, 216, 219 Nephthytis afzelii Schott var. graboensis Bogner, c/plate 122A Nephthytis bintuluensis A. Hay, Bogner & P.C. Boyce, 218 Nephthytis constricta N.E.Br., 219 Nephthytis hallaei (Bogner) Bogner, 218, 219 Nephthytis mayombensis Namur & Bogner, 218 Nephthytis poissonii (Engl.) N.E.Br., 219 Nephthytis swainei Bogner, 219 Neurochaetidae, 48 Nitidulidae, 48 Nypa Steck, 59 Nypa fruticans Wurmb, 225

O Oligogynium Engl., 216 Ophione Schott, 73, 130 Orchidaceae Juss., 11 Orontieae R.Br. ex Dumort., 67, 73, 74 Orontioideae Mayo, Bogner & P.C. Boyce, 6, 10, 15, 20, 25, 32, 44, 66, 67, 68, 70, 91, 346 Orontium L., c/plate 107B, 10, 14, 17, 18, 20, 23, 25, 28, 32, 44, 55, 72, 73, 74, 91, 93 Orontium aquaticum L., c/plate 107B, 38, 54, 91, 93 Orontium fossile Cockerell, 59 Orthotropooae Schott, 72 Otosma Raf., 232 Oxytropis DC., 40

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P Pachyzeugmaticae Schott, 72 Palmoxylon sahnii Rode, 60 Pandanaceae R.Br., 16, 59, 60, 61 Pandanus Parkinson, 45 Panzhuyuia Z.Y. Zhu, 283 Paradoxurus hermaphroditus Pallas, 50 Pauella Ramam. & Sebastine, 260 Pedicellarum M. Hotta, c/plate 108B, ix, 6, 25, 26, 32, 74, 100, 101 Pedicellarum paiei M. Hotta, c/plate 108B, 100, 101 Peltandra Raf., c/plate 126A, 14, 17, 24, 34, 44, 48, 52, 55, 58, 59, 247, 248 Peltandra primaeva Hickey, 58 Peltandra sagittifolia (Michx.) Morong, 247 Peltandra undulata Raf., 247 Peltandra virginica (L.) Raf., c/plate 126A, 30, 36, 38, 42, 82, 247, 248 Peltandreae Engl., 14, 18, 26, 34, 58, 69, 71, 245 Peltandripites davisii Wodehouse, 59 Peltandripites dubius Sah & Dutta, 59 Peristatogoneae Schott, 72 Petermanniaceae Hutch., 61 Phaeochrous camerunensis, 35 Pharomachrus mocinno De La Llave, 50 Philodendreae Schott, 13, 18, 26, 33, 49, 68, 79, 169 Philodendroideae Engl., 18, 38, 59, 66, 70, 73, 74 Philodendron Schott, c/plate 116C, ix, 6, 7, 8, 9, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 24, 26, 27, 28, 29, 33, 35, 37, 39, 40, 44, 45, 46, 47, 48, 49, 50, 51, 52, 54, 55, 56, 58, 169, 170-174 Philodendron Alliance, 71 Philodendron sect. Polyspermium Engl., 17 Philodendron subgen. Meconostigma (Schott) Engl., 21, 28, 51, 169, 175 Philodendron subgen. Philodendron, 17, 21, 169, 175 Philodendron subgen. Pteromischum (Schott) Mayo, 14, 15, 169, 175 Philodendron andreanum Devansaye, 41 Philodendron angustisectum Engl., 40, 171 Philodendron anisotomum Schott, 171 Philodendron aromaticum Croat & Grayum, 170 Philodendron bipennifolium Schott, 40, 171, 174 Philodendron bipinnatifidum Schott, 6, 10, 21, 41, 50, 54, 55, 171, 172, 173, 174 Philodendron blanchetianum Schott, 170, 174 Philodendron calatheifolium G.S. Bunting, 170 Philodendron callosum K. Krause, 170 Philodendron corcovadense Kunth, 40 Philodendron craspedodromum R.E. Schult., 311 Philodendron crassinervium Lindl., 81, 170 Philodendron erubescens K. Koch & Augustin, 40, 41, 55 Philodendron eximium Schott, 174 Philodendron fenzlii Engl., 40 Philodendron fibrillosum Poepp., 170 Philodendron fragrantissimum (Hook.) G. Don, 45 Philodendron frits-wentii G.S. Bunting, 170 Philodendron goeldii G.M. Barroso, 7, 22, 51, 169,171, 174, 175 Philodendron grandifolium (Jacq.) Schott, 169 Philodendron grazielae G.S. Bunting, 170 Philodendron hederaceum (Jacq.) Schott, 15 Philodendron heterophyllum Poepp., 170 Philodendron inaequilaterum Liebm., 174 Philodendron insigne Schott, 45, 172 Philodendron leal-costae Mayo & G.M. Barroso, 169, 173, 175 Philodendron limnestis Dilcher & Daghlian, 58 Philodendron linnaei Kunth, 45 Philodendron martianum Engl., 41, 80 Philodendron megalophyllum Schott, 22, 42, 51 Philodendron melanochrysum Linden & André, 9, 41 Philodendron melinonii Brongn. ex Regel, 172, 174 Philodendron myrmecophilum Engl., 22, 51

338

THE GENERA OF ARACEAE

Philodendron ornatum Schott, 38 Philodendron radiatum Schott, 40 Philodendron rigidifolium K. Krause, 173 Philodendron rugosum Bogner & G.S. Bunting, c/plate 116C Philodendron sagittifolium Liebm., 40 Philodendron scandens K. Koch & Sello, ix, 9, 41, 44, 55, 172 Philodendron scandens K. Koch & Sello subsp. oxycardium (Schott) Bunting, 40 Philodendron selloum K. Koch, 15, 28, 46 Philodendron squamiferum Poepp., 40, 80 Philodendron surinamense (Miq.) Engl., 169 Philodendron tripartitum (Jacq.) Schott, 41 Philodendron tuxtlanum G.S. Bunting, 40 Philodendron uliginosum Mayo Philodendron verrucosum Mathieu ex Schott, 172, 173 Philodendron sp. cf. verrucosum Mathieu ex Schott, 170 Philodendrum Schott, 169 Philonotion Schott, 182 Philydraceae Link, 38 Phoridae, 48 Phyllostachys edulis (Carr.) H. de Lehaie, 41 Phyllotaenium André, 207 Phymatarum M. Hotta, c/plate 118C, ix, 14, 33, 189, 191 Phymatarum borneense M. Hotta, c/plate 118C, 189, 191 Pinellia Ten., c/plate 128D, 6, 18, 25, 29, 34, 37, 44, 47, 49, 55, 69, 268, 269 Pinellia cordata N.E.Br., c/plate 128D, 269 Pinellia pedatisecta Schott, 269 Pinellia ternata (Thunb.) Makino, 18, 36, 41, 55, 269, 270 Pinellia tripartita Schott, 36, 40, 268, 269, 270 Pinellia tuberifera Ten., 268 Pinus L., 60 Piper sumatranum C.DC. var. andamanicum, 42 Piperaceae C. Agardh, 62 Piptospatha N.E.Br., c/plate 117D, 14, 17, 33, 48, 56, 184, 186 Piptospatha sect. Gamogyne (N.E.Br.) M. Hotta, 184 Piptospatha sect. Piptospatha, 184 Piptospatha burbidgei (N.E.Br.) M. Hotta, c/plate 117D, 186 Piptospatha elongata (Engl.) N.E.Br., 186 Piptospatha insignis N.E.Br., 184 Piptospatha ridleyi N.E.Br., 186 Pistia L., c/plate 130C, 7, 10, 14, 16, 17, 23, 24, 25, 28, 30, 34, 44, 47, 48, 51, 52, 58, 59, 63, 64, 65, 71, 74, 75, 286, 287 Pistia claibornensis E.W. Berry, 58 Pistia corrugata Lesq., 58 Pistia mazelii Saporta & Marion, 58 Pistia nordenskiöldii (Heer) E.W. Berry, 58 Pistia sibirica Dorof., 59 Pistia stratiotes L., c/plate 130C, 38, 42, 46, 58, 286, 287 Pistia wilcoxensis E.W. Berry, 58 Pistiaceae Rich. ex C. Agardh, 5 Pistieae Blume, 18, 20, 34, 69, 71, 286 Pistites loriformis Hosius & Marck, 60 Platanaceae Lestib. ex Dumort., 60 Platanus L., 60 Pleea Michx., 61 Plesmonium Schott, 235 Pleuriarum Nakai, 270 Pleurospa Raf., 230 Podolasia N.E.Br., c/plate 112C, 17, 32, 44, 73, 141, 142 Podolasia stipitata N.E.Br., c/plate 112C, 141, 142 Podospadix Raf., 103 Pollenites tranquillus R. Potonié, 60 Polygonaceae Juss., 40 Polygonum L., 63 Pontederiaceae Kunth, 38 Porosia verrucosa (Lesq.) Hickey, 60 Porphyrospatha Engl., 214 Potamogetonaceae Dumort., 63 Potha Kuntze, 98 Potheae Engl., 7, 31, 32, 61, 67, 68, 98 Pothocites grantonii Paterson, 60 Pothoeae Engl., 98

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Pothoideae Engl., 5, 10, 12, 13, 18, 19, 20, 23, 25, 26, 32, 38, 44, 61, 67, 70, 72, 73, 74, 75, 96 Pothoidium Schott, 10, 13, 14, 15, 16, 19, 32, 73, 74, 100, 102 Pothoidium lobbianum Schott, 100, 102 Pothos L., c/plate 108A, 2, 6, 7, 10, 11, 13, 14, 15, 16, 17, 18, 19, 23, 24, 25, 31, 32, 52, 58, 73, 74, 98, 99 Pothos sect. Allopothos Schott, 98 Pothos sect. Pothos, 98 Pothos ser. Goniuri Presl, 25, 98, 100 Pothos acaulis Jacq., 103 Pothos armatus C.E.C. Fisch., 98 Pothos aureus Linden & André, 56 Pothos barberianus Schott, 99 Pothos beccarianus Engl., 99 Pothos celatocaulis N.E.Br., 12 Pothos hosei Rendle, 311 Pothos insignis Engl., 98 Pothos junghuhnii de Vriese, 99 Pothos macrocephalus Scort. ex Hook.f., c/plate 108A Pothos mirabilis Merr., 98 Pothos motleyanus Schott, 99 Pothos officinalis Roxb., 123 Pothos pertusa Roxb., 118 Pothos rumphii (C. Presl) Schott, 24, 98, 99 Pothos sagittaefolia Rudge, 144 Pothos scandens L., 24, 98, 99 Protarum Engl., 34, 69, 74, 275, 277 Protarum sechellarum Engl., 275, 277 Proteinophallus Hook.f., 235 Provenzalia Adans., 146 Pseudodracontium N.E.Br., c/plate 125A, 18, 26, 34, 240, 241 Pseudodracontium anomalum N.E.Br., 240 Pseudodracontium lacourii (Lind. & André) N.E.Br., c/plate 125A, 240 Pseudodracontium latifolium Serebryanyi, 241 Pseudohomalomena A.D. Hawkes, 232 Pseudohydrosme Engl., c/plate 122C, 7, 17, 33, 40, 48, 221, 222 Pseudohydrosme sect. Chorianthera Engl., 221 Pseudohydrosme sect. Pseudohydrosme, 221 Pseudohydrosme sect. Zyganthera (N.E.Br.) Engl., 221 Pseudohydrosme buettneri Engl., 221 Pseudohydrosme gabunensis Engl., c/plate 122C, 35, 221, 222 Psychodidae, 48 Ptiliidae, 48 Ptiloris magnificus Vieillot, 50 Pycnonotus zeylanicus Gmelin, 50 Pycnospatha Thorel ex Gagnep., c/plate 111C, ix, 7, 32, 74, 135, 136 Pycnospatha arietina Thorel ex Gagnep., c/plate 111C, 136 Pycnospatha palmata Thorel ex Gagnep., 135 Pythion Mart., 235 Pythonium Schott, 235

R Raphidophora Hassk., 118 Remusatia Schott, c/plate 129D, 6, 13, 16, 17, 21, 34, 45, 52, 56, 69, 280, 281 Remusatia sect. Gonatanthus (Klotzsch) H. Li & A. Hay, 280 Remusatia sect. Remusatia, 280 Remusatia pumila (D. Don) H. Li & A. Hay, 281 Remusatia vivipara (Roxb.) Schott, c/plate 129D, 41, 45, 50, 280, 281 Remusatia yunnanensis (H. Li & A. Hay) H. Li & A. Hay, 280 Rensselaeria L.C. Beck, 247 Rhaphidophora Hassk., c/plate 109B, 6, 7, 9, 12, 14, 16, 17, 18, 19, 20, 21, 24, 32, 52, 55, 118, 119 Rhaphidophora africana N.E.Br., 119 Rhaphidophora beccarii Engl., 121 Rhaphidophora celatocaulis Alderw., 14 Rhaphidophora decursiva (Roxb.) Schott, 41, 119 Rhaphidophora foraminifera (Engl.) Engl., c/plate 109B, 119 Rhaphidophora glauca (Roxb.) Schott, 119

Rhaphidophora korthalsii Schott, 12 Rhaphidophora lacera Hassk., 118 Rhaphidophora pertusa (Roxb.) Schott, 118 Rhaphidophora pteropoda (Teijsm. & Binn.) Engl., 11 Rhaphidophora sylvestris (Blume) Engl., 119 Rhaphidophora tenuis Engl., 119 Rhaphiophallus Schott, 235 Rhapis L.f. ex Aiton, 15 Rhektophyllum N.E.Br., 228 Rhodospatha Poepp., c/plate 110B, 6, 7, 14, 15, 16, 32, 125, 127 Rhodospatha latifolia Poepp., 45, 125 Rhodospatha oblongata Poepp. & Endl., 127 Rhodospatha perezii G.S. Bunting, c/plate 110B Rhodospatha rubropunctata Croat, 127 Rhodospatha venosa Gleason, 15, 16, 125, 127 Rhopalostigma B.D.Jacks., 160 Rhopalostigmium Schott, 160 Rhynchopyle Engl., 184 Richardia Kunth, 232 Ringentiarum Nakai, 270 Rohdea Roth, 61 Rutelinae, 35

S Sahnipushpam Shukla, 60 Sahnipushpam glandulosum Prakash, 60 Sahnipushpam shuklai Verma, 60 Saruma Oliver, 24 Sauromatum Schott, c/plate 128B, 7, 34, 47, 48, 50, 52, 263, 264 Sauromatum brevipes (Hook.f.) N.E.Br., 264 Sauromatum guttatum Schott, 36, 41, 263 Sauromatum venosum (Aiton) Kunth, c/plate 128B, 7, 36, 38, 41, 46, 50, 56, 263, 264 Scaphidiidae, 48 Scaphispatha Brongn. ex Schott, c/plate 120B, 33, 46, 69, 71, 204, 206 Scaphispatha gracilis Brongn. ex Schott, c/plate 120B, 204, 206 Scarabaeidae, 35, 48 Scheuchzeria L., 22, 66 Scheuchzeriaceae Rudolphi, 63, 66 Schismatoglottideae Nakai, 14, 18, 19, 26, 33, 44, 46, 57, 69, 180 Schismatoglottis Zoll. & Moritzi, c/plate 117C, 14, 16, 17, 18, 23, 26, 33, 35, 44, 48, 52, 55, 182, 183-185 Schismatoglottis Alliance, 71 Schismatoglottis sect. Philonotion (Schott) Bunting, 182 Schismatoglottis acutifolia (Engl.) M. Hotta, 182 Schismatoglottis barbata Engl., 182 Schismatoglottis beccariana Engl., 182 Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi, 182, 184, 185 Schismatoglottis convolvula P.C. Boyce, 183, 346 Schismatoglottis crispata Hook.f., 184, 185 Schismatoglottis ferruginea Merr., c/plate 117C, 184 Schismatoglottis gillianae P.C. Boyce, 183, 184 Schismatoglottis hastifolia Hallier f. ex Engl., 184 Schismatoglottis homalomenoidea M. Hotta, 182 Schismatoglottis hottae Bogner & Nicolson, 183, 184 Schismatoglottis monoplacenta M. Hotta, 182 Schismatoglottis neoguinensis (André) N.E.Br., 185 Schismatoglottis “ruttenii”, 40 Schismatoglottis spruceana (Schott) G.S. Bunting, 182, 183, 184, 185 Schismatoglottis tecturata (Schott) Engl., 185 Schizocasia Schott ex Engl., 283 Sciaridae, 48 Scindapsites Gregor & Bogner, 59 Scindapsites crassus (Reid & Reid) Gregor & Bogner, 59 Scindapsus Schott, c/plate 109D, 9, 12, 14, 16, 24, 32, 45, 55, 122, 123 Scindapsus aureus Engl., 56 Scindapsus beccarii Engl., c/plate 109D, 122 Scindapsus hederaceus Schott, 9

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339

31 Scientific Index Acro 18/7/97 9:47 Page 340

Scindapsus occidentalis Poepp., 125 Scindapsus officinalis (Roxb.) Schott, 122, 123 Scindapsus perakensis Hook.f., 122 Scindapsus pictus Hassk., 14, 38, 122 Scindapsus pothoides Miq., 9 Scindapsus pteropodus Teijsm. & Binn., 11 Scindapsus rupestris Ridl., 122, 123 Scydmaenidae, 48 Seguinum Raf., 153 Serangium Wood ex Salisb., 123 Sericulus chrysocephalus Lewin, 50 Shuklanthus, 60 Silphidae, 48 Simuliidae, 48 Sloanea L., 52 Smilacaceae Vent., 61 Sparganiaceae Rudolphi, 36, 61 Sparganium L., 61 Spathantheum Schott, c/plate 116A, 14, 18, 33, 166, 167 Spathantheum intermedium Bogner, c/plate 116A, 89, 166, 167 Spathantheum orbignyanum Schott, 166, 167 Spathicarpa Hook., c/plate 116B, 33, 46, 166, 168 Spathicarpa burchelliana Engl. 168 Spathicarpa gardneri Schott, c/plate 116B Spathicarpa hastifolia Hook., 166, 168 Spathicarpa lanceolata Engl., 168 Spathicarpeae Schott, 13, 26, 33, 68, 70, 71, 74, 75, 155 Spathiphylleae Engl., 15, 20, 32, 35, 44, 67, 68, 70, 73, 75, 109 Spathiphyllopsis Teijsm. & Binn., 109 Spathiphyllum Schott, c/plate 108D, 5, 6, 14, 17, 18, 20, 31, 32, 44, 46, 47, 48, 49, 52, 55, 56, 58, 59, 60, 70, 73, 74, 109, 111 Spathiphyllum sect. Amomophyllum Engl., 110 Spathiphyllum sect. Chlaenophyllum Nicolson, 110 Spathiphyllum sect. Dysspathiphyllum Engl., 110 Spathiphyllum sect. Massowia (K. Koch) Engl., 26, 110 Spathiphyllum sect. Spathiphyllum, 110 Spathiphyllum beccarii Engl., 110 Spathiphyllum cannifolium (Dryand.) Schott, 15, 42, 56, 81, 111 Spathiphyllum ‘Clevelandii’, 111 Spathiphyllum cochlearispathum Engl., 111 Spathiphyllum floribundum N.E.Br., c/plate 108D, 56 Spathiphyllum humboldtii Schott, 111 Spathiphyllum kochii Engl. & K. Krause, 12 Spathiphyllum laeve Engl., 111 Spathiphyllum lanceifolium (Jacq.) Schott, 109 Spathiphyllum solomonense Nicolson, 110 Spathyema Raf., 96 Spaticarpa Schott, 166 Sphaeroceridae, 48 Sphagnum L., 146 Spirodela Schleid., 10, 22, 58, 63 Spirospatha Raf., 177 Staphylinidae, 48 Stauromatum Endl., 263 Staurostigma Scheidw., 160 Stemonaceae Engl., 61 Stenospermation Schott, c/plate 110C, 14, 15, 16, 18, 32, 45, 58, 128, 129 Stenospermation angustifolium Hemsl., 129 Stenospermation columbiense Engelh., 58 Stenospermation mathewsii Schott, 128 Stenospermation multiovulatum (Engl.) N.E.Br., c/plate 110C Stenospermation rusbyi N.E.Br., 129 Stenospermation ulei K. Krause, 128, 129 Stenospermatium Schott, 128 Stenozeugmaticae Schott, 72 Stenurus Salisb., 266 Steudnera K. Koch, c/plate 129C, 22, 34, 44, 69, 278, 279 Steudnera colocasiifolia K. Koch, c/plate 129C, 21, 278 Steudnera colocasioides Hook.f., 279 Steudnera discolor W. Bull, 279 Steudnera henryana Engl., 279

340

THE GENERA OF ARACEAE

Stichoporella, 60 Stratiotes aloides L., 36 Strelitzia Banks ex Dryander, 43 Strelitziaceae (K.Schum.) Hutch., 60 Strepsanthera Raf., 103 Stylochaeton Lepr., c/plate 114A, 17, 26, 33, 38, 45, 46, 70, 74, 151, 152 Stylochaeton sect. Cyclogyne Engl., 151 Stylochaeton sect. Spirogyne Engl., 151 Stylochaeton sect. Stylochaeton, 151 Stylochaeton bogneri Mayo, 152 Stylochaeton borumensis N.E.Br., 152 Stylochaeton hypogeum Lepr., 151 Stylochaeton lancifolius Kotschy & Peyr., 152 Stylochaeton natalensis Schott, 45, 151 Stylochaeton salaamicus N.E.Br., c/plate 114A Stylochaeton zenkeri Engl., 45, 151, 152 Stylochaetoneae Schott, 20, 33, 64, 68, 70, 151 Stylochiton Schott, 151 Swainsona Salisb., 40 Symplocarpeae Engl., 74 Symplocarpos Schult. & Schult.f., 96 Symplocarpus Salisb. ex Nutt., c/plate 107D, 10, 14, 23, 30, 32, 44, 48, 52, 55, 72, 74, 96, 97 Symplocarpus foetidus (L.) Nutt., 37, 73, 96, 97 Symplocarpus nipponicus Makino, 96 Symplocarpus renifolius Schott ex Miq., c/plate 107D, 96, 97 Synandriospadix Engl., 163 Synandrodieae Buchet, 243 Synandrogyne Buchet, 243 Synandrospadix Engl., c/plate 115D, 33, 56, 163, 164 Synandrospadix vermitoxicus (Griseb.) Engl., c/plate 115D, 30, 163, 164 Synantherias Schott, 235 Syngonium Schott, c/plate 121C, 6, 7, 9, 11, 12, 13, 14, 15, 17, 20, 28, 33, 44, 48, 51, 52, 55, 56, 69, 214, 215 Syngonium sect. Cordata Croat, 216 Syngonium sect. Oblongata Croat, 216 Syngonium sect. Pinnatiloba Croat, 216 Syngonium sect. Syngonium, 216 Syngonium armigerum (Standl. & L.O. Williams) Croat, 215 Syngonium auritum (L.) Schott, 214 Syngonium neglectum Schott, 215 Syngonium podophyllum Schott, 56, 215 Syngonium podophyllum Schott var. peliocladum (Schott) Croat, 17 Syngonium schottianum H. Wendl. ex Schott, 214, 215 Syngonium steyermarkii Croat, c/plate 121C, 214, 215 Syngonium vellozianum Schott, 215 Syrphidae, 48

T Tacca leontopetaloides (L.) Kuntze, 158 Taccaceae Dumort., 61 Taccarum Brongn. ex Schott, c/plate 115A, 33, 56, 158, 159 Taccarum caudatum Rusby, 89, 158 Taccarum warmingii Engl., 159 Taccarum weddellianum Brongn. ex Schott, c/plate 115A, 158, 159 Tapanava Adans., 98 Tapinocarpus Dalzell, 260 Teichosperma spadiciflorum Renner, 59 Telipodus Raf., 169 Thaumatophyllum Schott, 169 Theaceae D. Don, 60 Theriophonum Blume, 26, 34, 56, 260, 261 Theriophonum crenatum (Wight) Blume, 260 Theriophonum infaustum N.E.Br., 261 Theriophonum minutum (Willd.) Baill., 30, 261 Theriophonum sivaganganum (Ramam. & Sebastine) Bogner, 261 Thomsonia Wall., 235

31 Scientific Index Acro 18/7/97 9:47 Page 341

Thomsonieae Blume, 7, 13, 26, 34, 40, 49, 69, 74, 75, 235 Tofieldia Huds., 66, 68, 71 Tornelia Gutierrez ex Schott, 123 Trichopodaceae Hutch., 61 Tricyrtidaceae, 36 Trilliaceae Lindl., 61 Triuridiflorae Dahlgren, Clifford & Yeo, 43 Tupistra Ker Gawl., 61 Turdus merula L., 50 Typhaceae Juss., 29 Typhales Dumort., 61 Typhonium Schott, c/plate 128A, 10, 17, 31, 34, 46, 48, 52, 56, 260, 262 Typhonium sect. Diversifolia Sriboonma & J. Murata, 263 Typhonium sect. Gigantea Sriboonma & J. Murata, 263 Typhonium sect. Hirsuta Sriboonma & J. Murata, 263 Typhonium sect. Pedata Sriboonma & J. Murata, 263 Typhonium sect. Typhonium, 263 Typhonium albispathum Bogner, 45, 263 Typhonium blumei Nicolson & Sivad., 57 Typhonium brownii Schott, 38, 262 Typhonium bulbiferum Dalzell, 18 Typhonium fultum Ridl., 260 Typhonium giganteum Engl., 262 Typhonium hayatae Sriboonma & J. Murata, 260 Typhonium hirsutum (S.Y. Hu) J. Murata & Mayo, 87, 260, 262, 263 Typhonium nudibaccatum A. Hay, 260 Typhonium trilobatum (L.) Schott, c/plate 128A, 260, 262, 263 Typhonium violifolium Gagnep., 18 Typhonodoreae Engl., 245 Typhonodorum Schott, c/plate 126B, 6, 14, 17, 26, 28, 34, 46, 51, 52, 58, 247, 249 Typhonodorum lindleyanum Schott, c/plate 126B, 54, 55, 247, 249

U Ulearum Engl., c/plate 120A, 31, 33, 69, 202, 203 Ulearum reconditum Madison, 155 Ulearum sagittatum Engl., 31, 202, 203 Ulearum sagittatum Engl. var. viridispadix Bogner, c/plate 120A Uromyces sparganii, 61 Urophyllum K. Koch, 144 Urospatha Schott, c/plate 113A, 32, 48, 56, 73, 144, 145 Urospatha angustiloba Engl., 145 Urospatha desciscens Schott, 130 Urospatha sagittifolia (Rudge) Schott, 144, 145 Urospatha tonduzii Engl., c/plate 113A Urospatha wurdackii (G.S. Bunting) A. Hay, 144, 145 Urospathella G.S. Bunting, 144 Urospathella wurdackii G.S. Bunting, 144 Urospathites Gregor & Bogner, 59 Urospathites dalgasii (Hartz) Gregor & Bogner, 59 Urticaceae Juss., 41

V

X Xanthosoma Schott, c/plate 121A, 6, 7, 13, 16, 21, 29, 33, 35, 44, 47, 48, 53, 55, 58, 69, 209, 212 Xanthosoma sect. Acontias (Schott) Engl., 56, 211 Xanthosoma sect. Xanthosoma, 57, 211 Xanthosoma atrovirens K. Koch & Bouché, 53 Xanthosoma brasiliense (Desf.) Engl., 53 Xanthosoma caracu K. Koch & Bouché, 53 Xanthosoma helleborifolium (Jacq.) Schott, 212 Xanthosoma lindenii (André) Engl., 40 Xanthosoma mafaffa Schott, 53 Xanthosoma plowmanii Bogner, 89, 211, 212 Xanthosoma pubescens Poepp., 16, 209 Xanthosoma robustum Schott, 42, 53 Xanthosoma sagittifolium (L.) Schott, 22, 52, 53, 55, 209, 211 Xanthosoma striatipes (Kunth) Madison, 212 Xanthosoma tarapotense Engl., 13 Xanthosoma violaceum Schott, c/plate 121A, 17, 53, 212 Xanthosoma viviparum Madison, 209 Xenophya Schott, 283

Z Zala Lour., 286 Zamia L., 149 Zamioculcadeae Engl., 13, 18, 20, 33, 35, 64, 68, 70, 73, 74, 75, 146 Zamioculcas Schott, c/plate 113C, 7, 13, 15, 16, 17, 18, 23, 26, 33, 62, 146, 148 Zamioculcas boivinii Decne., 149 Zamioculcas loddigesii Schott, 146 Zamioculcas zamiifolia (Lodd.) Engl., c/plate 113C, 18, 46, 146, 148 Zantedeschia Spreng., c/plate 124A, 10, 11, 14, 17, 18, 22, 24, 29, 34, 47, 48, 55, 56, 232, 233 Zantedeschia aethiopica (L.) Spreng., c/plate 124A, ix, 38, 40, 41, 47, 52, 56, 57, 230, 232, 233 Zantedeschia albomaculata (Hook.f.) Baill., 56 Zantedeschia albomaculata (Hook.f.) Baill. subsp. albomaculata, 233 Zantedeschia odorata P.L. Perry, 232 Zantedeschia rehmannii Engl., 38, 40, 233 Zantedeschieae Engl., 13, 18, 26, 34, 68, 230 Zingiberaceae Lindl., 42, 60 Zingiberales Nakai, 36, 65 Zingiberiflorae Dahlgren, Clifford & Yeo, 43 Zomicarpa Schott, c/plate 119C, 28, 33, 69, 74, 199, 200 Zomicarpa pythonium (Mart.) Schott, 199 Zomicarpa riedeliana Schott, c/plate 119C Zomicarpa steigeriana Schott, 199 Zomicarpeae Schott, 13, 20, 26, 33, 66, 71, 199 Zomicarpella N.E.Br., c/plate 119D, 33, 69, 199, 201 Zomicarpella amazonica Bogner, c/plate 119D, 45, 199, 201, 202 Zomicarpella maculata N.E.Br., 199, 201, 202 Zosteraceae Dumort., 63 Zyganthera N.E.Br., 221

C

Vallisneria spiralis L., 36 Viracarpon Sahni, 60 Viracarpon elongatum Sahni, 60 Viracarpon hexaspermum Sahni, 60 Visnea L., 60

I N D E X TO S C I E N T I F I C N A M E S

341

32 Subject Index Acro 18/7/97 9:58 Page 342

32 S U B J E C T I N D E X

C

A adventitious roots, 10 alkaloidal glycosidase inhibitors (AGIs), 40 amines, 41 amphivasal stem vascular bundles, 15 anastomosing laticifers, 20 anchor roots, 9, 45 Andrade, I.M., 5 aneuploidy, 31 animal dispersal, 50 anterior division of leaf, 7, 8 anther, 26 anthocyanins, 38 aperigoniate Aroideae, 72 appendix, terminal, 26, 48 aquatic and subaquatic ornamentals, 55 aquatics, free-floating, 46 aquatics, submerged or periodically submerged, 46 Araceae, paraphyly of, 65 Araceae, phylogenetic relationships with other monocots, 61 Araceae, phylogenetic relationships within family, 66 Araceae, primitive characters, 62 Araceae, relationships with Acoraceae, 61 Araceae, relationships with Alismatiflorae, 62 Araceae, relationships with Lemnaceae, 63-65 arborescent plants, 6 aril, 28 Aroideana, 5 articles, 6 asarone, 42 axial stem bundles, 14

B Barabé, D., 5 Barros, C., 5 basal rib, 7, 8 Berlin, 4, 5 Besse, L., xi bicchu-shu, 53 biforines, 23 biogenic amines and alkaloids, 41 birds of paradise, 50 Birdsey, M., 5 blackbirds, 50 Blanc, P., 5, 6 Blume, C.L., 3 bog gardens, 55 Bogner, J., ix, xi, 5, 73, 74, Bown, D., ix Boyce, P.C., ix, 5 Brazil, 5 Brown, N.E., 4 Buchet, S., ix bud traces, 16 bulbils, 6 bulbul bird, 50 Bunting, G.S., ix, 5

C C-glycoflavones, 38 calcium oxalate crystals, 22, 36 Calicut, 5

342

THE GENERA OF ARACEAE

carbohydrates, 36 Carvell, W.N., 5 Catherine, E., ix, xi chemotaxonomy, 43 cataphylls, 6 chira take, 53 chiroko, 53 Chrétien, L., 5 chromosomes, primary basic number, 31 Chung, M., 5 cladistics of Araceae, 61-71 classifications, previous, 72 cocoyam, 53, 55, 211 colocasioid venation, 8, 311 common bush tanager, 50 common palm civet, 50 compound stem vascular bundles, 14-16 connectives, 26 conservation, 57 contact dermatitis, 40 continuation shoot, 6 contractile roots, 6, 10 Copenhagen, 5 cortical stem vascular bundles, 13 cotyledonar hyperphyll, 29 cotyledonar sheath, 29 cpDNA characters, 5, 61, 66 crassinucellate ovules, 30 Croat, T.B., 5 crop species, 53, 55 crystal idioblasts, 22 crystal sand, 22 cultivated aroids, 55 cut flowers, ix, 56 cuticle, 17 cyanogenic glucosides, 38 cytology, 31

D dimorphic roots, 6, 9 dispersal, 50 DNA studies, 67, 70 Dodoens, R., 2 dormancy, seasonal, of growth, 45, 56 dracontioid leaf form, 7 druses, 23

E ecology and life forms, 44 eddoe, 283 egumi-taste, 41 elephant yam, 53, 55 embryogeny types in Araceae, 30 embryology, 30 endemism, 52 endodermis of root, 12 endodermis of stem, 14 endosperm, 28, 30 endosperm development, 30, 62 endothecial thickenings, 27 endothelium, 30 Engler, A., 3-4, 72-73 epicuticular waxes, 43

32 Subject Index Acro 18/7/97 9:58 Page 343

epidermal cells, 10, 13, 16 epiphytes, 6, 45 essential oils, 42 exine ornamentation, 35 extrafloral nectaries and punctations, 22

F feeder roots, 9, 45 fibres, useful, 54 first leaf, 7, 29 flagelliform shoots, 6, 45 flavones, 38 flavonoids, 38 flavonols, 38 floral anatomy, 27 floral morphology, 25 floral sympodium, 6 Florida, 5 flowers, 26 food plants, 53 Forget, S., 5 fossil aroids, 58 fossils, excluded from Araceae, 60 free-floating aquatics, 46 French, J.C., x, 5, 9, 75 fruit presentation, 50 fruits, 28 fruits, fossil, 59 Fuchs, L., 2 funicle trichomes, 26

G Gagnepain, F., ix Genera Aroidearum, 2 geniculum, 18 geography, 52 geophytes, 6, 45 giant taro, 52, 53, 55 glucomannans, 36 Gonçalves, E.G., 5 Grayum, M.H., 5, 74 guttation, 22 gynoecium, 26

H half hardy species, 55 hardy species, 55 Harvard Forest, 5 haustorium, 29 Hay, A., ix, 5, 75 Hegnauer, R., x, 36 helophytes, 46 hemiepiphytes, 6, 46 Hernandez, F., 2 heteroblasty, 7, 45 Hetterscheid, W., 5 higher order leaf venation, 8, 18, 311 honeyeater birds, 50 Hooker, J.D., 5, 73 hornbills, 50 Hotta, M., ix, 5, 74 houseplants, ix, 55 Humboldt, A. von, 3 Hur, Y., 5 Hutchinson, J., 5, 73 hybrids, 56 hydathodes, 22 hydrochory, 51 hydroculture, 55 hydrophytes, 46

hypocotyl, 29 hypodermis, of leaf, 17 hypodermis, sclerotic, of root, 11 hypogeal stolons, 6

I Icones Aroidearum, 2 indoles, 41 inflorescence anatomy, 27 inflorescence morphology, 25 inflorescence odours, 41, 47 infructescence, 28, 50 infructescence, fossils of, 59 inner cortex of root, 11 International Aroid Society, 5 internode development, 16 intravaginal squamules, 22 irritants, 40-41

J Jacobsen, N., 5 Jacquin, N.J., 3 Jumelle, H., ix Jussieu, A.L., 2

K Kagoshima, 5 keladi, 207 Kew, 4, 5 Kite, G., 5 kolokasia, 283 konjac, 53 Krause, K., 3, 4 Kunming, 5 Kunth, C.S., 3 Kyoto, 5

L Labrecque, M., 5 latex, 21 latex cytology, 21 laticifers, 20 leaf blade shape, 7, 313 leaf epidermis, 16 leaf fossils, 58 leaf mesophyll, 17 leaf morphology, 7-8 leaf punctations, 22 leaf structure of Acorus, 19 leaf tubercles and regeneration, 18 leaf vasculature, 18 leaf venation, 8, 18, 311 Li Heng, 5 life forms, 44 ligule, 19 Linnaeus, C., 2 Lins, A., 5 lithophytes, 45 long-tailed manakin, 50

M Mabberley, D., 5, 75 Madison, M.T., ix, 5 magical and ritual uses, 54 Major Group Proto-Araceae, 66 Major Group True Araceae, 67 Marchant, C., 5, 74

SUBJECT INDEX

343

32 Subject Index Acro 18/7/97 9:58 Page 344

Martius, C.F.P. von, 3 Mayo, S.J., ix, 5 medicinal uses, 54 megaspore tetrads, 30 Meletemata botanica, 2, 72 mesophytic habit, 48 mesophyll, leaf, 17 midrib, 7 Mikan, J.C., 3 mineral crystals, 22 mineral deposits and primary metabolites, 36 molecular studies, 5, 61, 62, 63, 64, 66, 67, 70, 71, 75 monkey dispersal, 51 Monographiae Phanerogamarum, 3, 73 Moscow, 5 mucilage cells, canals, and cavities, 21 mucilages, 36 Munich, 4, 5 Murata, J., 5 myrmecochory, 51

Nadruz Coelho, M.A., 5 Nakai, T., 5 nama konjaku, 53 Natterer, K., 3 naturalized taxa, 52 nest roots, 10 Nicolson, D.H., ix, 5, 74 non-allergenic skin irritations, 41 Ntépé-Nyame, C., 5 nucellar cap, 30

phytochemistry, 36 phytosterols, 42 pistil, see gynoecium pistillodes, 49 placentation, 26 plumule, 28, 29 Pohl, J., 3 pollen aperture type, 35 pollen, fossil, 59 pollen grain size, 35 pollen mother cell division, 30 pollen shape, 35 pollen starch, 35 pollination biology, 47 polyhydroxy alkaloids, 40 polyploidy, 31 pond gardens, 55 posterior divisions, 7, 8 prickles, 17 primary lateral veins, 7, 18 primary root, 9 primitive characters in Araceae, 62 prisms, 22 procyanidins, 38 Prodromus systematis Aroidearum, 2, 72 prophyll, 6 protection of developing fruits and seeds, 28 proto-Araceae, 66 protogyny, 49 pseudomonomery, of gynoecia, 27 pseudostem, 6 pulvinus, of leaf, 18 punctations, leaf, 22

O

Q

odours, of inflorescence, 41, 47 Oesterreiches Botanisches Wochenblatt, 2 Oesterreichische Botanische Zeitschrift, 2 operculum, 28 origin of aroids, 61 ornamental Araceae, 55 ornamental plants, 54, 55, 56 ornithochory, 50 osmophores, 26, 27, 49, ovular trichomes, 26, 27 ovule morphology, 26 ovule vasculature, 27 oxalic acid, 36

qolqas, 283

N

P palisade parenchyma, of leaf, 17 palynology, 35 paper-making, 54 parallel-pinnate leaf venation, 8, 18, 311 paraphyly, 65 pectins, 37 peduncle, 25 perforated (fenestrate) leaves, 7 periderm formation, 12 perigoniate Aroideae, 70 periplasmodial anther tapetum, 30 perisperm, 30 Petersen, G., 5, 31 petiole anatomy, 18, 19 Pflanzenfamilien, die natürlichen, 3 Pflanzenreich, Das, 3 phenolic compounds, 38 phloem cytology, 24 phylogenetic relationships with other monocots, 61 phylogenetic relationships within Araceae, 66

344

THE GENERA OF ARACEAE

R Ramalho, F.C., 5 range disjunctions, 52 raphides, 22-23, 36, 41 Ray, J., 2 Ray, T., 5, 6 regant bowerbird, 50 regeneration, from leaf, 18 resin canals and cavities, 21 resplendent quetzal, 50 reticulate leaf venation, 8, 18, 311 rheophytes, 46 rheophytes, cultivation, 56 rhizoids, 29 Riedl, H., 3, 5 Rio de Janeiro, 3 robins, 50 rock gardens, 55 root anatomy, 9-12 root apex, 12 root collar, 29 root cortex, 11 root epidermis, 10 root exodermis, 11 root growth, 10 root hairs, 10-11 root morphology, 6 root periderm formation, 12 root vascular cylinder, 12 rosulate plants, 6 rust fungus, 61 Rutgers, 5

32 Subject Index Acro 18/7/97 9:58 Page 345

S

T

Sakuragui, C.M., 5 saponins, 37 Sarasota, 5 Schönbrunn, 2, 3 Schott, H.W., 2-3, 72 sclerotic hypodermis of root, 11 seasonally dormant genera, cultivation, 56 secondary metabolites, 37 secretory anther tapetum, 30 seed morphology, 28 seedling morphology, 29 seeds, fossil, 59 Serebryanyi, M., 5 Seubert, E., 5, 23, 29 shina-shu, 53 shingle plants, 7, 45 shoot organization, 6 sister group relationships of Araceae, 61-65 Sivadasan, M., 5 skin irritation, non-allergenic, 41 skototropism, 45 Soares, M.L., 5 spadix, 25 spadix fossils, 59 spadix stipe, 25 spathe constriction, 25, 49 spathe, 32 Spix, J.B. von, 3 spongy mesophyll, 17 squamules, intravaginal, 22 St. Louis, 5 stamens, 26, 49 stamen vasculature, 27 staminodes, 49 starch grain morphology, 24, 37 starch in leaves, 36 starch sources, 53 starch storage, 37 stem anatomy, 13-16 stem central cylinder, organization of, 14 stem cortex, 13 stem endodermis, 14 stem epidermis, 13 stem morphology, 6-7 stem vascular bundles, course of, 15 sticky seeds, 28 stigma, 26 stomata, 17 strophiole, 28 stylar region, 26 styloids, 22 submerged aquatics, 46 subsidiary cells of stomata, 17 successive pollen mother cell division, 30 swamp taro, 53, 55 Sydney, 5 sympodial leaf, 6 sympodial units, 6 sympodium, 6 synandria, 26 syncarp, 28 synhospitality, 47

tannia, 211 tapetum, types, 30 taro, 36, 53, 55, 283 tawny-capped euphonia, 54 temperate ornamental species, 55 tenuinucellate ovules, 30 tepals, 26 terminal appendix, of spadix, 26, 48 testa, 28 Thanikaimoni, G., 5, 73 theca dehiscence, 26 Theophrastus, 2 thermogenesis, 48 threatened species, 57 Tillich, H.-J., 5, 29 Tokyo, 5 Tomlinson, P.B., 5 Tournefort, J.P. de, 2 toxic effects, 40, 54 trichomes, 16-17 trichosclereids, 19 triglochinin, 40 triterpenes, 42 tropical aquatics and helophytes, 56 tropical ornamental species, 55-56 true Araceae, 67 tubercles, 18 tubers, 6, 13, 45

U useful aroids, 53-54

V vascular system, 12, 14-16, 18 vegetative anatomy, 9-24 venation, leaf, 8, 18, 311 vessels, 24 Vienna, 2, 3 volatile amines and indoles, 41

W Waechter, J., 5 Washington DC., 5 Waterbury, Betty, xi wax crystalloids, 17 Wiener Zeitschrift für Kunst, Literatur, Theater und Mode, 2 Wit, H.C.D. de, 5 wood thrush, 50 woodland gardens, 55

Y Yaoundé, 5

Z zairai-shu, 53

C

SUBJECT INDEX

345

33 New Taxa Acro 18/7/97 10:00 Page 346

33 D E S C R I P T I O N S O F N E W TA X A

C

Subfamily Orontioideae Mayo, Bogner & P.C. Boyce, subfam. nov. Typus: Orontium L. Herbae palustres; caudex rhizomatosus, crassus, erectus, sympodialis, sympodii ramus novus ex axilla folii supremi ortus; petiolus apice non geniculatus, petioli vagina longa; folii lamina integer, oblongo-elliptica, venae tenues reticulatae; flores bisexuales, perigonio instructi; tepala libera, fornicata, stamina filamentis distinctis munita; antherae terminales; connectivum gracile; thecae longitudinaliter dehiscentes; pollinis granum unisulcatum; seminis testa laevis, tenuis vel maturitate nulla; embryo grandis; endospermium vix adest vel omnino abest.

Schismatoglottis convolvula P.C. Boyce, sp. nov. Typus: Malaysia, Sarawak, J. Mamit S.42101 (SAR!, holotypus; L!, US!, isotypi). See p. 183 (Plate 49 (i) E). Schismatoglottis convolvula ab omnibus speciebus generis ceteris caulibus longis tenuis repentibus folios dispersos ferentibus distinguitur. Caules, ab origine vulgari orientes, hapaxanthi videntur, caule quoque vel paene ita basaliter innovationes post anthesin parienti, tum morientes.

346

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 107. A, Gymnostachys anceps; B, Orontium aquaticum; C, Lysichiton camtschatcensis; D, Symplocarpus renifolius.

COLOUR PLATES

347

C

A

B

C

D

Plate 108. A, Pothos macrocephalus; B, Pedicellarum paiei ; C, Anthurium flavolineatum; D, Spathiphyllum floribundum.

348

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 109. A, Amydrium medium; B, Rhaphidophora foraminifera; C, Epipremnum falcifolium; D, Scindapsus beccarii.

COLOUR PLATES

349

C

A

B

C

D

Plate 110. A, Monstera adansonii; B, Rhodospatha perezii; C, Stenospermation multiovulatum; D, Dracontium changuango.

350

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 111. A, Dracontioides desciscens; B, Anaphyllopsis americana; C, Pycnospatha arietina; D, Anaphyllum beddomei.

COLOUR PLATES

351

C

A

B

C

D

Plate 112. A, Cyrtosperma carrii; B, Lasimorpha senegalensis; C, Podolasia stipitata; D, Lasia spinosa (form with entire leaf blades).

352

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 113. A, Urospatha tonduzii; B, Calla palustris ; C, Zamioculcas zamiifolia; D, Gonatopus marattioides.

COLOUR PLATES

353

C

A

B

C

D

Plate 114. A, Stylochaeton salaamicus; B, Dieffenbachia maculata; C, Bognera recondita; D, Mangonia tweedieana.

354

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 115. A, Taccarum weddellianum; B, Asterostigma riedelianum; C, Gorgonidium vermicidum; D, Synandrospadix vermitoxicus.

COLOUR PLATES

355

C

A

B

C

D

Plate 116. A, Spathantheum intermedium; B, Spathicarpa gardneri; C, Philodendron rugosum; D, Furtadoa sumatrensis.

356

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 117. A, Homalomena geniculata; B, Anubias gigantea; C, Schismatoglottis ferruginea; D, Piptospatha burbidgei.

COLOUR PLATES

357

C

A

B

C

D

Plate 118. A, Hottarum lucens; B, Bucephalandra motleyana; C, Phymatarum borneense; D, Aridarum annae.

358

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 119. A, Lagenandra nairii; B, Cryptocoryne longicauda; C, Zomicarpa riedeliana; D, Zomicarpella amazonica.

COLOUR PLATES

359

C

A

B

C

D

Plate 120. A, Ulearum sagittatum; B, Scaphispatha gracilis; C, Caladium tuberosum; D, Jasarum steyermarkii.

360

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 121. A, Xanthosoma violaceum; B, Chlorospatha longipoda; C, Syngonium steyermarkii; D, Hapaline celatrix

COLOUR PLATES

361

C

A

B

C

D

Plate 122. A, Nephthytis afzelii var. graboensis ; B, Anchomanes abbreviatus ; C, Pseudohydrosme gabunensis ; D, Aglaonema tenuipes.

362

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 123. A, Aglaodorum griffithii ; B, Culcasia saxatilis ; C, Cercestis ivorensis ; D, Montrichardia linifera.

COLOUR PLATES

363

C

A

B

C

D

Plate 124. A, Zantedeschia aethiopica; B, Callopsis volkensii; C, Amorphophallus prainii; D, Amorphophallus aphyllus.

364

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 125. A, Pseudodracontium lacourii; B, Arophyton crassifolium; C, Carlephyton glaucophyllum; D, Colletogyne perrieri.

COLOUR PLATES

365

C

A

B

C

D

Plate 126. A, Peltandra virginica (mature infructescences); B, Typhonodorum lindleyanum; C, Arisarum simorrhinum; D, Ambrosina bassii.

366

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 127. A, Arum maculatum; B, Eminium lehmannii; C, Dracunculus vulgaris; D, Helicodiceros muscivorus.

COLOUR PLATES

367

C

A

B

C

D

Plate 128. A, Typhonium trilobatum; B, Sauromatum venosum; C, Biarum ditschianum; D, Pinellia cordata.

368

THE GENERA OF ARACEAE

C

A

B

C

D

Plate 129. A, Arisaema ovale var. ovale; B, Ariopsis peltata; C, Steudnera colocasiifolia; D, Remusatia vivipara.

COLOUR PLATES

369

C

A

B

C

D

Plate 130. A, Colocasia esculenta; B, Alocasia brisbanensis; C, Pistia stratiotes; D, Acorus calamus (mature infructescence).

370

THE GENERA OF ARACEAE