A Field Guide to Tropical Plant Families 3031059417, 9783031059414

Table of contents :
Foreword by David Mabberley
Foreword by Jitka Klimešová
Acknowledgements
Contents
About the Author
User Manual and Architectural Concepts
1 Introduction
1.1 Scope of the Book
1.2 Why Consider Families?
1.3 Photos or Drawings?
1.4 Sources and Tests
1.5 Structure of the Book
References
2 How to Use This Book
References
3 Architectural Concepts and Key to the Models
3.1 Concepts and Statements
3.2 Key to the Architectural Models
References
Identification Keys
4 Key to the Groups of Families of Trees and Shrubs
5 Keys to Families of Trees and Shrubs
6 Supplementary Keys
6.2 Key B—Tribes and Main Genera of the Malvaceae
6.3 Key C—Families of the Order Sapindales + Picramniaceae
6.4 Key D—Tribes and Main Genera of the Leguminosae
6.5 Key E—Some Genera of the Rubiaceae
References
7 Keys to the Groups of Families of Lianas and Woody Epiphytes
8 Keys to Families of Lianas and Woody Epiphytes
Description of the Families
9 Acanthaceae to Asteraceae
9.1 Acanthaceae: 36-37-38-56
9.2 Achariaceae: 15-16-18-20-27
9.3 Actinidiaceae: 24-28
9.4 Anacardiaceae: 3-24-28-29-44-C
9.5 Ancistrocladaceae: 54
9.6 Anisophylleaceae: 13-20
9.7 Annonaceae: 10-54
9.8 Apocynaceae: 3-4-53
9.9 Aquifoliaceae: 18
9.10 Araliaceae: 19-26-47-48-49-57
9.11 Araucariaceae: 1-42
9.12 Arecaceae: 1-57 (Monocotyledons)
9.13 Aristolochiaceae: 12-55
9.14 Asteraceae: 21-23-38-40-56
References
10 Bignoniaceae to Buxaceae
10.1 Bignoniaceae: 21-40-43-49-51-52-56-57
10.2 Bixaceae: 2-15-47
10.3 Boraginaceae (incl. Cordiaceae, Ehretiaceae): 20-21-24-25-30
10.4 Burseraceae: 3-46-C
10.5 Buxaceae: 39-44
References
11 Cactaceae to Cycadaceae
11.1 Cactaceae: 21
11.2 Canellaceae (C) + Winteraceae (W): 13-23-29-30
11.3 Capparaceae (incl. Cleomaceae): 8-16-21-22-27-29-30-47
11.4 Caricaceae: 3-20-49
11.5 Casuarinaceae: 37
11.6 Celastraceae: 4-8-13-16-18-21-30-33-34-45-53-54-56
11.7 Chloranthaceae: 32
11.8 Chrysobalanaceae: 2-5-7-8-13
11.9 Clethraceae (CL) + Cyrillaceae (CY): 28-29
11.10 Clusiaceae + Calophyllaceae + Bonnetiaceae + Hypericaceae: 2-3-4-11-13-24-29-38-41-42
11.11 Combretaceae: 21-22-24-30-40-41-56
11.12 Connaraceae: 12-21-48-49-53-57
11.13 Convolvulacaeae: 53-54-57
11.14 Cornaceae (C) + Nyssaceae (N): 13-29
11.16 Cunoniaceae (C) + Brunelliaceae (B): 32-50
11.17 Cycadaceae + Zamiaceae: 1 (Gymnosperms)
11.18 Cyclanthaceae: 1-57 (Monocotyledons)
References
12 Dichapetalaceae to Dipterocarpaceae
12.1 Dichapetalaceae: 7-8-54
12.2 Dilleniaceae: 14-26-28-54
12.3 Dioscoreaceae: 55-56-57 (Monocotyledons)
12.4 Dipterocarpaceae: 2-5-7-8-15-18
References
13 Ebenaceae to Euphorbiaceae
13.1 Ebenaceae: 11-12-40
13.2 Elaeocarpaceae: 15-16-18-27-34
13.3 Ericaceae: 12-13-20-29-30
13.4 Erythropalaceae: 3-12-13
13.5 Erythroxylaceae: 8-32
13.6 Euphorbiaceae + Peraceae: 2-4-6-8-14-16-17-18-19-20-22-27-32-34-47-53-55-A
References
14 Fagaceae
14.1 Fagaceae: 8-9-18
References
15 Gentianaceae to Goupiaceae
15.1 Gentianaceae: 37-38
15.2 Gnetaceae: 36-39-53-56
15.3 Goodeniaceae: 24-29
15.4 Goupiaceae: 9
References
16 Hamamelidaceae to Humiriaceae
16.1 Hamamelidaceae: 5-6-8-19-29 (incl. Altingiaceae)
16.2 Hernandiaceae: 20-55-57
16.3 Huaceae: 8
16.4 Humiriaceae: 9-13-22-30
References
17 Icacinaceae s.l. to Irvingiaceae
17.1 Icacinaceae s.l. (Icacinaceae s.str. I + Metteniusaceae M + Stemonuraceae S) + Cardiopteridaceae C: 12-13-25-30-44-54-56
17.2 Irvingiaceae: 5
18 Juglandaceae
18.1 Juglandaceae: 48
References
19 Lamiaceae to Lythraceae
19.1 Lamiaceae: 35-38-39-43-44-45-51-52-56
19.2 Lauraceae: 12-20-21-23-24-29-30-39-44
19.3 Lecythidaceae: 9-10-13-15-18-25-29
19.4 Leguminosae—Introduction
19.5 Leguminosae–Cercideae–Cassieae–Detarieae–Caesalpinieae: 47-48-55-D
19.6 Leguminosae–Mimoseae–Acacieae–Ingeae: 20-30-46-57-D
19.7 Leguminosae–Papilionoideae: 2-6-8-9-46-47-50-53-54-57-D
19.8 Linaceae–Hugonioideae (H) + Ixonanthaceae: 13-18-29-30-54 (I)
19.9 Loganiaceae + Gelsemiaceae: 32-33-35-38-56
19.10 Lythraceae: 33-37-38-41
References
20 Magnoliaceae to Myrtaceae
20.1 Magnoliaceae: 5-14
20.2 Malpighiaceae: 4-17-33-40-44-56
20.3 Malvaceae: 5-6-8-14-15-47-55-B
20.4 Marcgraviaceae: 13-54
20.5 Melastomataceae: 35-37-39-41-44
20.6 Meliaceae: 12-24-48-51-C
20.7 Menispermaceae: 19-55-57
20.8 Monimiaceae s.l.: 40-44-45
20.9 Montiniaceae: 44
20.10 Moraceae: 2-4
20.11 Moringaceae: 48
20.12 Muntingiaceae: 6
20.13 Myricaceae: 30
20.14 Myristicaceae: 3-12
20.15 Myrtaceae: 4-23-39-40-41-42-44
References
21 Nyctaginaceae
21.1 Nyctaginaceae: 36-41-44-56
Reference
22 Ochnaceae to Oxalidaceae
22.1 Ochnaceae: 8-9-18-34-50
22.2 Olacaceae (incl. Schoepfiaceae): 12-13-21
22.3 Oleaceae: 35-40-44-45-51-52-56-57
22.4 Opiliaceae: 13-30
22.5 Oxalidaceae: 27-48-49-54
Reference
23 Pandaceae to Putranjivaceae
23.1 Pandaceae: 8-A
23.2 Pandanaceae: 1-54 (Monocotyledons)
23.3 Passifloraceae: 6-7-17-22-55-57
23.4 Pentaphylacaceae: 12-23-29-30
23.5 Phyllanthaceae: 2-3-7-8-27-47
23.6 Physenaceae: 30
23.7 Phytolaccaceae (incl. Petiveriaceae): 30-54
23.8 Picrodendraceae: 31-33-44-49-52-A
23.9 Pinaceae: 1-37 (Gymnosperms)
23.10 Piperaceae: 13-20
23.11 Pittosporaceae: 29
23.12 Poaceae (Bambuseae): 1 (Monocotyledons)
23.13 Podocarpaceae: 1
23.14 Polygalaceae: 11-54
23.15 Polygonaceae: 5-14-54
23.16 Primulaceae: 11–23-29
23.17 Proteaceae: 25-44-45-48
23.18 Putranjivaceae: 8-9-13-A
References
24 Rhabdodendraceae to Rutaceae
24.1 Rhabdodendraceae: 23
24.2 Rhamnaceae: 6-8-9-17-18-31-32-33-54-55-56
24.3 Rhizophoraceae: 32-33-34
24.4 Rosaceae: 7-11-15-46
24.5 Rubiaceae: 32-56-E
References
25 Sabiaceae to Symplocaceae
25.1 Sabiaceae: 25-30-48-54
25.2 Salicaceae (incl. Lacistemataceae): 6-7-8-9-13-17-18-22-30
25.3 Santalaceae (S) + Loranthaceae (L): 12-30-37-44
25.5 Sapotaceae: 2-3-4-24
25.6 Schisandraceae: 23
25.7 Simaroubaceae (S) + Picramniaceae (P): 11-21-29-46-48-C
25.8 Smilacaceae: 55 (Monocotyledons)
25.9 Solanaceae: 11-12-21-30-54
25.10 Styracaceae: 12-22
25.11 Surianaceae: 21
25.12 Symplocaceae: 30
References
26 Tetramelaceae to Torricelliaceae
26.1 Tetramelaceae: 20
26.2 Theaceae: 12-29-30
26.3 Thymelaeaceae: 10-25-44-54-56
26.4 Torricelliaceae: 26
Reference
27 Ulmaceae s.l. to Urticaceae
27.1 Ulmaceae (U) + Cannabaceae (C): 5-6-8-31
27.2 Urticaceae: 2-6-14-15-31-47
Reference
28 Verbenaceae to Vochysiaceae
28.1 Verbenaceae + Callicarpa: 23-35-38-40-41-44-56
28.2 Violaceae: 8-9-18-33-34-54
28.3 Vitaceae: 5-6-14-46-47-55-57
28.4 Vochysiaceae: 33-42
References
29 Zygophyllaceae s.l. (incl. Balanitaceae)
29.1 Zygophyllaceae (incl. Balanitaceae): 48–50
30 Herbaceous Versus Arborescent Habit in the Families and Orders of Flowering Plants—Evolutionary Trends
30.1 A Diagrammatic Representation of Angiosperm Evolution
30.2 References to Phylogenetic Studies
30.3 Discussion and Hypothesis
References
Appendix_1
Index

Citation preview

Roland Keller

A Field Guide to Tropical Plant Families

A Field Guide to Tropical Plant Families

Roland Keller

A Field Guide to Tropical Plant Families

123

Roland Keller Lausanne, Switzerland

ISBN 978-3-031-05941-4 ISBN 978-3-031-05942-1 https://doi.org/10.1007/978-3-031-05942-1

(eBook)

© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Foreword by David Mabberley

Botanists, formerly (and with differing motives), collected, described and named the world’s trees and other plants as the principal task of the plant sciences. The practical necessity to classify plants according to their uses is at the root of plant systematics, which is therefore one of the oldest of all sciences, being evident in the historical records of all major civilisations. Despite this longevity and an enormous literature, both printed and online, the fact remains that we still cannot accurately state how many plant species there are (or, very often, even agree on what constitutes a species), while details of their overall morphology and ecology often not known at all. Each year some 2000 new plant species are described as new, and we do not know how many undescribed ones remain undiscovered in the field or lie fallow as preserved specimens in museum collections. Yet, despite the hullabaloo about conservation of resources, endless conferences, ‘declarations’, ‘accords’ and pious sentiments from politicians, it is clear that, following, and greatly accelerating, the environmental modifications wrought by their forebears, modern humans have caused the extermination of hordes of animal and plant species we know about—and, undoubtedly, many that were never documented. We live in the Anthropocene and are witnessing the fastest of the great extinction events ever seen on earth—and it is our fault. Besides the rising tide of alarm about climate change and this extinction, there are increasing public concerns about, or general interest in plants in general: organic food, herbal remedies and supplements, biofuels, ‘superfoods’, aromatherapy, genetically modified organisms and invasive species; gardening and the mere presence of plants are increasingly seen as good therapy. In short, people are engrossed by plants and, I would argue, the fathoming of the magnificent natural world (incidentally usually feminine as in Mother Earth and Mother Nature) of which we form an increasingly large part, is perhaps the greatest contribution that we scientists, as part of that magnificence, can possibly do. A 1946 paper of Corner’s in New Phytologist was prescient, ‘I fear lest all the virgin lowland forest of the tropics may be destroyed before botany awakes: even our children may never see the objects of our delight which we have not cared for in their vanishing ….there is still a vast amount of hack-work to be done ….’. Contrary to what the concerned layperson might believe, such work, so basic to the conservation effort, is not at the forefront of botanical endeavour, now swept along by a universal and mindless ‘business model’ v

vi

Foreword by David Mabberley

requiring publication of papers of theoretical interest, in ‘high-impact’ journals, without which truly Neronian fiddling young scientists cannot advance their careers. It is particularly lamentable that this now obtains even in the countries which retain most of the greatest biodiversity. Nonetheless, like the guardians of human artefacts in museums, those of us, who are able, owe it to the coming generations to at least try to collect and describe what remains. How very refreshing then is this mighty book: a practical tool in tune with the concerns of everyman and everywoman rather than grant-giving bodies in the most ivory of academic towers! Roland Keller is in the grand tradition of pioneering naturalists, rightly awestruck by the majesty of the natural world, rather than obsessed with green matter in test-tubes. He follows modern exponents like Francis Hallé and Roelof Oldeman, these inspired by John Corner, who built on the work of Arthur Harry Church, paradoxically one of the least travelled of botanists (in body, though not mind). Ecological and conservation work in the tropics entails identifying large numbers of trees, which are often not in flower or fruit: consideration has to be given to vegetative characters that can be used in the field, characters often not even mentioned in floristic accounts or monographs. Roland Keller, with wide experience across the tropical world, has drawn together knowledge of such characters, most derived from personal experience and observation. His book is full of original information not to be found elsewhere: it is a thesaurus of botanical facts and is written in an accessible and effervescent style. Unlike most such guides or keys, this superbly illustrated book can be used anywhere in the tropics and provides in a straightforward three-step process identification to the level of families, which are now circumscribed according to molecular as well as morphological characters in the universally accepted scheme of the Angiosperm Phylogeny Group. Here that scheme is bolstered by the consideration of important and useful field characters—from overall architecture to bark characters (particularly revelatory in Lauraceae for example), to stipular scars, nature of any latex, odours of bruised tissues (and their taste!) besides the more familiar characters of indumentum and armature, for example. Within the family accounts, there is information concerning important economic plants with notes on the larger genera and, particularly helpfully, discussion of families readily confused and how to separate them. Particularly delightful are extracts from the author’s field-books, showing exquisite drawings of architecture and other features of plants as they are found in the forest. This book is unique—and a triumph. David Mabberley Wadham College University of Oxford Oxford, England Department of Natural Sciences Macquarie University Sydney, Australia

Foreword by Jitka Klimešová

As an ecologist, I like keys for determination of plants according to vegetative characters. When assessing species composition of a vegetation, majority of plants is not flowering but still they are residents of the community, interacting with other species and affecting ecosystem functions and I want to know them. As a morphologist, I know that not all characters useful for plant determination are commonly known to authors of identification keys. It concerns plant belowground organs in herbs or branching patterns in trees. I appreciate that book A Field Guide to Tropical Plant Families that is mainly devoted to trees. Moreover, branching and architectural models are integrated into the determination keys and plant descriptions. Although these traits are interesting per se, the main strength of this attempt is the treatment of a plant in its whole appearance. Currently, we know large part of the world flora only as a plant name and a flowering twig in herbarium, and such books as this from Roland Keller may encourage botanists to take whole plant into account when looking for determination characters and note it for others and future research. Personally, after two years of travel restrictions due to pandemic, I feel excited when imagining myself outside of camp in tropics observing a plant in its natural habitat, examining its branching, bark, exudates or odour of crushed leaves as is suggested in the section ‘It is better to observe the following characters in the field, before bringing the samples back to camp’. Jitka Klimešová Institute of Botany Czech Academy of Sciences Trebon, Czech Republic

vii

Acknowledgements

I wish to thank the following for their assistance, their friendship or their encouragements Daniel Atuany (Taman Nasional Manusela, Seram) Rupert Barneby † (New York Botanical Garden) Claude Edelin (CNRS, Montpellier) Peter Endress (Institut für Systematische und Evolutionäre Botanik, Zürich) Miguel Estaba (Boca Nichare, Venezuela) Jacques Fournet † (INRA, Pointe-à-Pitre) Pierre-André Loizeau, Nicolas Fumeaux, Fred Staufer, Laurent Gautier (Conservatoire Botanique de Genève) René Hebding † (Jardin Botanique Les Cèdres, Saint-Jean-Cap-Ferrat) Pak Ijun † (BIOTROP, Bogor) André Kostermans † (Kebun Raya, Herbarium Bogoriense) Anton Leeuwenberg † (Herbarium Vadense, Wageningen) Pierre Lombion (Morne-à-l’Eau, Guadeloupe) Germán López (Belize’s National Herbarium, Belmopan) Paul Maas (Rijksuniversiteit Utrecht) Francis Ng (Forest Research Institute Malaysia, Kepong) Jeanine Raharilala † (Jardin botanique de Tsimbazaza, Tananarive) Mamisoa Rapanoelina (WWF, Diego Suarez) Elio Sanoja (Universidad Nacional Experimental de Guayana, Upata) Nanu Sasidharan (Kerala Forest Research Institute, Peechi) Prachaya Srisanga (Queen Sirikit Botanical Garden, Chiang Mai) Sarawood Sungkaev (Jatujak University, Bangkok) Max van Balgooy (Universiteit Leiden) John van Thuyne (Van Thuyne Ridge Research Center, Chobe, Botswana) Jasper Zanco (Universidade do Sul de Santa Catarina) In addition, I thank Marie-Claude Gothuey, Claude Maillefer, Arnaud Pradervand, Caroline Sonnay, Daniel Schlaepfer, Laurent Rivier and Jean-Denis Briod for their practical assistance and moral support. I am indebted to the Corporación Venezolana de Guayana, the Queen Sirikit Botanical Garden and the Conservatoire Botanique de Genève, who generously helped me for logistical support or herbarium assistance. My heartfelt appreciation to Francis Hallé who was an invaluable guide throughout my years of research into the architecture of plants. ix

x

My humble thanks to David Mabberley for his flamboyant preface; his ‘Plant-Book’ was a faithful companion in the course of my last trips in Indonesia. Michael Heads has kindly provided many improvements to the concepts and the botanical terminology; thank you, Michael, for your wise suggestions. I was pleased to discover the work of Jitka Klimešová. Her book «Temperate Herbs—An Architectural Analysis» is particularly important as it should encourage more frequent exchanges between botanists specialising in tropical plants and those of mid-latitudes. I am very grateful to Rosemary Lees for her contribution to the translation of the French version into English.

Acknowledgements

Contents

Part I

User Manual and Architectural Concepts

1

Introduction . . . . . . . . . . . . . . . . . . . 1.1 Scope of the Book . . . . . . . . 1.2 Why Consider Families? . . . . 1.3 Photos or Drawings? . . . . . . . 1.4 Sources and Tests . . . . . . . . . 1.5 Structure of the Book . . . . . . References . . . . . . . . . . . . . . . . . . . . .

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3 3 3 4 4 4 5

2

How to Use This Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

7 10

3

Architectural Concepts and Key to the Models . . . 3.1 Concepts and Statements . . . . . . . . . . . . . . . . 3.2 Key to the Architectural Models . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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13 13 18 25

4

Key to the Groups of Families of Trees and Shrubs . . . . . . . .

29

5

Keys to Families of Trees and Shrubs . . . . . . . . . . . . . . . . . . .

37

6

Supplementary Keys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.1 Key A—Tribes and Main Genera of the Euphorbiaceae s.l. . . . . . . . . . . . . . . . . . . . . . 6.2 Key B—Tribes and Main Genera of the Malvaceae 6.3 Key C—Families of the Order Sapindales + Picramniaceae . . . . . . . . . . . . . . . . . . 6.4 Key D—Tribes and Main Genera of the Leguminosae . . . . . . . . . . . . . . . . . . . . . . . . . 6.5 Key E—Some Genera of the Rubiaceae . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Part II

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Identification Keys

7

Keys to the Groups of Families of Lianas and Woody Epiphytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

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Keys to Families of Lianas and Woody Epiphytes . . . . . . . . . 123 xi

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Contents

Part III

Description of the Families . . . . . . . . . . . . . . . .

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141 141 143 145 147 149 150 151 154 158 159 162 163 167 168 171

10 Bignoniaceae to Buxaceae . . . . . . . . . . . . . . . . . . . . 10.1 Bignoniaceae . . . . . . . . . . . . . . . . . . . . . . . . . 10.2 Bixaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.3 Boraginaceae (incl. Cordiaceae, Ehretiaceae) 10.4 Burseraceae . . . . . . . . . . . . . . . . . . . . . . . . . . 10.5 Buxaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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173 173 176 178 181 184 184

11 Cactaceae to Cycadaceae. . . . . . . . . . . . . . . . . . . . . . . . . . 11.1 Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.2 Canellaceae + Winteraceae . . . . . . . . . . . . . . . . . . . 11.3 Capparaceae (incl. Cleomaceae) . . . . . . . . . . . . . . . 11.4 Caricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.5 Casuarinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.6 Celastraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.7 Chloranthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.8 Chrysobalanaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 11.9 Clethraceae + Cyrillaceae . . . . . . . . . . . . . . . . . . . . 11.10 Clusiaceae + Calophyllaceae + Bonnetiaceae + Hypericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.11 Combretaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.12 Connaraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.13 Convolvulacaeae . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.14 Cornaceae + Nyssaceae . . . . . . . . . . . . . . . . . . . . . . 11.15 Corynocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.16 Cunoniaceae + Brunelliaceae . . . . . . . . . . . . . . . . . 11.17 Cycadaceae + Zamiaceae . . . . . . . . . . . . . . . . . . . . 11.18 Cyclanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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185 185 186 187 191 192 194 196 198 200

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202 205 209 212 215 216 217 218 219 220

9

Acanthaceae to Asteraceae . . . 9.1 Acanthaceae . . . . . . . . . 9.2 Achariaceae . . . . . . . . . . 9.3 Actinidiaceae . . . . . . . . . 9.4 Anacardiaceae . . . . . . . . 9.5 Ancistrocladaceae . . . . . 9.6 Anisophylleaceae. . . . . . 9.7 Annonaceae . . . . . . . . . . 9.8 Apocynaceae . . . . . . . . . 9.9 Aquifoliaceae . . . . . . . . 9.10 Araliaceae . . . . . . . . . . . 9.11 Araucariaceae . . . . . . . . 9.12 Arecaceae . . . . . . . . . . . 9.13 Aristolochiaceae . . . . . . 9.14 Asteraceae . . . . . . . . . . . References . . . . . . . . . . . . . . . . .

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xiii

12 Dichapetalaceae to Dipterocarpaceae . . . 12.1 Dichapetalaceae . . . . . . . . . . . . . . . 12.2 Dilleniaceae. . . . . . . . . . . . . . . . . . 12.3 Dioscoreaceae . . . . . . . . . . . . . . . . 12.4 Dipterocarpaceae . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . .

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221 221 222 224 225 228

13 Ebenaceae to Euphorbiaceae . . . . . 13.1 Ebenaceae . . . . . . . . . . . . . . . 13.2 Elaeocarpaceae . . . . . . . . . . . 13.3 Ericaceae . . . . . . . . . . . . . . . . 13.4 Erythropalaceae . . . . . . . . . . . 13.5 Erythroxylaceae . . . . . . . . . . . 13.6 Euphorbiaceae + Peraceae . . . References . . . . . . . . . . . . . . . . . . . . .

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229 229 231 234 237 239 241 248

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14 Fagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249 14.1 Fagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252 . . . . . .

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253 253 256 258 258 259

16 Hamamelidaceae to Humiriaceae . . . . . . . . . . 16.1 Hamamelidaceae (incl. Altingiaceae) . . . 16.2 Hernandiaceae . . . . . . . . . . . . . . . . . . . . 16.3 Huaceae . . . . . . . . . . . . . . . . . . . . . . . . 16.4 Humiriaceae . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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261 261 262 264 265 266

15 Gentianaceae to Goupiaceae . . 15.1 Gentianaceae . . . . . . . . . 15.2 Gnetaceae . . . . . . . . . . . 15.3 Goodeniaceae . . . . . . . . 15.4 Goupiaceae . . . . . . . . . . References . . . . . . . . . . . . . . . . .

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17 Icacinaceae s.l. to Irvingiaceae . . . . . . . . . . . . . . . . . . . . . . . . . 269 17.1 Icacinaceae s.l. + Cardiopteridaceae . . . . . . . . . . . . . . . . 269 17.2 Irvingiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271 18 Juglandaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275 18.1 Juglandaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276 19 Lamiaceae to Lythraceae . . . . . . . . . . . . . . . . . . . . . . . . . 19.1 Lamiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19.2 Lauraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19.3 Lecythidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19.4 Leguminosae—Introduction . . . . . . . . . . . . . . . . . . 19.5 Leguminosae–Cercideae–Cassieae–Detarieae– Caesalpinieae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19.6 Leguminosae–Mimoseae–Acacieae–Ingeae . . . . . . . 19.7 Leguminosae–Papilionoideae . . . . . . . . . . . . . . . . . . 19.8 Linaceae–Hugonioideae + Ixonanthaceae . . . . . . . .

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286 291 297 302

xiv

Contents

19.9 Loganiaceae + Gelsemiaceae . . . . . . . . . . . . . . . . . . . . . . 303 19.10 Lythraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 20 Magnoliaceae to Myrtaceae. . . 20.1 Magnoliaceae . . . . . . . . 20.2 Malpighiaceae . . . . . . . . 20.3 Malvaceae . . . . . . . . . . . 20.4 Marcgraviaceae . . . . . . . 20.5 Melastomataceae . . . . . . 20.6 Meliaceae . . . . . . . . . . . 20.7 Menispermaceae . . . . . . 20.8 Monimiaceae s.l. . . . . . . 20.9 Montiniaceae . . . . . . . . . 20.10 Moraceae. . . . . . . . . . . . 20.11 Moringaceae . . . . . . . . . 20.12 Muntingiaceae . . . . . . . . 20.13 Myricaceae . . . . . . . . . . 20.14 Myristicaceae . . . . . . . . 20.15 Myrtaceae . . . . . . . . . . . References . . . . . . . . . . . . . . . . .

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309 309 310 312 320 321 324 328 329 331 331 335 336 336 337 339 342

21 Nyctaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345 21.1 Nyctaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345 Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347 . . . . . . .

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349 349 351 353 355 357 358

23 Pandaceae to Putranjivaceae . . . . . . . . . . . . . 23.1 Pandaceae . . . . . . . . . . . . . . . . . . . . . . . 23.2 Pandanaceae . . . . . . . . . . . . . . . . . . . . . 23.3 Passifloraceae . . . . . . . . . . . . . . . . . . . . 23.4 Pentaphylacaceae. . . . . . . . . . . . . . . . . . 23.5 Phyllanthaceae. . . . . . . . . . . . . . . . . . . . 23.6 Physenaceae . . . . . . . . . . . . . . . . . . . . . 23.7 Phytolaccaceae (incl. Petiveriaceae) . . . 23.8 Picrodendraceae . . . . . . . . . . . . . . . . . . 23.9 Pinaceae . . . . . . . . . . . . . . . . . . . . . . . . 23.10 Piperaceae . . . . . . . . . . . . . . . . . . . . . . . 23.11 Pittosporaceae . . . . . . . . . . . . . . . . . . . . 23.12 Poaceae (Bambuseae) . . . . . . . . . . . . . . 23.13 Podocarpaceae . . . . . . . . . . . . . . . . . . . . 23.14 Polygalaceae . . . . . . . . . . . . . . . . . . . . . 23.15 Polygonaceae . . . . . . . . . . . . . . . . . . . .

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359 359 360 361 363 365 368 369 370 371 371 373 373 375 376 378

22 Ochnaceae to Oxalidaceae . . . . . . . . . . . . 22.1 Ochnaceae . . . . . . . . . . . . . . . . . . . 22.2 Olacaceae (incl. Schoepfiaceae) . . . 22.3 Oleaceae . . . . . . . . . . . . . . . . . . . . 22.4 Opiliaceae . . . . . . . . . . . . . . . . . . . 22.5 Oxalidaceae . . . . . . . . . . . . . . . . . . Reference . . . . . . . . . . . . . . . . . . . . . . . . .

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xv

23.16 Primulaceae . . . . . . 23.17 Proteaceae . . . . . . . 23.18 Putranjivaceae . . . . References . . . . . . . . . . . . .

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380 384 386 388

24 Rhabdodendraceae to Rutaceae . . . 24.1 Rhabdodendraceae. . . . . . . . . 24.2 Rhamnaceae . . . . . . . . . . . . . 24.3 Rhizophoraceae . . . . . . . . . . . 24.4 Rosaceae . . . . . . . . . . . . . . . . 24.5 Rubiaceae . . . . . . . . . . . . . . . 24.6 Rutaceae . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . .

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389 389 390 392 395 397 403 407

25 Sabiaceae to Symplocaceae . . . . . . . . . . . 25.1 Sabiaceae . . . . . . . . . . . . . . . . . . . 25.2 Salicaceae (incl. Lacistemataceae) . 25.3 Santalaceae + Loranthaceae . . . . . . 25.4 Sapindaceae. . . . . . . . . . . . . . . . . . 25.5 Sapotaceae . . . . . . . . . . . . . . . . . . 25.6 Schisandraceae . . . . . . . . . . . . . . . 25.7 Simaroubaceae + Picramniaceae . . 25.8 Smilacaceae. . . . . . . . . . . . . . . . . . 25.9 Solanaceae . . . . . . . . . . . . . . . . . . 25.10 Styracaceae . . . . . . . . . . . . . . . . . . 25.11 Surianaceae . . . . . . . . . . . . . . . . . . 25.12 Symplocaceae . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . .

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409 409 410 414 416 420 424 424 426 427 430 430 431 432

26 Tetramelaceae to Torricelliaceae . . 26.1 Tetramelaceae . . . . . . . . . . . . 26.2 Theaceae . . . . . . . . . . . . . . . . 26.3 Thymelaeaceae . . . . . . . . . . . 26.4 Torricelliaceae . . . . . . . . . . . . Reference . . . . . . . . . . . . . . . . . . . . .

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433 433 434 435 437 437

27 Ulmaceae s.l. to Urticaceae . . . . . . . 27.1 Ulmaceae + Cannabaceae . . . 27.2 Urticaceae . . . . . . . . . . . . . . . Reference . . . . . . . . . . . . . . . . . . . . .

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28 Verbenaceae to Vochysiaceae . . . . . 28.1 Verbenaceae + Callicarpa . . . 28.2 Violaceae . . . . . . . . . . . . . . . 28.3 Vitaceae . . . . . . . . . . . . . . . . 28.4 Vochysiaceae . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . .

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29 Zygophyllaceae s.l. (incl. Balanitaceae) . . . . . . . . . . . . . . . . . . 455 29.1 Zygophyllaceae (incl. Balanitaceae) . . . . . . . . . . . . . . . . 455

xvi

30 Herbaceous Versus Arborescent Habit in the Families and Orders of Flowering Plants—Evolutionary Trends . 30.1 A Diagrammatic Representation of Angiosperm Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30.2 References to Phylogenetic Studies . . . . . . . . . . . . . 30.3 Discussion and Hypothesis . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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List of Identified Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477

About the Author

Roland Keller is an independent botanist living in Switzerland. He was trained in the ‘Montpellier School’ of botanists, where he was initiated into the fields of plant architecture and botanical illustration, aspects which have remained important throughout his career. He has worked as a Research Associate at the Institute of Systematic Botany and Geobotany of the University of Lausanne and has carried out most of his fieldwork in the tropical forests of numerous countries, as well as in alpine areas. He has also been involved with surveys of biodiversity in Switzerland. This book is a completely revised version of his previous field guide: Identification of Tropical Woody Plants in the Absence of Flowers, which was published almost 20 years ago.

xvii

Part I User Manual and Architectural Concepts

1

Introduction

1.1

Scope of the Book

This guide covers the families of trees, shrubs and lianas of tropical lowland forests and savannas. Tropical terrestrial environments exist on four land masses: America, Africa, Asia and Australia as well as on many islands of Indomalesia and Oceania. Because only a few important families, i.e. those consisting of more than a hundred species, are restricted to one continental mass, the pantropical option has been retained. For example, the process of identification for a given member of the Moraceae family is similar whether one considers the whole tropical region or only Africa. There are around 425 families of Spermaphytes (or Seed plants), encompassing more than 350,000 species and 13,000 genera. Nearly 350 of these families have tropical representatives and 230 comprise woody tropical species. The numbers decrease to approximately 160 when considering families of more than twenty woody species. The present book helps to identify about 140 families, mostly consisting of more than thirty species, as well as some other, much smaller, but inevitable, families (Bixaceae, Moringaceae, Tetramelaceae, etc.). It is evident that large families are generally keyed out several times. For instance, Euphorbiaceae, holding the record of polymorphism among Angiosperms, are quoted in more than twenty-five items.

1.2

Why Consider Families?

The reason is that in herbaria, plants are always classified by families and in the absence of a reasonably priced and easily portable flora, the botanist working in the tropics will usually consult the local or national herbarium. Once the family of the specimen has been determined, it is relatively easy, using dried plant collections, to make an identification at the species level. However, without knowledge of the family, this process would be extremely tedious. I spent many hours in the herbarium of the Botanical Museum of Geneva trying to identify a specimen from Indonesia, which finally turned out to be a Proteaceae of the genus Helicia. This plant was unknown to my local guide in Manusela National park (Seram, Indonesia) although he was an excellent botanist. For both of us, the plant had an exotic appearance and we were not expecting to find, in Indonesia, a tree belonging to a family which exists mainly in South Africa and Australia.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 R. Keller, A Field Guide to Tropical Plant Families, https://doi.org/10.1007/978-3-031-05942-1_1

3

4

1.3

1

Introduction

Photos or Drawings?

Whereas some families can be identified at first sight, such as Palms or Arecaceae, most families are identified in an analytical way relying on a combination of several traits. As families are as a rule very variable, one needs to formalize their description with the help of graphical concepts. Photos are precise pictures of an individual, but these pictures can be misleading. A photo provides too much and not enough information at the same time; even though photos can be useful for identifying a species when one is familiar with the local flora, they cannot show certain traits which may be necessary for taxonomic identification at the family or genus level. However, carefully selected photos do help recognition, and make families …more familiar.

1.4

Sources and Tests

This book is a completely revised edition of the Field Guide published by the author about two decades ago [1, 2]. It is the result of a trial and error process based on innumerable tests. Data have been collected after studies in Belize, Northern Botswana, Southern Brazil, Cameroon, French Guiana, Guadeloupe, India (Kerala), Indonesia (Sumatra, Bali, Flores, Seram), Madagascar, Malaysia, Thailand, and Venezuela. Botanical gardens helped too (especially the Bogor Kebun Raya, Java). Tests have been carried out during each journey, directly in the field or soon after at the camp. Plant parts have been collected and helped as voucher specimens. Because of the lack of an existing field flora based on vegetative characters, herbaria were an invaluable tool for identification, at least at the family level (commonly at the genus level). Except a very few ones, all illustrations were made by the author. Digital photos were mainly taken during the period 2006–2019, which is why the majority of the pictures are from those more recently visited regions. However, older slides (period 1987–2004: Brazil, Cameroon, Guadeloupe, Guianas, Indonesia, Madagascar, Venezuela) have been scanned when contributing relevant information, although their quality is not as good. Apart from the matchless Mabberley’s Plant-Book [3, 4], many other field guides or floras provide valuable information: Aubréville [5], Da Silva [6–8], Gardner et al. [9], Gentry [10], Harden et al. [11, 12], Heywood [13], Hutchinson and Dalziel [14], LaFrankie [15], Lorenzi [16, 17], Ng [18], Palgrave [19], van Balgooy [20], Whitmore [21], and the Kew Tropical Handbook [22] helped too.

1.5

Structure of the Book

The book consists of three parts: I. User manual and botanical concepts II. Identification keys III. Description of the families The first part also gives explanations of architectural concepts, although architectural models are not strictly necessary for identification, they add very important information. In the second part (keys), morphological concepts are introduced using small illustrations alongside the keys. The third part is devoted to descriptions of the families, which are in alphabetical order. Each family is described in terms of its morphological traits (vegetative ones are emphasized) and currently accepted architectural models, geographical distribution, a list of important or distinctive genera, its economic or ethnobotanical importance, and tips indicating superficial resemblances with other families concerning vegetative traits. When appropriate, the family description ends with a short anecdote. Photos

References

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and line drawings complete these descriptions. This part also comprises ideas concerning angiosperm evolution. Of course, further improvements may be necessary. When considering more than fifty thousand arborescent species of the tropical belt, the keys may fail in some cases, although I have made tests with almost every species I found in the field. Floras and herbaria also helped greatly for constructing and testing the keys. Some families showing a narrow endemism or “strange” morphological adaptations are not included in this book, for example those characteristic of the distinctive flora of New Caledonia (especially on ultramafic rocks) and Southern Madagascar.

References 1. Keller, R. 1996. Identification of tropical woody plants in the absence of flowers—A field guide. 1st and 2nd ed. Birkhäuser, Basel: Berlin & Boston. 229 pp. 2. Keller, R. 2004. Identification of tropical woody plants in the absence of flowers—A field guide. 1st and 2nd ed. Birkhäuser, Basel, Berlin & Boston. 294 pp. 3. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3d ed., 1022 pp. Cambridge: Cambridge University Press. 4. Mabberley, D.J. 2017. Mabberley’s Plant-Book, 4th ed., 1102 pp. Cambridge: Cambridge University Press. 5. Aubréville, A. 1959. Flore forestière de la Côte d’Ivoire, Vol. I-II, 344–372. Nogent-sur-Marne: CTFT. 6. Da Silva Júnior, M.C. 2005. 100 Árvores do cerrado. Rede do Sementes do Cerrado, Brasilia. 278 pp. 7. Da Silva Júnior, M.C., and B.A. Da Silva Pereira. 2009. 100 Árvores do cerrado—matas de galeria. Brasilia. Rede do Sementes do Cerrado, 288 pp. 8. Da Silva Júnior, M.C., and M.R. Costa e Lima. 2010. 100 Árvores urbanas Brasilia. Brasilia: Rede do Sementes do Cerrado, 292 pp. 9. Gardner, S., P. Sidisunthorn, and V. Anusarnsunthorn. 2000. A field guide to forest trees of Northern Thailand, 545 pp. Bangkok: Kobfai Publising Project. 10. Gentry, A. H. 1996. A field guide to the families and genera of woody plants of Northwest South America. Chicago: The University of Chicago Press. xxiv + 896 pp. 11. Harden, G., B. McDonald, and J. Williams. 2006. Rainforest trees and shrubs: a field guide to their identification in Victoria, New South Wales and subtropical Queensland using vegetative features. Nambucca Heads: Gwen Harden Publishing. xxxiv + 266 pp. 12. Harden, G., B. McDonald, and J. Williams. 2007. Rainforest climbing plants: a field guide to their identification in Victoria, New South Wales and subtropical Queensland using vegetative features. Nambucca Heads: Gwen Harden Publishing. xvi + 176 pp. 13. Heywood, V.H. 1985. Flowering Plants of the World, 336 pp. London and Sydney: Croom Helm. 14. Hutchinson, J., and J.M. Dalziel. 1954. In Flora of West Tropical Africa, ed. R.W.J. Keay I (I-2), 2nd ed. London: Crown Agents. 15. LaFrankie, J.V. 2010. Trees of Tropical Asia: An Illustrated Guide to Diversity, 748 pp. Philippines: Black Tree Publication Inc. 16. Lorenzi, H. 1998. Árvores Brasileiras, vol. 1, 3rd ed., 384 pp. Nova Odessa: Instituto Plantarum da Estudos da Flora LTDA. 17. Lorenzi, H. 2000. Árvores Brasileiras, vol. 2, 3rd ed., 384 pp. Nova Odessa: Instituto Plantarum da Estudos da Flora LTDA. 18. Ng, F.S.P. 1978–1989. Tree Flora of Malaya, vols. 3–4, 340 pp/350 pp. London: Longman. 19. Palgrave, K.C., and M. Palgrave. 2003. Trees of Southern Africa, 960 pp. Cape Town: Random House Struik. 20. Van Balgooy, M.M.J. 1997. Malesian Seed Plants, vol. 1—Spot Characters, 154 pp. Leiden: Rijksherbarium. 21. Whitmore, T.C. 1972–1973. Tree Flora of Malaya, vols. 1–2, 472 pp./444 pp. London: Longman. 22. Utteridge, T., and G. Bramley. 2020. Tropical Plant Families Identification Handbook, 240 pp. Kew: The Boards of Trustees of the Royal Botanic Gardens.

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How to Use This Book

Explanations relative to keys Identification of a family involves three steps. The first step leads to one of 52 different groups of families. The second step leads to the family, except for the families of the order Sapindales. The third step helps to identify the different families of the Sapindales and to a more precise identification of large families such as Euphorbiaceae s.l., Malvaceae and Leguminosae. Concepts are not explicitly defined but are graphically presented by small illustrations in the margins. Keys cannot be used for the identification of saplings or very young plants which have not yet branched. Identification of architectural models are, in most cases, not necessary for taxonomic identification, however a basic knowledge of these models can help in many cases. When a liana is observed, the use of keys 53 to 57 will speed up identification. Precautions to be taken in the observation of certain characters It is better to observe the following characters in the field, before bringing the samples back to camp: • Exudates: These should be carefully searched for by making a tangential cut in the trunk and by sectioning leaves and young twigs. • Rhytidome characters: these concern the peripheral, dead tissues of trunks and branches. These characters are especially useful in the families of the order Myrtales and the EuphorbiaceaePhyllanthaceae. • Bark characters: these refer to the inner living bark, and have to be observed in branches or trunks sufficiently thick (the existence of network structures must be carefully searched for). • The monopodial or sympodial character of the branches and, if possible, of the trunk. • Presence of stems with two-ranked leaf arrangement is to be carefully searched for. The alternative is an overall spiral arrangement of the leaves. Remember that on main stems (trunks), leaf arrangement is usually spiral or opposite, very rarely two-ranked. • The odour of the crushed leaves and, for those species without latex, the taste (caution with Anacardiaceae and Euphorbiaceae!). • The mechanical properties of the twigs, such as flexibility and rigidity (e.g. for Salicaceae, Euphorbiaceae-Phyllanthaceae, Sapindaceae and Meliaceae).

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In the camp the following characters should be carefully examined (with a hand lens or against light, if necessary): • Presence or absence of stipules, these can be very reduced, especially for Aquifoliaceae (Ilex), Celastraceae, Dichapetalaceae, Euphorbiaceae, Lythraceae (Lagerstroemia), Malpighiaceae, Myrtaceae (Psidium), Ulmaceae, Violaceae and other taxa. Stipules can be modified into spines in many species of Acacia, Capparis and Ziziphus. • Presence or absence of glands and translucent dots on the leaves. • Indumentum characters such as the presence of stellate or appressed hairs. • A sunken or raised venation on the upper side of the blade. Taxonomy Taxonomy follows the systematic treatment of Mabberley’s Plant-Book [1] which follows in almost all cases the APG IV classification system upon which orders are named. The term ‘Dicotyledons’ (or ‘Dicots’) is kept here, but written between quotes, in spite that there are no longer a monophyletic group. Field of application All keys apply principally to Phanerogams of tropical lowland forests. The identification of the family to which a plant belongs can be guaranteed in a vast majority of cases covering the woody species in the tropical land masses of America, Africa and Asia. About one hundred and forty families are concerned. Arborescent Ferns are excluded. In order to specify more precisely the range of application, it must be pointed out that: • The key applies to ramified woody plants, however, some monocaulous trees (CORNER’s, HOLTTUM’s and CHAMBERLAIN’s models) are identifiable at family or genus level. • Certain very infrequently encountered or highly endemic families have been deliberately excluded due to lack of observation in the field. Conventions Syntax When an item consists of more than one proposition, these propositions are separated by a full stop. A full stop could be replaced by «and» without any change in the significance of the two propositions: thus «and» is implied; «or» is always non-exclusive. For example: a Spiral leaf arrangement. Leaves simple. b Leaf arrangement not spiral or leaves compound. Thus items a and b, each with two propositions, are complementary. Optional characters The propositions concerning optional taxonomical characters are printed between brackets, but if one of them is confirmed this increases the probability of an exact identification. Propositions that are not in brackets should be sufficient for identification.

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How to Use This Book

9

Architectural models are usually cited as optional characters and are written in brackets. Name of a family written in italics It occurs in a few cases, then the family is not described in Part III. Genera cited as examples The names of identified families are given in capital letters. For each land mass where a family is represented (with the relevant characters given by the key at that point) an effort has been made to cite at least one genus for each main land mass, as in the following case from the supplementary key B: MALVACEAE Theobroma (AM, cult.), Scaphopetalum (AF), Heritiera (AF, MA, AS) This means that Theobroma is exclusively American (but widely cultivated), Scaphopetalum is exclusively African and that Heritiera is paleotropical. Genera having pantropical distribution (i.e. at least in America, Africa and Asia) are mentioned at the end of the list, without land mass specification, as in the following case in group 5: ULMACEAE Gironniera (AS), Chaetachme (AF, MA), Celtis Taxonomic information can sometimes be insufficient to be able to cite an example of genus for one of the large land masses where the family might exist, e.g. in group 21: 8a Periderm remains green for several consecutive GUs. Young internodes angular or grooved. (AUBRÉVILLE). LAURACEAE e.g. Aniba (AM) and one should not deduce that Lauraceae conforming to the AUBRÉVILLE’s model are absent in Africa or Asia. If a genus is mentioned without being preceded by «e.g.», this genus alone is identified at this point in the key, as for example: THYMELAEACEAE Gonystylus (AS, OC) When a genus is mentioned in the key as an example for all the characters given, this does not imply that the characters exist in all species of the genus. Nevertheless, the characters stipulated are in general fairly typical of a genus. Abbreviations and acronyms ap. m. ab. = apical meristem abortions cult. = cultivated for its agronomic or timber value, hence possibly outside of its area of origin f. s. spp. = for some species ornam. = cultivated as ornamental, hence possibly outside of its area of origin L. = leaves or leaflets Rhy/Per = rhytidome (i.e. dead bark) or periderm GU = growth unit(s), equivalent to UE (unit of extension) p.p. = pro parte, i.e. for some species of a genus spp. = some species of a genus do exhibit the aforementioned trait s.t. = sometimes V II/III/IV = second / third / fourth level veinlets AF: tropical Africa; AM: tropical America; AS: tropical Asia (mainly Indomalesia); AU tropical Australia; EU: Europe, OC: Oceania; MA: Madagascar and neighbouring islands, NC: New Caledonia; NG: New Guinea; Paleo: paleotropical (AF, and AS or AU); WI: West Indies. -C, -E, -N, -W, -S for Central, East, North, West and South. Examples: AM-C, AF-W: Central America, West Africa.

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Explanations relative to the descriptions of the families Numbers given in the headings Numbers refer to groups given by the keys. For example, «ACANTHACEAE 36–37-38–56» means that the family Acanthaceae appears in groups 36, 37, 38 and 56. Some families, such as Aquifoliaceae are stereotyped and appear only in group 18 and are keyed out only one time, other families are more or less polymorphic, the record being established by the Euphorbiaceae (in eighteen groups) and are keyed out with 27 different possibilities! Common or vernacular names When a vernacular name does not come from the English language, it is set in quotes; e.g.: ‘buah rao’ (Dracontomelon dao) in Indonesia, ‘buah’ designates a fruit in Malayan language; ‘mureillo’ (Erisma uncinatum) in Venezuela; ‘oku baku’ (Okoubaka aubrevillei) in West Africa. Architectural models One or more examples of architectural models are cited for each family, the list is not exhaustive! Sources are usually not explicitly mentioned but refer mainly to [2], in some cases to [3, 4]. However in some cases information comes merely from the personal observations of the author. I accept that there may still be some errors. Uses The ‘Plant-Book’ [1, 5] helped to mention many agricultural, pharmaceutical, and other uses of plants. In addition, especially for Indomalesia, the monumental work of Burkill [6] provided interesting information. Other less known works were also useful: [7–11], among others. Acronyms When a picture or a drawing was made in an institution, the latter is cited by an acronym BG FIBV: Botanical garden of the Fundación Instituto Botánico de Venezuela, Caracas. BG Geneva: Jardin botanique de Genève. BG La Mortola: Giardini Botanici Hanbury, Ventimiglia, Italia. BG LC: Jardin botanique Les Cèdres, St Jean-Cap-Ferrat, France. BG Singapore: Singapore Botanic Gardens, Singapore. BG Sirikit: Botanical garden of the Queen Sirikit, Chiang Mai, Thailand. BG Tsimbazaza: Parc botanique et zoologique de Tsimbazaza, Antananarivo, Madagascar. CBG H: Herbier du Conservatoire Botanique de Genève. INRA: Institut National de Recherches Agronomiques, France. FRIM: Forest Research Institute of Malaya, Kuala Lumpur, Malaysia. Kebun Raya: Bogor botanical garden, Java. KFRI: Kerala Forest Research Institute, Thrissur, India. Sakaerat: Sakaerat Environmental Research Station, Thailand. Shipstern: Shipstern Conservation and Management Area, Belize.

References 1. 2. 3. 4.

Mabberley, D.J. 2017. Mabberley’s Plant-Book, 4th ed, 1102 pp. Cambridge: Cambridge University Press. Hallé, F., and R. Keller. 2019. Mais d’où viennent les plantes?, 180 pp. Actes Sud. Hallé, F. 2004. Architectures de plantes, 162 pp. Montpellier: JPC Editions. Hallé, F., R.A.A. Oldeman, and P.B. Tomlinson. 1978. Tropical Trees and Forests—An Architectural Analysis, 442 pp. Springer, Berlin. 5. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3rd ed, 1022 pp. Cambridge: Cambridge University Press.

References

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6. Burkill, I.H. 1935. A Dictionary of the Economic Products of the Malay Peninsula, vols. I and II, 2402 pp. Oxford: University Press. 7. Aubréville, A. 1959. Flore forestière de la Côte d’Ivoire, vols. I–II, 372 pp.–344 pp. Nogent s/Marne: CTFT. 8. Nair, K.K.N. 2000. Non-wood Forest Products of Kerala, 450 pp. Peechi: Kerala Forest Research Institute. 9. Roodt, V. 1998. Trees and Shrubs of the Okavango Delta, 224 pp. Shell Oil Botswana. 10. Schnee, L. 1984. Plantas comunes de Venezuela, 822 pp. Universidad Central de Venezuela: Colección ciencias biológicas VIII. 11. Soukup, J. 1970. Vocabulario de los nombres vulgares de la flora peruana, 384 pp. Lima: Colegio Salesiano.

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Architectural Concepts and Key to the Models

3.1

Concepts and Statements

Basic concepts Plant architecture is the science of stem arrangements in plants. A stem (or axis) consists of at least one internode bearing a lateral leaf and an apical meristem. Axes can be very different in their appearance. Interpreting the observed morphological structures is a challenging task in botany. The classic work of reference by Bell [1] is recommended for those who wish to study this subject in more detail. Illustrations of this topic are numbered according to paragraph numbers. Orientating Oneself on an Axis Orientation is obvious for the main stem of a treelet. However branches can bend in all directions and orientation becomes relative to a reference axis (‘upper’ or ‘lower’ parts have no meaning). To overcome this problem, basal (b) and distal (d) are more suitable terms: the basal/distal part of a lateral axis is that which is closer to / farther from the main axis (Fig. 3.1). Fig. 3.1 Basal (b) and distal (d) parts of an axis

Monopodial versus sympodial A monopodial axis is characterized by a linear series of modules, without terminal flowering. In the architectural sense, they are of two different types (Fig. 3.2a and b). ‘x’ marks the position of an aborted apical meristem. A sympodial axis is characterized by juxtaposition of modules with terminal flowering (inflorescences in red), (Fig. 3.2c).

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 R. Keller, A Field Guide to Tropical Plant Families, https://doi.org/10.1007/978-3-031-05942-1_3

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Fig. 3.2 (a) Monopodial axis, (b) monopodial axis with abortion of apical mersitems, (c) sympodial axis, (x = abortion of an apical meristem)

Growth rhythms Growth can be rhythmic, growth units, or GUs, are readily determined by the existence of shorter internodes or a periderm/young bark varying in aspect). Growth is said to be nearly continuous when the limits of GUs are not readily detectable (Fig. 3.3). However, the absence of morphological markers does not mean that rhythmic growth has not occurred, for instance Palm trees show a constant internode length although it is possible that during the dry season almost no stem elongation occurs, hence, in relation to time, growth is also rhythmic. Fig. 3.3 Rhythmic and continuous growth, (GU = growth unit)

Branching (or ramification), axillary buds and prophylls In Eudicotyledons, the prophylls are the two first leaves of a lateral axis, these are named a and b and are usually opposite (Fig. 3.4a). Prophylls are usually smaller than the ‘normal’ leaves of the individual. Axillary buds (b) are lateral axes too, but very small and with prophylls reduced to scaleleaves. Monocots have only one prophyll (Fig. 3.4b) and this one is adjacent to the main axis.

Fig. 3.4 Prophylls (a–b) in (a) Dicotyledons and (b) Monocotyledons

3.1 Concepts and Statements

15

Immediate or delayed branching Branching is immediate when a lateral axis develops in the course of development of the main axis (i.e. on the last GU formed). Branching is delayed when a lateral axis develops on an older GU of the main axis (usually the next to last GU) (Fig. 3.5). Fig. 3.5 Immediate (imm) and delayed (del) branching

Ramification (i.e. branching) and reiteration Branching (for French-speaking botanists: ‘ramification’), (Fig. 3.6, ram) is an automatic, genetically programmed, process. Reiteration (Fig. 3.6, reiter) looks very like branching but, when delayed in older plant trunks or branches, is an adaptative process. Consequently, when considering plant architecture one should distinguish between automatic processes and processes depending on external events. Fig. 3.6 (a) Branching, (b) reiteration (R)

Tropisms An axis is orthotropic (Fig. 3.7, ortho) when its growth tends to the vertical direction. An axis is plagiotropic (Fig. 3.7, plagio) when its growth tends to horizontality. A special type of plagiotropism, i.e. plagiotropism by apposition growth (Fig. 3.7, plagio appo), refers to a plagiotropic compound axis whose modules are at first plagiotropic (with one or two basal long internodes) then becoming orthotropic (with shorter internodes), see further AUBRÉVILLE’s and FAGERLIND’s models.

Fig. 3.7 Orthotropism, plagiotropism and plagiotropism by apposition growth

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Architectural Concepts and Key to the Models

Reorientation of a main axis In some cases an axis which is at first orthotropic may bend abruptly to become plagiotropic, see MANGENOT’s model), this phenomenon occurs during primary growth. Another case of reorientation occurs for axes being at first plagiotropic then becoming orthotropic, this phenomenon occurs in secondary growth (see TROLL’s model further) (Fig. 3.8).

Fig. 3.8 (o-p) Axis becoming plagiotrop in the course of primary growth, (p-o) axis becoming orthotropic in the course of secondary growth

Caution: When a new trunk leader develops by delayed branching without going through an early plagiotropic stage, one cannot speak of reorientation. This helps to define a ‘delayed’ KORIBA’s model which is in fact not a TROLL’s model (e.g. Butea monosperma, Olea salicifolia). Trunk branching, growth and tropisms, up and down processes A lateral axis developing on a trunk can be orthotropic in two ways: by a distal and delayed branching, a process named here orthotropic-up (Fig. 3.9, ortho-up), or by a more or less basal and immediate branching, this process is named here orthotropic-down (Fig. 3.9, ortho-down). For orthotropic-down AUBRÉVILLE’s models, at the beginning of their development, the first GUs of lateral branches are orthotropic, plagiotropy becoming established later in a global way for the whole branch. Fig. 3.9 Orientation of lateral branches in relation with the growth unit

3.1 Concepts and Statements

17

A lateral axis developing on a trunk has only one way to become plagiotropic: by distal branching of the next to last GU, i.e. plagiotropic-up process (Fig. 3.9, plagio-up). Basal immediate branching almost never produces plagiotropic axes, hence the plagiotropic-down process (Fig. 3.9, plagiodown) is against nature! However see Phyllanthus juglandifolius in the description of Phyllanthaceae. A special case of branching It is worth mentioning the peculiar case of the NOZERAN’s model exemplified by the apex of an individual of Theobroma cacao (Fig. 3.10). One can see that when plagiotropic A2 develop, the trunk apex GUn continues to grow but only for a short time and its apical meristem soon aborts (x). Growth will certainly continue by means of a delayed axis which will develop from an axil of GU(n-1), a process leading to a NOZERAN’s model (see Fig. 9b in the presentation of the architectural models). Hence, NOZERAN’s model appears as a MASSART’s model which goes wrong! This “technical hitch” may explain also the rarity of the model.

Fig. 3.10 Development of lateral branches in a young trunk of Theobroma cacao

Preliminary remarks to architectural models For basic knowledge about architecture, the reader may consult Bathélémy & Caraglio [2], Edelin [3], Hallé [4], Hallé et al. [5], Heuret et al. [6], Isnard et al. [7], Millet [8]. Plant architecture is a science of ‘axis arrangement’ and, practically, only vegetative axes and location of inflorescences are considered. However, architecture can also be applied to inflorescences, see Endress [9] for a review of the subject. Treating with herbs, architectural analysis becomes tricky (one goes a bit out of the paradigm because it becomes difficult to tell apart «vegetative» axes and inflorescence axes). See Klimešová [10] for an architectural approach adapted to herbs. The student must not confuse ‘architectural models’ with architectural analysis. Models are concepts while architectural analysis is the study of branching and reiteration processes. Models are presented in this book because they are taxonomically sound, and the author is convinced by what has become, not merely an artificial classification of plant structures, but a representation of essentially different growth processes. Architectural models do not follow a clear phylogenetic sequence. Although the same model is frequently found in closely related species, it is not unusual for an “exotic” model to be present in a group where a different model is the norm, e.g. MANGENOT’s model in the RAUH-MASSART continuum of Pinaceae and SCHOUTE’s model in Arecaceae.

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Architectural Concepts and Key to the Models

Another striking case can be found in Averrhoa (Oxalidaceae) where there are only two species but they have completely different architecture Ch. 22.5, [11], despite resembling each other in many other ways. Similarly, six different models of architecture can be found within the group Hibiscus [11]. Hence it seems more logical to consider the models as resurgences, rather than a gradual transformation in a phylogenetic series. Of course, each model may have appeared at a particular time during the evolution of a certain plant group. For example, KORIBA’s model, quite frequent in ‘Dicotyledons’, is absent in Gymnosperms and Monocotyledons. However, where and when this model arose during the evolution of the ‘Dicotyledons’ remains an unanswered question. As described later in this book, it is possible that the origin of TROLL’s model is to be found during the evolution of the Magnoliales on the Laurasian landmass about 100 Myr ago. Architectural models are based on specific combinations of characters. Not all character combinations are realized. Nature has rules that logic alone cannot discover. See, for instance, item 19c for description of a hypothetical model. As a warning, let finally expose the following statements: • Exact and stable definitions of models still do not exist, and will never exist, because they are varying among authors by subtle differences in primacy of characters, interpretation of processes, etc. • Limits of models can be fuzzy, especially between the pairs LEEUWENBERG-KORIBA, RAUHMASSART and CHAMPAGNAT-OLDEMAN for which continua represent better the reality. • Architectural models are more easily identifiable in young trees, before reiteration occurs. Reiteration is a duplication process of the whole model or a subset of it (e.g. branches reiterate to give other branches, but not other trunks). Some tree species do not reiterate (unitary trees, e.g. Cecropia spp.), but most of them do. • Architectural analysis may be relatively simple, for example when treating with “monopodial trees” of equatorial forests. The more the climate is changeable (long dry seasons, alternation of cold and hot seasons, etc.), the more interpretations (model diagnoses) become problematic. • Architectural analysis becomes very complicated for sympodial adult trees where the original model is recognizable only in the numerous peripheral small structures, but not in the whole tree. In fact the tree will develop a trunk appearing to be monopodial, but being anatomically sympodial (see Ch. 15.4: Goupia glabra). In order to describe this behaviour accurately, new concepts have been established, see [2, 3]. Nevertheless, a pagoda tree (Terminalia catappa), a mangosteen (Garcinia mangostana) and a ‘saman’ (Samanea saman) differ sufficiently enough in their structure to maintain architectural concepts.

3.2

Key to the Architectural Models

Models are illustrated on Figs. 3.11, 3.12, 3.13 and 3.14, and are numbered according to item numbers in the following key. Caution! KORIBA’s, RAUH’s and TROLL’s models, each have two types. 1a Plant monocaulous, its trunk monopodial or sympodial (see 2a, 2b, and 2c). 2 2a Trunk monopodial, lateral flowering. (Fig. 3.11). CORNER 2b Trunk monopodial, terminal flowering, (plant hapaxanthic). (Fig. 3.11). HOLTTUM 2c Trunk sympodial, terminal flowering. (Fig. 3.11). CHAMBERLAIN

3.2 Key to the Architectural Models

19

1b Trunk polycaulous (plant appearing branched at first sight). 3 3a Branching by dichotomy (trichotomy) of the terminal bud., frequent in plants living during the Mesozoic Era, actually a relict process). (Fig. 3.11). SCHOUTE 3b Branching lateral (i.e. monopodial or sympodial). 4 4a Trunk or main stems (A1) with sympodial branching (possibly frequent reiteration on main axis). 5 5a Main stem (trunk) with basal branching (usually palms or herbs with subterranean shoots and adventitious roots). Various aerial stems present or only one stem at a time. (Fig. 3.11). TOMLINSON 5b Main stem (trunk) with distal branching (s.t. basal reiteration). 6 6a Development of branches by immediate branching of the trunk and abortion of trunk apical meristem. 7 7a Flowering terminal (s.t. lateral). One or several lateral axes become(s) erect. 8 8a All lateral axes equivalent and becoming erect. (Fig. 3.11). LEEUWENBERG 8b Only one lateral axis becomes erect, the other oblique. (Fig. 3.11). KORIBA 7b Distal lateral axes stay plagiotropic. (New trunk leader with delayed development). 9 9a Lateral axes sympodial. (Fig. 3.12). PRÉVOST 9b Lateral axes monopodial. (Fig. 3.12). NOZERAN 6b Trunk modules with delayed branching (usually frequent reiteration starting from branches or trunks). 10 10a Branching is dominant in younger stages (reiteration will occur when the flowering stage is reached). 11 11a All axes orthotropic. 12 12a All lateral axes become erect (architecture somewhat unbalanced by lack of synchronisation between branches of the same storey). (Fig. 3.11). LEEUWENBERG 12b Only one lateral axis becomes erect, the other remains oblique. Caution: this architecture looks like the TROLL’s model but the new trunk leaders are not plagiotropic at their early stages (architecture somewhat unbalanced). (Fig. 3.11). KORIBA 11b ‘Mixed’ axes, i.e. in part plagiotropic and in part orthotropic. 13 13a Distal lateral branches at first plagiotropic then becoming erect during secondary growth (branching usually sympodial) or trunk distally plagiotropic but becoming erect during development (branching monopodial) or something in between. (Flowering usually lateral). (Fig. 3.12). TROLL 13b Trunk module at first orthotropic then bending abruptly to become plagiotropic. (Fig. 3.12). MANGENOT 10b Plant undergoing frequent reiterations on their main axes. 14 14a Decumbent shrub or tree (reiteration basal). (Fig. 3.12). CHAMPAGNAT 14b Liana with lateral branches (reiteration apical). (Fig. 3.12). OLDEMAN

20

3

Architectural Concepts and Key to the Models

4b Trunk (first order axis, A1) of the young plant with monopodial branching (reiteration appears in later stages). 15 15a Trunk with rhythmic branching. 16 16a Branches with monopodial ramification (second order axis, A2). 17 17a A2 plagiotropic, with lateral flowering (when A3 present, then flowering lateral or terminal). (Fig. 3.13). MASSART ‘down’ (if ‘up’, see NOZERAN) 17b A2 orthotropic, with lateral flowering (when A3 present, then flowering lateral or terminal). (Fig. 3.13). RAUH ‘up’ and ‘down’ 17c A2 orthotropic, with terminal flowering (when A3 present, then flowering terminal). (Fig. 3.13). SCARRONE 16b Branches with sympodial ramification (A2). 18 18a A2 plagiotropic by apposition, flowering lateral. (Fig. 3.13). AUBRÉVILLE 18b A2 plagiotropic by apposition, flowering terminal. (Fig. 3.13). FAGERLIND 15b Trunk (A1) with continuous branching. 19 19a Branches (A2) orthotropic. 20 20a Basal branches orthotropic, as vigorous as the main stem (herb or shrub). (Fig. 3.14). ACOSTA 20b Branches (A2) orthotropic, weaker than trunk, hierarchy of axes stronger. (Tree or shrub). 21 21a Branches with lateral flowering (when A3 present, flowering is lateral or terminal). (Fig. 3.14). ATTIMS 21b Branches with terminal flowering (when A3 present, flowering is terminal). (Fig. 3.14). STONE 19b Branches (A2) plagiotropic, but not by apposition. 22 22a Branches with lateral flowering (when A3 present, flowering is lateral or terminal). (Fig. 3.14). ROUX 22b Branches with terminal flowering (when A3 present, flowering is terminal). Branch modules not apically orthotropic. (Fig. 3.14). PETIT 19c Branches (A2) plagiotropic by apposition (hence branch modules apically orthotropic). Flowering lateral. (Fig. 3.14). Hypothetical model 15c Main axis (trunk) plagiotropic and usually subterranean. Secondary axes aerial and orthotropic, ramified or not. (Herb or shrub). (Fig. 3.14). BELL

3.2 Key to the Architectural Models

21

Fig. 3.11 Architectural models: CORNER, HOLTTUM, CHAMBERLAIN, SCHOUTE, TOMLINSON, LEEUWENBERG and KORIBA

22

3

Architectural Concepts and Key to the Models

Fig. 3.12 Architectural models: CHAMPAGNAT, OLDEMAN, MANGENOT, TROLL, PRÉVOST and NOZERAN

3.2 Key to the Architectural Models

Fig. 3.13 Architectural models: MASSART, RAUH, SCARRONE, AUBRÉVILLE and FAGERLIND

23

24

3

Architectural Concepts and Key to the Models

Fig. 3.14 Architectural models: ACOSTA, ATTIMS, STONE, ROUX, PETIT, BELL and hypothetical

References

25

References 1. Bell, A.D. 2008. Plant Form: An Illustrated Guide to Flowering Plant Morphology, 432 pp. Portland: Timber Press. 2. Barthélémy, D., and Y. Caraglio. 2007. Plant architecture: A dynamic, multilevel and comprehensive approach to plant form, structure and ontogeny. Annals of Botany 99: 375–407. 3. Edelin, C. 1991. Nouvelles données sur l’architecture des arbres sympodiaux: le concept de plan d’organisation. In: L’Arbre—Biologie et Développement, ed. C. Edelin, 127–154. Naturalia Monspeliensia, numéro hors série. 4. Hallé, F. 2004. Architectures de plantes, 162 pp. JPC Edition. 5. Hallé, F., R.A.A. Oldeman, and P.B. Tomlinson. 1978. Tropical trees and forests—An architectural analysis, 442 pp. Berlin: Springer. 6. Heuret, P., E. Nicolini, C. Edelin, and J.-C. Roggy. 2003. Approche architecturale pour l’étude des arbres de forêt tropicale humide guyanaise. Revue Forestière Française. Numéro spécial: 158–178. 7. Isnard, S., J. Prosperi, S. Wanke, S.T. Wagner, M.-S. Samain, S. Trueba, L. Frenzke, C. Neinhuis, and N. P. Rowek. 2012. Growth form evolution in Piperales and its relevance for understanding angiosperm diversification: An integrative approach combining plant architecture, anatomy, and biomechanics. International Journal of Plant Sciences 173: 610–639. 8. Millet, J. 2012. L’architecture des arbres des régions tempérées, 397 pp. Québec: Editions Multimondes. 9. Endress, P.K. 2010. Disentangling confusions in inflorescence morphology: Patterns and diversity of reproductive shoot ramification in angiosperms. Journal of Systematics and Evolution 48: 225–239. 10. Klimešová, J. 2018. Temperate herbes: an architectural analysis, 274 pp. Praha: ACADEMIA. 11. Hallé, F., and R. Keller. 2019. Mais d’où viennent les plantes? Actes Sud. 180 pp.

Part II Identification Keys

4

Key to the Groups of Families of Trees and Shrubs

See also groups 53 to 57 if lianescent in habit. Unillustrated families written in italics. 1a Venation parallel or indistinct, axillary buds infrequent or absent. Petiolar base decurrent on the stem (Gymnosperms). Prophyll appressed to the stem and sheathing leaf base (Fig. 4.1a), (Monocotyledons). Group 1: ARACEAE, ARAUCARIACEAE, ARECACEAE, ASPARAGACEAE, CUPRESSACEAE, CYCADACEAE+ZAMIACEAE, CYCLANTHACEAE, PANDANACEAE, PINACEAE, POACEAE - BAMBUSEAE, PODOCARPACEAE 1b Venation reticulate or indistinct, not parallel, presence of buds, prophylls a−b lateral, (Fig. 4.1b). DICOTYLEDONS and GYMNOSPERMS pp. 2 2a Leaves simple (entire, toothed or lobate) (For 2b, see p. 35). 3 3a Whitish, opalescent or coloured latex, or gum or resin in bark, stems or leaves. S.t. inner bark turning reddish and with an astringent taste. Late exudation of gum, which could happen when bark is wounded, is not to be considered. 4 4a Leaves alternate (i.e. spiral or two-ranked). 5 5a Leaves stipulate (Fig. 4.1c). Group 2: BIXACEAE, CHRYSOBALANACEAE, CALOPHYLLACEAE, DIPTEROCARPACEAE, EUPHORBIACEAE, LEGUMINOSAE, MORACEAE, PHYLLANTHACEAE, SAPOTACEAE, URTICACEAE 5b Leaves not stipulate. Group 3: ANACARDIACEAE, APOCYNACEAE, BURSERACEAE, CALOPHYLLACEAE, CARICACEAE, ERYTHROPALACEAE, MYRISTICACEAE, PHYLLANTHACEAE, SAPOTACEAE 4b Leaves opposite or whorled (Fig. 4.1d). Group 4: APOCYNACEAE, CELASTRACEAE, CLUSIACEAE s.l., EUPHORBIACEAE, MALPIGHIACEAE, MORACEAE, MYRTACEAE, SAPOTACEAE 3b Such an exudate absent in all plant parts (leaves, stems and bark). 6 6a Leaves alternate (For 6b, see p. 33). 7 7a Presence of twigs with two-ranked leaf arrangement (Fig. 4.1e) (For 7b, see p. 31). 8 8a Leaves stipulate. 9 9a Young stems with annular stipular scars (Fig. 4.1f). © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 R. Keller, A Field Guide to Tropical Plant Families, https://doi.org/10.1007/978-3-031-05942-1_4

29

30

4

Key to the Groups of Families of Trees and Shrubs

Group 5: CHRYSOBALANACEAE, DIPTEROCARPACEAE, HAMAMELIDACEAE, IRVINGIACEAE, MAGNOLIACEAE, MALVACEAE, POLYGONACEAE, SARCOLAENACEAE, ULMACEAE s.l., VITACEAE 9b Stipular scars not annular.

10

Fig. 4.1

10a Leaf venation palmate or tripliveined (Fig. 4.2a). Group 6: EUPHORBIACEAE, HAMAMELIDACEAE, LEGUMINOSAE, MALVACEAE, MUNTINGIACEAE, PASSIFLORACEAE, RHAMNACEAE, SALICACEAE, ULMACEAE s.l., URTICACEAE, VITACEAE 10b Leaf venation pinnate or supra-tripliveined (Fig. 4.2b). 11a Presence of leaves with glands.

11

Group 7: CHRYSOBALANACEAE, DICHAPETALACEAE, DIPTEROCARPACEAE, PASSIFLORACEAE, PHYLLANTHACEAE, ROSACEAE, SALICACEAE 11b Leaves without glands. 12a Leaf margin entire (Fig. 4.2c).

12

Group 8: CAPPARACEAE, CELASTRACEAE, CHRYSOBALANACEAE, DICHAPETALACEAE, DIPTEROCARPACEAE, ERYTHROXYLACEAE, EUPHORBIACEAE, FAGACEAE, HAMAMELIDACEAE, HUACEAE, LEGUMINOSAE, MALVACEAE, OCHNACEAE, PANDACEAE, PHYLLANTHACEAE, PUTRANJIVACEAE, RHAMNACEAE, SALICACEAE, ULMACEAE s.l., VIOLACEAE 12b Leaf margin toothed (th), crenulate (cr) or lobate (Fig. 4.2d). Group 9: APHLOIACEAE, FAGACEAE, GOUPIACEAE, HUMIRIACEAE, LACISTEMATACEAE, LECYTHIDACEAE, LEGUMINOSAE, OCHNACEAE, PUTRANJIVACEAE, RHAMNACEAE, SALICACEAE, VIOLACEAE 8b Leaves not stipulate (careful observation with a hand lens!). 13 13a Inner bark with a network of fibres (bark difficult to tear and detaching in strips, Fig. 4.2e). Group 10: ANNONACEAE, LECYTHIDACEAE, THYMELAEACEAE 13b Inner bark different.

14

Group 11: CALOPHYLLACEAE, EBENACEAE, POLYGALACEAE, PRIMULACEAE, ROSACEAE, SIMAROUBACEAE, SOLANACEAE 14a Stems or leaves with glands, translucent dots or dark dots (glandular trichomes), (Fig. 4.2f). (Caution! epiphytic colonies of microorganisms look like trichomes). 14b No glands, nor translucent dots or glandular trichomes. 15 15a Pubescent stem apices or buds, or indumentum of scaly hairs (hand lens!).

4

Key to the Groups of Families of Trees and Shrubs

31

Group 12: ARISTOLOCHIACEAE, CONNARACEAE, EBENACEAE, ERICACEAE, ERYTHROPALACEAE, ICACINACEAE s.l., LAURACEAE, MELIACEAE, MYRISTICACEAE, OLACACEAE, PENTAPHYLACACEAE, PERACEAE, SANTALACEAE, SOLANACEAE, STYRACACEAE, THEACEAE 15b Apices and buds glabrous (hand lens!). Group 13: ANISOPHYLLEACEAE, CANELLACEAE, CELASTRACEAE, CHRYSOBALANACEAE, CLUSIACEAE, CORNACEAE, ERICACEAE, ERYTHROPALACEAE, HUMIRIACEAE, ICACINACEAE, IXONANTHACEAE, LECYTHIDACEAE MARCGRAVIACEAE, OLACACEAE, OPILIACEAE, PIPERACEAE, PUTRANJIVACEAE, SALICACEAE

Fig. 4.2

7b All stems with spiral leaf arrangement (Fig. 4.3a). 16 16a Leaves stipulate (hand lens!). 17 17a Young stems with annular stipular scars or petiolar base sheathing the stem (Fig. 4.3b). Group 14: DILLENIACEAE, EUPHORBIACEAE, MAGNOLIACEAE, MALVACEAE, POLYGONACEAE, URTICACEAE, VITACEAE 17b Scars not annular and petiolar bases different. 18a Inner bark with a network of fibres.

18

Group 15: ACHARIACEAE, BIXACEAE, DIPTEROCARPACEAE, ELAEOCARPACEAE, LECYTHIDACEAE, MALVACEAE, ROSACEAE, URTICACEAE 18b Inner bark different. 19a Petiole distally swollen into a pulvinus (Fig. 4.3c).

19

Group 16: ACHARIACEAE, CAPPARACEAE, CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, MEDUSANDRACEAE 19b Petiole different. 20a Presence of leaves with glands (Fig. 4.3d).

20

Group 17: EUPHORBIACEAE, MALPIGHIACEAE, PASSIFLORACEAE, RHAMNACEAE, SALICACEAE 20b L. without glands (s.t. dark dots, i.e. ‘glandular’ trichomes). Group 18: ACHARIACEAE, AQUIFOLIACEAE, CELASTRACEAE, CORYNOCARPACEAE, DIPTEROCARPACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, FAGACEAE, IXONANTHACEAE, LECYTHIDACEAE, LINACEAE, OCHNACEAE, RHAMNACEAE, SALICACEAE, VIOLACEAE 16b Leaves not stipulate. 21a Leaf venation palmate or tripliveined (Fig. 4.3e). 22a Petiole distally enlarged into a pulvinus.

21 22

32

4

Key to the Groups of Families of Trees and Shrubs

Group 19: ARALIACEAE, EUPHORBIACEAE, HAMAMELIDACEAE, MENISPERMACEAE 22b Petiole different. Group 20: ACHARIACEAE, ANISOPHYLLEACEAE, BORAGINACEAE, CARICACEAE, ERICACEAE, EUPHORBIACEAE, HERNANDIACEAE, LAURACEAE, PIPERACEAE, TETRAMELACEAE 21b Leaf venation pinnate or supra-tripliveined. (parallel V II: phyllodes bearing Acacia, lianescent ARACEAE climbing by means of their roots: Heteropsis, AM). 23 23a Presence of lateral or terminal (i.e. AUBRÉVILLE) short shoots with very short internodes (plant s.t. thorny or spiny) (Fig. 4.3f).

Fig. 4.3

Group 21: ASTERACEAE, BIGNONIACEAE, BORAGINACEAE, CACTACEAE, CAPPARACEAE, CELASTRACEAE, COMBRETACEAE, CONNARACEAE, LAURACEAE, OLACACEAE, RUTACEAE, SOLANACEAE, SIMAROUBACEAE, SURIANACEAE 23b No such twigs and plant not spiny. 24 24a Leaves with glands, or translucent dots or dark dots (Fig. 4.4a), (glandular trichomes), or waxy indumentum or stellate/scaly hairs. 25 25a Glands or stellate/indumentum of minute scaly hairs (Fig. 4.4b). Group 22: CAPPARACEAE, COMBRETACEAE, EUPHORBIACEAE, HUMIRIACEAE, PASSIFLORACEAE, SALICACEAE, STYRACACEAE 25b Leaves with translucent or dark dots (The last not to be confused with epiphytic colonies of microorganisms!) or underside of leaves with waxy indumentum. Group 23: ASTERACEAE, LAURACEAE, MYRTACEAE, PENTAPHYLACACEAE, PRIMULACEAE, RHABDODENDRACEAE, RUTACEAE, SAPINDACEAE, SCHISANDRACEAE, VERBENACEAE, WINTERACEAE 24b Leaves without glands, translucent/dark dots or stellate hairs. 26a Branches plagiotropic by apposition (Fig. 4.4c).

26

Group 24: ACTINIDIACEAE, ANACARDIACEAE, BONNETIACEAE, BORAGINACEAE, COMBRETACEAE, GOODENIACEAE, LAURACEAE, MELIACEAE, RUTACEAE, SAPOTACEAE 26b Branches different. 27 27a Inner bark fibrous, with a networked of fibres or external wood furrowed (Fig. 4.4d).

4

Key to the Groups of Families of Trees and Shrubs

33

Group 25: BORAGINACEAE, ICACINACEAE, LECYTHIDACEAE, PROTEACEAE, SABIACEAE, THYMELAEACEAE 27b Bark and stems different. 28 28a Petiolar base sheathing (s.t. stipuliform petiolar expansions) or base of petiole canaliculate or grooved (Fig. 4.4e). Group 26: ARALIACEAE, DILLENIACEAE, TORRICELLIACEAE 28b Petiole different. 29a Petiole distally swollen or pulvinate.

29

Group 27: ACHARIACEAE, CAPPARACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, LEPIDOBOTRYACEAE, OXALIDACEAE, PHYLLANTHACEAE 29b Petiole different. 30a Veins II or III secant on lamina margin (Fig. 4.4f).

30

Group 28: ACTINIDIACEAE, ANACARDIACEAE, CLETHRACEAE, DILLENIACEAE 30b Venation not secant.

31

Fig. 4.4

31a Branches with growth strongly rhythmic denoted by shorter internodes and sequences of growth units without abortion of apical meristems (Fig. 4.5a). Group 29: ANACARDIACEAE, BONNETIACEAE, CLETHRACEAE, CANELLACEAE, CYRILLACEAE, DAPHNIPHYLLACEAE, ERICACEAE, GOODENIACEAE, HAMAMELIDACEAE, IXONANTHACEAE, LAURACEAE, LECYTHIDACEAE, NYSSACEAE, PENTAPHYLACACEAE, PITTOSPORACEAE, PRIMULACEAE, SIMAROUBACEAE, THEACEAE 31b Branches without series of shorter internodes or branches with frequent ap. m. ab. (Fig. 4.5b), these denoted by discontinued pith. Group 30: BORAGINACEAE, CANELLACEAE, CAPPARACEAE, CELASTRACEAE, COMBRETACEAE, ERICACEAE, HUMIRIACEAE, ICACINACEAE, IXONANTHACEAE, LAURACEAE, LEGUMINOSAE (phyllodinous Acacia), MYRICACEAE, OPILIACEAE, PENTAPHYLACACEAE, PHYSENACEAE, PHYTOLACCACEAE, SABIACEAE, SALICACEAE, SANTALACEAE, SOLANACEAE, SYMPLOCACEAE, THEACEAE 6b Leaves opposite or whorled (Fig. 4.5c). 32a Leaves stipulate. 33a Leaf venation palmate or tripliveined.

32 33

34

4

Key to the Groups of Families of Trees and Shrubs

Group 31: CANNABACEAE, EUPHORBIACEAE, RHAMNACEAE, URTICACEAE 33b Leaf venation pinnate (Fig. 4.5d). 34 34a Stipular scars connecting petiolar bases (stipules interpetiolar) or stipules intrapetiolar (Fig. 4.5e). Group 32: CHLORANTHACEAE, CTENOLOPHONACEAE, CUNONIACEAE, ERYTHROXYLACEAE, EUPHORBIACEAE, LOGANIACEAE s.l., RHAMNACEAE, RHIZOPHORACEAE, RUBIACEAE, TRIGONIACEAE 34b Stipular scars not connecting the petiolar bases, relatively small and lateral. 35 35a Leaf margin entire. Group 33: CELASTRACEAE, CLUSIACEAE, LOGANIACEAE, LYTHRACEAE, MALPIGHIACEAE, PICRODENDRACEAE, RHAMNACEAE, RHIZOPHORACEAE, VIOLACEAE, VOCHYSIACEAE 35b Leaf margin toothed or crenulate. Group 34: CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, OCHNACEAE, RHIZOPHORACEAE, VIOLACEAE 32b Leaves not stipulate. 36a Nodes with interpetiolar ridge (Fig. 4.5f). 37a Three or five veins starting from leaf base or slightly above it.

36 37

Group 35: LAMIACEAE, LOGANIACEAE s.l., OLEACEAE, MELASTOMATACEAE, VERBENACEAE, VIBURNACEAE 37b Venation different.

38

Fig. 4.5

38a Nodes or base of internodes swollen (Fig. 4.1a). Group 36: ACANTHACEAE, AMARANTHACEAE, GNETACEAE, NYCTAGINACEAE 38b Nodes and base of internodes different. 39 39a Venation indistinct (V II not visible) or leaves like small scales or needles (Fig. 4.6b). Group 37: ACANTHACEAE, CASUARINACEAE, CUPRESSACEAE, GENTIANACEAE, LORANTHACEAE, LYTHRACEAE, MELASTOMATACEAE, PINACEAE 39b Leaves and venation different. Group 38: ACANTHACEAE, ASTERACEAE, CLUSIACEAE s.l., CRYPTERONIACEAE, GENTIANACEAE, LAMIACEAE, LOGANIACEAE s.l., LYTHRACEAE, VERBENACEAE 36b Nodes without interpetiolar ridge. 40a Three or five veins starting from leaf base or slightly above it (Fig. 4.6c).

40

4

Key to the Groups of Families of Trees and Shrubs

35

Group 39: BUXACEAE, GENTIANACEAE, LAMIACEAE, LAURACEAE, STOMATACEAE, MYRTACEAE, SAPINDACEAE, SCROPHULARIACEAE

MELA-

40b Venation different. 41a Leaves with small dots, peltate trichomes, stellate hairs or glands.

41

Group 40: ASTERACEAE, BIGNONIACEAE, COMBRETACEAE, EBENACEAE, MALPIGHIACEAE, MONIMIACEAE, MYRTACEAE, OLEACEAE, RUTACEAE, TRIMENIACEAE, VERBENACEAE 41b Leaves different or glabrous. 42 42a Bark peeling in the form of straw-like fibres or small scales (Fig. 4.6d). Group 41: CLUSIACEAE s.l., COMBRETACEAE, LYTHRACEAE, MELASTOMATACEAE, MYRTACEAE, NYCTAGINACEAE, ONAGRACEAE, VERBENACEAE 42b Bark different. 43 43a Leaves with intramarginal or fimbrial veins or venation parallel (Fig. 4.6e). Group 42: ARAUCARIACEAE, CALOPHYLLACEAE, MYRTACEAE, VOCHYSIACEAE 43b Leaves and venation different. 44a Petiole distally pulvinate (Fig. 4.6f).

44

Group 43: BIGNONIACEAE, LAMIACEAE 44b Petiole different. 45a Leaf margin entire.

45

Group 44: ANACARDIACEAE, BUXACEAE, ICACINACEAE, LAMIACEAE, LAURACEAE, MALPIGHIACEAE, MELASTOMATACEAE, MONIMIACEAE, MONTINIACEAE, MYRTACEAE, NYCTAGINACEAE, OLEACEAE, PICRODENDRACEAE, PROTEACEAE, SANTALACEAE, THYMELAEACEAE, VERBENACEAE

Fig. 4.6

45b Leaf margin toothed or crenulate. Group 45: CELASTRACEAE, LAMIACEAE, MONIMIACEAE, OLEACEAE, PROTEACEAE 2b Leaves compound (Fig. 4.7a). 46a Leaf arrangement alternate. 47a Leaves stipulate. 48a Leaves pinnate (Fig. 4.7b).

46 47 48

Group 46: BURSERACEAE, LEGUMINOSAE, ROSACEAE, RUTACEAE, SAPINDACEAE, SIMAROUBACEAE, STAPHYLEACEAE, VITACEAE 48b Leaves palmate or trifoliolate (Fig. 4.7c).

36

4

Key to the Groups of Families of Trees and Shrubs

Group 47: ARALIACEAE, BIXACEAE, CAPPARACEAE, CARYOCARACEAE, EUPHORBIACEAE, LEGUMINOSAE, MALVACEAE, PHYLLANTHACEAE, URTICACEAE, VITACEAE 47b Leaves not stipulate. 49a Leaves pinnate or bifoliolate (Fig. 4.7d).

49

Group 48: ARALIACEAE, BERBERIDACEAE, CONNARACEAE, JUGLANDACEAE, LEGUMINOSAE, MORINGACEAE, OXALIDACEAE, PICRAMNIACEAE, PROTEACEAE, SABIACEAE, SAPINDALES (MELIACEAE, RUTACEAE, etc.), ZYGOPHYLLACEAE 49b Leaves palmate or trifoliolate. Group 49: ARALIACEAE, BIGNONIACEAE, CARICACEAE, CONNARACEAE, OXALIDACEAE, PICRODENDRACEAE, RUTACEAE, SAPINDALES (SAPINDACEAE, etc.) 46b Leaf arrangement opposite or whorled. 50a Leaves stipulate.

50

Group 50: BRUNELLIACEAE, CARYOCARACEAE, CUNONIACEAE, LEGUMINOSAE, OCHNACEAE, STAPHYLEACEAE, ZYGOPHYLLACEAE 50b Leaves not stipulate. 51a Leaves pinnate with more than three leaflets/pinnae (Fig. 4.7e).

51

Group 51: BIGNONIACEAE, LAMIACEAE, MELIACEAE, OLEACEAE, RUTACEAE, SAPINDACEAE, VIBURNACEAE 51b Leaves palmate or trifoliolate. Group 52: BIGNONIACEAE, LAMIACEAE, PICRODENDRACEAE, RUTACEAE, SAPINDACEAE

Fig. 4.7

5

Keys to Families of Trees and Shrubs

Group 1—Foliar base sheathing the stem or its base decurrent (Fig. 5.1a); leaves simple or compound; venation indistinct or parallel. (Gymnosperms or Monocots). ARACEAE, ARAUCARIACEAE, ARECACEAE, ASPARAGACEAE, CUPRESSACEAE, CYCADACEAE+ZAMIACEAE, CYCLANTHACEAE, PANDANACEAE, PINACEAE, POACEAEBAMBUSEAE, PODOCARPACEAE 1a Leaves simple. 2a Alternate leaf arrangement in trunk and branches. 3a Leaves needle-like, in groups of 2, 3 or 5. (Fig. 5.1b)

2 3 PINACEAE Pinus (temperate AS, AM)

3b Leaves different. 4 4a Paleo. Leaf arrangement in three imbricate spirals (Fig. 5.1c). Large, stretched leaves. (Trunk bearing adventitious roots). CORNER, LEEUWENBERG, SCARRONE, STONE). PANDANACEAE Pandanus (AF, MA, AS, OC) 4b Leaf arrangement simply spiral or two-ranked. 5 5a All stems with spiral leaf arrangement or plant monocaulous. 6 6a Leaves long, with petiolar base sheathing the stem or leaf sheath tubular. (See 7a to 7c). 7a Ramified plant. Leaf lamina flat, with an amplexicaulous base (Dracaena). ASPARAGACEAE e.g. Yucca (AM), Dracaena (Paleo), Cordyline (AM, AS, AU) 7b Plant ramified bearing drooping adventitious roots (Fig. 5.1d) ARACEAE e.g. Heteropsis (AM-S) 7c Plant monocaulous. Leaves plicate (Fig. 5.1e). 8 8a Trunk erect, not ramified. (Leaves bifid). Leaves with tubular sheath. ARECACEAE e.g. Geonoma (AM), Johannesteijsmannia (Indomalesia) (e.g. sapling of young Coconut tree)

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 R. Keller, A Field Guide to Tropical Plant Families, https://doi.org/10.1007/978-3-031-05942-1_5

37

38

5

Keys to Families of Trees and Shrubs

8b AM. Plant weakly woody. Trunk prostrate or young stage of a climbing plant. Foliar sheath not tubular. CHAMBERLAIN, leaves bifid: e.g. Asplundia; CORNER, leaves simple: e.g. Ludovia. CYCLANTHACEAE e.g. Asplundia, Evodianthus, Ludovia (AM) 6b Foliar base decurrent on stem, not sheathing (Fig. 5.1f). plant ramified.

9

Fig. 5.1

9a Not in AF. Existence of shoots arranged in a same plane. Leaves coriaceous and acuminate. (MASSART, RAUH). ARAUCARIACEAE Araucaria (AM, NG, AU, OC, cult.) 9b (Highlands). RAUH. All branches with a spiral arrangement. (Leaf lamina not pungent, s.t. flat). PODOCARPACEAE e.g. Podocarpus 5b Presence of two-ranked leaf arrangement. (Shrub). 10 10a Twigs bearing very large and long leaves. Growth rhythmic. Twigs drooping, or erect at first, then becoming plagiotropic (MANGENOT Fig. 5.2a). ASPARAGACEAE Cordyline (AS) 10b Aerial stems without rhythmic growth. Stems swollen at nodes. Distinct serial buds. L. differentiated into lamina and sheath (Fig. 5.2b). POACEAE-BAMBUSOIDEAE e.g. Dendrocalamus, Gigantochloa (AS), Bambusa 2b Leaves opposite. 11a Not in AF. (MASSART). Leaves flat, with parallels veins

11 ARAUCARIACEAE Agathis (AS, NG, OC)

11b AU, OC. Leaves like scales (Fig. 5.2c).

CUPRESSACEAE Callitris (AU, New Caledonia)

1b Leaves compound. 12 12a Leaflets plicate (Fig. 5.2d). 13 13a Leaves pinnate. Leaf sheath tubular. (Leaves heterophyllous, those of the young stage simpler in form than leaves of adult stage. Plant not ramified or plant sprouting at its base (e.g. Bactris, Chrysalidocarpus). ARECACEAE e.g. Bactris, Euterpe, Roystonea (AM) Chrysalidocarpus (MA), Elaeis (AF, cult.), Veitchia (OC) 13b Leaves palmately compound. 14 14a Foliar base forming a cylindrical tubular sheath (Fig. 5.2e). Not ramified tree or plant sprouting at its base. (Branches aerial, e.g. Hyphaene, subtropical, Paleo). ARECACEAE e.g. Mauritia (AM), Borassus (AF, AS) Phoenix (EU, AF, AS), Livistona (AF, AS, AU)

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Keys to Families of Trees and Shrubs

14b AM. Foliar sheath not tubular.

39

CYCLANTHACEAE e.g. Carludovica (AM)

12b Divisions of leaves flat, not plicate, their base decurrent on the rachis (Fig. 5.2f). CYCADACEAE+ZAMIACEAE e.g. Zamia (AM), Ceratozamia (AM-C), Cycas (Paleo)

Fig. 5.2

Group 2—Presence of a white, opalescent or coloured exudate; alternate arrangement of leaves; leaves simple, stipulate. BIXACEAE, CHRYSOBALANACEAE, CALOPHYLLACEAE, DIPTEROCARPACEAE, EUPHORBIACEAE, LEGUMINOSAE, MORACEAE, PHYLLANTHACEAE, SAPOTACEAE, URTICACEAE 1a Petiole distally swollen (sometimes weakly) or pulvinate. (see 2a to 2d). (Fig. 5.3a). 2a Exudate white, reddish, brownish or opalescent. (S.t. leaves with glands). Very polymorphous family identifiable usually by its supple twigs and a young bark longitudinally fissured. EUPHORBIACEAE See key A, e.g. Elateriospermum (AS) 2b Exudate red, aqueous. Twigs lenticellate. Leaf margin entire. (TROLL, fluted trunk). LEGUMINOSAE e.g. Inocarpus fagifer (AS, OC) 2c AS. Sectioned bark producing a translucent gum (damar). Rhythmic growth (long and short internodes). Venation scalariform (Fig. 5.3b). DIPTEROCARPACEAE e.g. Vatica (AS) 2d Leaves entire, tripliveined (Fig. 5.3c). (Latex more or less colourless). URTICACEAE e.g. Coussapoa (AM) 1b Petiole distally neither swollen, nor pulvinate. 3 3a Stipules hood-like (Fig. 5.3d). 4 4a Exudate brownish or becoming so. Leaf margin entire or palmatilobate (Cecropia). Leaves tripliveined: Coussapoa, Pourouma. (Fimbrial vein: Pourouma). (Hemi-epiphyte: Coussapoa). URTICACEAE e.g. Coussapoa, Cecropia, Pourouma (AM) 4b Exudate white, whitish or pink. (Venation brochidodromous). MORACEAE e.g. Perebea, Pseudolmedia (AM), Treculia (AF), Ficus 3b Stipules different (small-sized or incompletely annular). 5 5a All stems with spiral arrangement of leaves (RAUH, AUBRÉVILLE). 6 6a Exudate orange-coloured, aqueous. Leaves simple (s.t. lobate) and without glands. Hairs simple (neither fasciculate, nor stellate), s.t. like minute circular scales (Bixa). Inner bark fibrous (Fig. 5.3e). BIXACEAE Amoreuxia, Bixa (AM, cult.), Cochlospermum

40

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Keys to Families of Trees and Shrubs

6b At least, at least one of these characters different. 7 7a Twigs lenticellate. Latex white, pink or opalescent, milky or somewhat viscous. Leaves without stellate hairs or glands. Stipules small or hood-like. Venation densely reticulate (Fig. 5.3f). (White latex and lobate leaves in Broussonetia sp., URTICACEAE).

Fig. 5.3

MORACEAE e.g. Artocarpus (AS), Ficus 7b Twigs not lenticellate. (See 8a to 8d). 8a Latex white. Stipules minute. Leaves without glands or stellate hairs. (Inner bark pink-orange). SAPOTACEAE e.g. Ecclinusa (AM), Mimusops, Palaquium (AS, OC) 8b Exudate white, whitish, reddish or opalescent. Quite supple twigs. (Sometimes lamina or petiole with glands). EUPHORBIACEAE e.g. Gymnanthes, Sebastiania (AM), Ostodes (AS) 8c AF. Exudate brownish. Small tree with stilt-roots (Fig. 5.4a). Petiole quite long. RAUH. PHYLLANTHACEAE Uapaca (AF, MA) 8d AM. Exudate yellowish or scarce. Leaf margin entire, without glands. (False stipules: Mahurea). CALOPHYLLACEAE Mahurea (AM) 5b Presence of two-ranked arrangement of leaves (Fig. 5.4b). (ROUX, TROLL). 9 9a Latex white, pink or opalescent, milky or somewhat viscous. Leaves without stellate hairs or glands. Stipules small or hood-like. Venation densely reticulate (Fig. 5.4c) (L. toothed: Clarisia, Sorocea, L. spiny: Streblus). MORACEAE e.g. Brosimum, Clarisia, Sorocea (AM), Artocarpus (AS), Streblus (MA, AS, OC) 9b Latex white (Fig. 5.4d). Venation not densely reticulate. L. toothed. Twigs supple. (Young bark with thin longitudinal slits). EUPHORBIACEAE e.g. Sebastiania (AM) 9c Inner bark pinkish producing a red, aqueous and astringent (Fig. 5.4e). Trunk or twigs lenticellate (Fig. 5.4f). CHRYSOBALANACEAE e.g. Licania (AM) DIPTEROCARPACEAE are part of this group because they are a source of oleoresins, but exudates are observable only in large trees which were artificially wounded.

Fig. 5.4

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Keys to Families of Trees and Shrubs

41

Group 3—Presence of a white, opalescent or coloured exudate; alternate arrangement of leaves; leaves simple, not stipulate. 9 ANACARDIACEAE, APOCYNACEAE, BURSERACEAE, CALOPHYLLACEAE, CARICACEAE, ERYTHROPALACEAE, MYRISTICACEAE, PHYLLANTHACEAE, SAPOTACEAE 1a Exudate red, reddish (Fig. 5.5a) or orange-coloured (s.t. no exudate but inner bark turning reddish) or bark astringent. 2 2a Bark astringent, not resinous. (MASSART). MYRISTICACEAE e.g. Virola (AM), Pycnanthus (AF), Myristica (Paleo, cult.) 2b Exudate neither astringent nor resinous. 3 3a Petiole distally swollen. ERYTHROPALACEAE Coula (AF) 3b Petiole not distally swollen. 4 4a ROUX’s model (Fig. 5.5b), twigs not lenticellate. SAPOTACEAE e.g. Chrysophyllum (AM) 4b AM. Exudate turning reddish. Twigs lenticellate. Venation not densely reticulate (Fig. 5.5c). (Fimbrial vein). (MASSART, RAUH). APOCYNACEAE Aspidosperma (AM) 1b Exudate neither red/reddish nor astringent. (See 5a to 5d). 5a Exudate in bark, or at least in leaves, viscous, white, beige, s.t. turning brown in the course of time. V II abruptly curved near lamina margin (Fig. 5.5d). ANACARDIACEAE e.g. Anacardium (AM, cult.), Mangifera (AS, cult.) 5b Translucent, aromatic, gum-resin. Leaves ‘unifoliolate’ with petiole distally pulvinate (Fig. 5.5e). BURSERACEAE Protium sp. (AM) 5c Exudate whitish. Small monocaulous or sparsely branched trees bearing palmate leaves. Large trees in equatorial Africa. CARICACEAE Cylicomorpha (AF), Carica, Jacaratia (AM, cult.) 5d Plant with other characters. 6 6a AM, AF. Latex whitish or beige in leaves, stems or bark, not viscous. Presence of twigs with two-ranked leaf arrangement. Stem periderm remaining green for several consecutive GUs. (ROUX, MANGENOT). ERYTHROPALACEAE e.g. Heisteria (AM), Coula (AF) 6b Plant different. (See 7a to 7c). 7 7a Twigs not lenticellate. 8 8a Latex white, viscous, seldom yellow or opalescent. Leaves without stellate hairs and without glands (s.t. numerous parallel V II, Fig. 5.5f). (RAUH, AUBRÉVILL E, ROUX, TROLL). Short twigs: Sideroxylon, hairs rust-coloured: Chrysophyllum, tiny stipules: Manilkara). SAPOTACEAE e.g. Sideroxylon (AM), Baillonella (AF), Chrysophyllum, Manilkara, Pouteria

Fig. 5.5

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Keys to Families of Trees and Shrubs

8b AM. Resin pale yellow (Fig. 5.6a). (Leaves with stellate hairs). CALOPHYLLACEAE (see CLUSIACEAE) Caraipa, Kielmeyera (AM) 7b Twigs lenticellate (Fig. 5.6b). 9 9a AM. Exudate white or turning brownish. Venation not densely reticulate (Fig. 5.6c). (L. clearly discoloured, fimbrial vein). APOCYNACEAE Aspidosperma, Himatanthus (AM), Plumeria (AM, cult), Cerbera (AS, OC) 9b AF. Small tree with stilt-roots. Leaf margin entire and long-petiolate. RAUH. (Exudate brownish). PHYLLANTHACEAE Uapaca (AF, MA) Group 4—Exudate white, opalescent or coloured; leaves simple, opposite or whorled arrangement. APOCYNACEAE, CELASTRACEAE, CLUSIACEAE s.l., EUPHORBIACEAE, MALPIGHIACEAE, MORACEAE, MYRTACEAE, SAPOTACAE 1a Leaves stipulate (s.t. stipules minute). 2 2a Stipular scar conspicuous. Venation densely reticulate (Fig. 5.6d). MORACEAE Bagassa (AM, Guianas), Ficus (AS) 2b Stipules or venation different. (see 3a to 3e). 3a Trees. Leaves coriaceous. Latex pale yellow, whitish or reddish. Stipules minute. CLUSIACEAE+CALOPHYLLACEAE e.g. Platonia (AM), Symphonia (AM, MA), Garcinia (e.g.: AF), Endodesmia (AF) 3b Small shrubs. Latex white (Fig. 5.6e). Leaves thin, not coriaceous. EUPHORBIACEAE Euphorbia sect. Chamaesyce 3c Tree. Twigs not lenticellate. (Latex white). SAPOTACEAE e.g. Pradosia (AM) 3d Shrub or small tree. Twigs lenticellate. Buds or young leaves with appressed hairs. (Stipules intrapetiolar). MALPIGHIACEAE e.g. Lophanthera (Brazil), Spachea (AM) 3e Woody liana. Exudate red. CELASTRACEAE Prionostemma (AM) 1b Leaves not stipulate. 4 4a Latex (gutta) yellow, orange-coloured or red. Leaf aestivation with young leaves of a same pair appressed (Fig. 5.6f). (Lamina bearing numerous trichomes or tiny dark dots: Vismia). (L. with resiniferous ducts (rd) looking like veinlets: Garcinia). CLUSIACEAE e.g. Clusia (AM), Allanblackia (AF), Garcinia

Fig. 5.6

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Keys to Families of Trees and Shrubs

43

CALOPHYLLACEAE (See CLUSIACEAE) e.g. Marila (AM), Mesua (AS), Mammea (mostly AS), Calophyllum HYPERICACEAE (See CLUSIACEAE) Vismia (AM, AF), Harungana (AF, MA) 4b Latex white, yellow, orange-coloured or red. Leaf aestivation different or young leaves not observable at the time. (See 5a to 5e). 5a Tree, shrub or liana. Latex white, seldom pale yellow. Twigs lenticellate (Fig. 5.7a). (Short twigs modified into spines: Carissa). APOCYNACEAE Aspidosperma (AM), Carissa (AS, AU), Funtumia (AF), Alstonia (mainly AS) Tabernaemontana, Rauvolfia, etc. 5b Tree or shrub. Latex (gutta) yellow or orange. Twigs not lenticellate. CLUSIACEAE s.l. Clusia, Mammea, Symphonia (AM), Cratoxylum (AS), Garcinia 5c Small liana. Latex white or pale yellow. APOCYNACEAE Asclepias (AM), Calotropis (AF, AS), Hoya (AS, OC), etc. 5d Rhythmic growth (Fig. 5.7b). Exudate smelling resin. Midrib abruptly curved near the margin (Fig. 5.7c). ANACARDIACEAE Protorhus (AF-S), Bouea (AS) 5e Exudate sticky, without resin smell. MYRTACEAE e.g. Syncarpia (AU) Group 5—Absence of a coloured or viscous exudate; two-ranked arrangement of leaves; leaves simple, stipulate; annular stipular scars. CHRYSOBALANACEAE, DIPTEROCARPACEAE, HAMAMELIDACEAE, IRVINGIACEAE, MAGNOLIACEAE, MALVACEAE, POLYGONACEAE, SARCOLAENACEAE, ULMACEAE s.l., VITACEAE 1a Leaf venation pinnate (Fig. 5.7d). 2 2a Stipules persisting, cylindrical (sometimes reduced into a ring) (Fig. 5.7e). POLYGONACEAE e.g.: Coccoloba, Ruprechtia, Triplaris (AM) 2b Stipules different. 3 3a Stipules adnate to petiole (Fig. 5.7f). VITACEAE Leea (AF, MA, AS, ornem.) 3b Stipules not adnate to petiole. 4

Fig. 5.7

4a Stipules clearly visible, large. 5a Venation brochidodromous (Fig. 5.8a).

5 MAGNOLIACEAE e.g.: Magnolia (AM, AS)

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Keys to Families of Trees and Shrubs

5b Secondary veins somewhat parallel, regularly incurved. Venation camptodromous (Fig. 5.8b). DIPTEROCARPACEAE e.g.: Dipterocarpus (AS) 4b Stipules tiny or narrow. 6 6a Paleotropics. Stipules and stem apices glabrous. Stipules thin and long (Fig. 5.8c). IRVINGIACEAE Irvingia (AF, AS) 6b AM. Stipules and stem apices pubescent. (See 7a to 7c). 7a AS. Stipules minute. ULMACEAE s.l. Gironniera (AS) 7b MA. Leaves with two false longitudinal veins (orig. leaf-folding). Stem apices with appressed hairs. (TROLL). SARCOLAENACEAE e.g.: Leptolaena, Sarcolaena (MA) 7c AS. Appressed hairs. Midrib raised slightly above the upper side of the lamina. CHRYSOBALANACEAE Atuna (AS, OC) 1b Venation palmate or L. tripliveined (Fig. 5.8d). 8 8a Leaves palmatilobate. 9 9a Stipules flat, appressed on to another (Fig. 5.8e). HAMAMELIDACEAE Exbucklandia (AS) 9b Stipules different. Bark fibrous (Fig. 5.8f). MALVACEAE Triplochiton (AS) 8b Leaves not palmatilobate. ULMACEAE s.l. Gironniera (AS), Chaetachme (AF, MA), Celtis Group 6—Leaves simple; no coloured or viscous exudate; two-ranked arrangement of leaves; leaves stipulate; stipular scars not annular; venation palmate or leaves tripliveined. EUPHORBIACEAE, HAMAMELIDACEAE, LEGUMINOSAE, MALVACEAE, MUNTINGIACEAE, PERIDISCACEAE, PASSIFLORACEAE, RHAMNACEAE, SALICACEAE, ULMACEAE s.l., URTICACEAE, VITACEAE

Fig. 5.8

1a Inner bark with a network of fibres. 2 2a Leaves with stellate or peltate hairs (Fig. 5.9a), or petiole distally pulvinate. 3 3a Tree or shrub. Petiole pulvinate or not. Leaves with stellate hairs. (Leaves lobate or cordate). (TROLL). MALVACEAE e.g. Apeiba (AM), Durio (AS), Grewia (Paleo), Pterospermum (AS) 3b Small tree with continuous flowering. TROLL. Petiole not pulvinate. Leaves with stellate hairs. MUNTINGIACEAE Muntingia (AM, cult) 2b Indument different or none. Petiole different. (ROUX Fig. 5.9b, TROLL Fig. 5.9c). 4 4a Leaves whitish beneath. Inner bark with long parallel fibres. URTICACEAE Leucosyke (AS, OC)

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Keys to Families of Trees and Shrubs

45

4b Leaves not whitish beneath, tripliveined.

ULMACEAE s.l. e.g.: Trema, Celtis 1b Inner bark different (s.t. difficult to break but without visible network). 5 5a Petiole distally pulvinate (Fig. 5.9d). 6 6a Base and upper side of the lamina bearing two glands. EUPHORBIACEAE e.g.: Glycydendron (AM) 6b Base and underside of the lamina bearing two glands. PERIDISCACEAE Peridiscus (AM, Brazil) 6c Leaves ‘unifoliolate’. No glands. (Leaves tripliveined). LEGUMINOSAE e.g. Flemingia (Paleo) 5b Petiole not distally pulvinate. 7 7a Leaves or petioles with glands. (Vine or liana). PASSIFLORACEAE Passiflora (AM, cult.) 7b Leaves without glands. 8 8a Plant self-supporting (not climbing). 9 9a Midrib raised on upper side of lamina (Fig. 5.9e). Leaves toothed or crenulate, 3–5-veined. (ROUX). SALICACEAE e.g. Prockia (AM) 9b Midrib sunken on upper side of lamina (Fig. 5.9f). (See 10a to 10c). 10a Leaf margin toothed. Venation camptodromous and clearly scalariform. (ROUX). RHAMNACEAE Ziziphus

Fig. 5.9

10b AM. Leaf margin entire, tripliveined, glabrous. (TROLL). HAMAMELIDACEAE Matudaea (AM-C) 10c Venation camptodromous (Fig. 5.10a). (ROUX, leaf margin entire or toothed, tripliveined, glabrous). ULMACEAE s.l. Celtis 8b Liana or potential liana. (See also groups 53 to 57). 11 11a Venation camptodromous and scalariform (Fig. 5.10b). (Lianas with prehensile twigs: Ventilago or tendrils: Gouania). (Leaves toothed). RHAMNACEAE Ventilago (Paleo), Gouania 11b Leaf margin toothed and venation secant. (Tendrils oppositifoliate Fig. 5.10c). VITACEAE e.g. Cissus 11c Leaf margin entire. Venation densely reticulate (L. indented or bilobate). (Liana). LEGUMINOSAE See key D, Bauhinia

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Keys to Families of Trees and Shrubs

Group 7—Absence of a coloured or viscous exudate; two-ranked arrangement of leaves; leaves simple, stipulate; stipular scars not annular; leaf venation pinnate; leaves with glands. CHRYSOBALANACEAE, DICHAPETALACEAE, DIPTEROCARPACEAE, PASSIFLORACEAE, PHYLLANTHACEAE, ROSACEAE, SALICACEAE 1a AF. Base of midrib with one solitary gland (Fig. 5.10d).

DIPTEROCARPACEAE Marquesia, Monotes (AF) 1b AF. Shrub. ROUX. Twigs hollow inhabited by ants. Glands in stipular position: Barteria. (Underside of lamina glandular: Paropsia). PASSIFLORACEAE Barteria (AF), Paropsia (AF, MA) 1c Plant neither as 1a nor as 1b. 2 2a Glands scattered on the underside of the lamina or close to, or along its margin (Fig. 5.10e). (See 3a to 3c). 3a AS. Venation camptodromous. Branching monopodial (Fig. 5.10f). (ROUX, RAUH). DIPTEROCARPACEAE Shorea, Vatica (AS) 3b Twigs stiff. Young leaves conduplicate. (TROLL). CHRYSOBLANACEAE e.g. Dactyladenia (AF), Parinari (AM) 3c Tree or liana. Glands on the underside of the lamina or disposed along the midrib. Appressed hairs. (Twigs bearing linear series of lenticels). (ROUX).

Fig. 5.10

DICHAPETALACEAE e.g. Tapura (AM), Dichapetalum 2b Base of lamina or petiole with glands (Fig. 5.11a). (See 4a to 4d). 4a Twigs supple. Young bark with thin longitudinal slits. PHYLLANTHACEAE e.g. Aporosa (AS, OC) 4b Glands disposed on lamina base, somewhat remote from midrib and leaf margin (Fig. 5.11b). CHRYSOBALANACEAE e.g. Chrysobalanus 4c Glands on lamina base. Twigs lenticellate. Stem apices and buds glabrous or with hairs not appressed. Venation brochidodromous. (RAUH, TROLL). ROSACEAE Prunus (AM, AS) 4d Glands at junction between lamina and petiole. Leaf margin hyaline (Fig. 5.11c). SALICACEAE e.g. Banara, Laetia (AM) Group 8—Absence of a coloured or viscous exudate; two-ranked leaf arrangement; leaves simple, entire, stipulate; stipular scars not annular; leaf venation pinnate; leaves and stems without glands. CAPPARACEAE, CELASTRACEAE, CHRYSOBALANACEAE, DICHAPETALACEAE, DIPTEROCARPACEAE, ERYTHROXYLACEAE, EUPHORBIACEAE, FAGACEAE, HAMAMELIDACEAE, HUACEAE, LEGUMINOSAE, MALVACEAE, OCHNACEAE, PANDACEAE, PHYLLANTHACEAE, PUTRANJIVACEAE, RHAMNACEAE, SALICACEAE, ULMACEAE s.l., VIOLACEAE

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Keys to Families of Trees and Shrubs

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1a External wood furrowed (Fig. 5.11d).

FAGACEAE e.g. Castanopsis, Lithocarpus (AS) 1b External wood not furrowed. 2 2a Strong rhythmic growth denoted by series of shorter internodes (Fig. 5.11e). 3 3a Stipules intrapetiolar. Venation not densely reticulate. (Lamina with or without false longitunal veins). ERYTHROXYLACEAE Erythroxylum 3b Numerous parallel V II or intercostal supplementary V II between main V II (Fig. 5.11f). Midrib and V II raised on the upper side of the lamina. (Stipules s.t. intrapetiolar). OCHNACEAE e.g. Ochna (AF, AS), Ouratea 2b Series of shorter internodes absent. (When petioles distinctly distally swollen, see key A to EUPHORBIACEAE). 4

Fig. 5.11

4a Stems with sequential GUs without apical meristem abortion (Fig. 5.12a). 5 5a Leaves with translucent dots or lines or leaves aromatic. Young leaves involute or hyaline margin (Fig. 5.12b). SALICACEAE e.g. Laetia, Ryania (AM) 5b Leaves without translucent dots or lines. 6 6a Twigs and leaves with numerous peltate hairs (Fig. 5.12c). Leaf margin entire. (Venation slightly tripliveined). MALVACEAE e.g. Durio (AS) 6b No peltate hairs. 7 7a Midrib raised on upper side of the lamina or young bark with thin longitudinal slits. 8 8a Lamina with a hyaline margin. (ROUX, TROLL). SALICACEAE e.g. Casearia spp. 8b Lamina without a hyaline margin. 9 9a Midrib ending in a mucro (Fig. 5.12d) (TROLL). CAPPARACEAE Capparis 9b Midrib different. (See 10a to 10c). 10 10a Leaf margin entire. Twigs supple. Petiole not distally swollen. (ROUX, TROLL). (Fimbrial vein: Bridelia). PHYLLANTHACEAE e.g. Breynia, Bridelia (Paleo), Phyllanthus 10b AS. Leaf margin entire. (Hairs fasciculate, looking like stellate hairs: Shorea). (MASSART, ROUX). DIPTEROCARPACEAE e.g. Hopea, Shorea (AS) 10c AS. Leaf margin entire or toothed. Venation camptodromous (Fig. 5.12e). (ROUX). RHAMNACEAE e.g. Alphitonia (AS), Colubrina

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Keys to Families of Trees and Shrubs

7b Midrib sunken on the underside of lamina (Fig. 5.12f). Young bark different. (See 11a to 11c). 11a AM. Venation camptodromous. (ROUX). ULMACEAE Ampelocera (AM) 11b AS. (Petiole slightly swollen distally, domatia present: Hopea). (Hairs fasciculate, looking like stellate hairs: Shorea). (MASSART, ROUX). DIPTEROCARPACEAE e.g. Hopea, Shorea (AS) 11c TROLL. (Leaves ‘unifoliolate’ with articulate or pulvinate petiole). (Spiny twigs or fimbrial vein: Dalbergia). LEGUMINOSAE e.g. Dalbergia

Fig. 5.12

4b Stem apices frequently aborting (Fig. 5.13a). (Sympodial branches). 12 12a TROLL’s model. 13 13a Branches or trunk distinctly lenticellate (Fig. 5.13b). (See 14a to 14c). 14 14a Twigs stiff. Lenticels clearly visible. CHRYSOBALANACEAE e.g. Couepia (AM), Hirtella (AM, AF), Licania (AM, AS) 14b Leaves with a hyaline margin (Fig. 5.13c). (Leaf-folding involute). SALICACEAE e.g. Homalium 14c MA-AS. Leaves glabrous and without hayline margin. (Stipules foliaceous: Maingaya). HAMAMELIDACEAE Dicoryphe (MA), Distylium, Maingaya (AS) 13b Lenticels absent or not clearly visible. 15 15a Midrib ending in a mucro or an indentation (Fig. 5.13d). (Young leaf-folding conduplicate). CAPPARACEAE e.g. Capparis 15b Midrib different. Young twigs supple. (Young leaf-folding conduplicate). EUPHORBIACEAE-PHYLLANTHACEAE e.g. Chaetocarpus (AM, AS), e.g. Savia (AM, AF, MA) 12b ROUX’s, PETIT’s, MASSART’s, FAGERLIND’s or MANGENOT’s model. 16 16a Stem apices or young leaves with appressed hairs (Fig. 5.13e). 17 17a Lamina not whitish beneath. ROUX. (L. with glands, Midrib sunken on the upper side of the lamina). DICHAPETALACEAE Tapura (AM), Dichapetalum 17b Lamina whitish beneath. EUPHRONIACEAE-TRIGONIACEAE Euphronia (AM), Trigoniastrum (AS) 16b Stem apices or young leaves without appressed hairs. 18 18a Underside of leaves with “satin finish”. (Stipular scars relatively extended). VIOLACEAE e.g. Leonia (AM), Rinorea (AM, AF) 18b Lamina different, its underside dull or shiny. 19

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Keys to Families of Trees and Shrubs

19a AF. Stem apices or buds with stellate hairs (Fig. 5.13f). Venation densely reticulate. Bark or leaves with smell of garlic.

49

HUACEAE Afrostyrax (AF)

19b Stem apices and buds different. (See 20a to 20c).

Fig. 5.13

20a Leaves coriaceous, shiny, glabrous. Young internodes angular. (ROUX). (Base of lamina asymmetrical). PUTRANJIVACEAE e.g. Drypetes 20b Leaves glabrous. Branches disposed more or less in tiers. Young internodes angular. (laticiferous threads in leaves Fig. 5.14a). CELASTRACEAE e.g. Maytenus (AM) 20c Leaves with a hyaline margin. (Young L. folding involute). SALICACEAE e.g. Casearia Group 9—Absence of a coloured or viscous exudate; two-ranked leaf arrangement; leaves simple, stipulate; stipular scars not annular; leaf margin toothed or crenulate; leaf venation pinnate; leaves without glands. APHLOIACEAE, FAGACEAE, GOUPIACEAE, HUMIRIACEAE, LACISTEMATACEAE, LECYTHIDACEAE, LEGUMINOSAE, OCHNACEAE, PUTRANJIVACEAE, RHAMNACEAE, SALICACEAE, VIOLACEAE 1a Stipules minute and lamina with vestigial teeth (v-t). Fibrous bark (attempts to break the stem often result in a strip of bark remaining attached (Fig. 5.14b). LECYTHIDACEAE Eschweilera, Lecythis (AM) 1b Stipules or teeth clearly visible or bark different. 2 2a External wood furrowed (Fig. 5.14c). FAGACEAE e.g. Castanea (AM-N, EU), Castanopsis (AS) 2b External wood not furrowed. 3 3a Midrib and V II raised on the upper side of the lamina. Numerous V II or supplementary parallel intercostal V II (Fig. 5.14d). Stipules intrapetiolar. OCHNACEAE e.g. Ouratea (AM) 3b Venation different or stipules not intrapetiolar. 4 4a Venation camptodromous and scalariform (Fig. 5.14e). RHAMNACEAE e.g. Maesopsis (AF) 4b Venation brochidodromous or not scalariform. 5 5a TROLL’s or MANGENOT’s model (Fig. 5.14f). 6 6a Laticiferous threads in leaves. (Young leaf-folding involute). (TROLL). HUMIRIACEAE Sacoglottis gabonensis (AF-W) 6b Leaves without laticiferous threads. (See 7a to 7d).

Fig. 5.14

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7a Young leaves involute (Fig. 5.15a). Hyaline lamina margin. (Inner bark with sclerenchymatous orange-coloured inclusions (Fig. 5.15b), leaves with translucent or dark dots or lines, leaves toothed). Young internodes somewhat winged. (TROLL). SALICACEAE e.g. Aphloia (APHLOIACEAE, AF-E, MA), Casearia, Homalium 7b AM. Shrub. Leaves small, pubescent. VIOLACEAE e.g. Hybanthus (AM) 7c AM. Small tree. Leaves shiny, glabrous, their teeth acute. LEGUMINOSAE Zollernia (AM) 7d AM. Tree. Leaves shiny, glabrous, their teeth obtuse. HUMIRIACEAE e.g. Humiriastrum (AM) 5b Other model, e.g. ROUX (Fig. 5.15c). 8 8a AM. Stipular scar almost annular. Leaf venation pinnate Fig. 5.15d. (ROUX). LACISTEMATACEAE Lacistema (AM) 8b Stipular scars different (See 9a to 9c). 9 9a ROUX. Young leaf involute. Lamina with a hyaline margin (Fig. 5.15e). (Leaves toothed or with translucent dots or lines). Spines: Xylosma. SALICACEAE e.g. Casearia, Xylosma 9b ROUX. Young internodes grooved or angular (Fig. 5.15f). Plant glabrous. PUTRANJIVACEAE e.g. Drypetes 9c AM. ROUX. Young internodes angular. Spreading hairs. GOUPIACEAE Goupia (AM) Group 10—Absence of a coloured or viscous exudate; bark with a network of fibres (attempts to break the stem often result in a strip of bark remaining attached); two-ranked leaf arrangement; leaves simple, not stipulate. ANNONACEAE, LECYTHIDACEAE, THYMELAEACEAE 1a Bark aromatic. Venation densely reticulate (Venation IV-V visible). (ROUX, PETIT, TROLL). (Stellate or peltate hairs: e.g. Duguetia). ANNONACEAE e.g. Annona (AM, cult.), Duguetia, Guatteria (AM) Cleistopholis, Monodora (AF), Polyalthia (Paleo), Xylopia

Fig. 5.15

1b Bark not aromatic. Leaf margin entire. Stem apices or young leaves with appressed hairs Fig. 5.16a. (Leaves pellucid-dotted, intramarginal vein: Gonystylus). THYMELAEACEAE e.g. Aquilaria (AS), Gonystylus (AS, OC), Dicranolepis (AF) 1c Bark not aromatic. Leaf margin toothed (s.t. vestigial glands). Stem apices and leaves glabrous. LECYTHIDACEAE e.g. Couratari, Eschweilera (AM), Napoleonaea (AF)

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Group 11—Absence of a coloured or viscous exudate; bark without network of fibres; two-ranked leaf arrangement; leaves simple, not stipulate; stems or leaves with glands or leaves with glandular trichomes or translucent dots. CALOPHYLLACEAE, EBENACEAE, POLYGALACEAE, PRIMULACEAE, ROSACEAE, SIMAROUBACEAE, SOLANACEAE 1a Glands scattered on the underside of the lamina (Fig. 5.16b). 2 2a Black layer between bark and wood (Fig. 5.16c). Buds or stem apices with appressed hairs. (MASSART). EBENACEAE Diospyros 2b Characters different. 3 3a Underside of lamina with glands. Glands scattered on underside of lamina, plugged on veinlets (Fig. 5.16d). Leaves turning yellowish when drying. POLYGALACEAE e.g. Xanthophyllum (AS) 3b Translucent dots disposed on the underside of the lamina, between veinlets. SIMAROUBACEAE Quassia indica (syn. Samadera indica) (MA, AS) 1b Glands disposed otherwise or underside of lamina with glandular trichomes. 4 4a Leaves with glands. 5 5a Petiole or lamina base with glands. (RAUH, TROLL). ROSACEAE Prunus (AS) 5b Glands in stipular position. POLYGALACEAE e.g. Carpolobia (AF), Securidaca (AM, AF) 4b Glandular trichomes (s.t. minute, use a hand lens!). 6 6a MASSART (Fig. 5.16e) or ROUX (Fig. 5.16f). (Liana weakly prostrate with decumbent branches: Embelia). PRIMULACEAE e.g. Ardisia (AM), Embelia (Paleo)

Fig. 5.16

6b (Neither MASSART’s nor ROUX’s model, yellow exudate really absent ?). CALOPHYLLACEAE (See CLUSIACEAE) Caraipa (AM) 6c Shrub. Leaves not coriaceous. (Crushed leaves giving out a curious smell). SOLANACEAE e.g. Cestrum (AM) Group 12—Absence of a coloured or viscous exudate; bark without network of fibres; two-ranked leaf arrangement; leaves simple, not stipulate; leaves and stems without glands or glandular trichomes; stem apices or young leaves pubescent (s.t. indumentum of minute scaly hairs). ARISTOLOCHIACEAE, CONNARACEAE, EBENACEAE, ERICACEAE, ERYTHROPALACEAE, ICACINACEAE s.l., LAURACEAE, MELIACEAE, MYRISTICACEAE, OLACACEAE, PENTAPHYLACACEAE, PERACEAE, SANTALACEAE, SOLANACEAE, STYRACACEAE, THEACEAE

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1a Inner bark astringent or turning red when cut. (MASSART). (Stem apices and buds covered by a rust-coloured indumentum). MYRISTICACEAE e.g. Virola (AM), Pycnanthus (AF), Myristica (Paleo) 1b Inner bark different. 2 2a Leaves and young internodes with stellate or peltate hairs Fig. 5.17a. (See 3a to 3d). 3a Bark sloughing off in scales (Fig. 5.17b). Periderm suberizing early (i.e. on the last or next to last GU). Leaves with stellate hairs. STYRACACEAE Styrax (AS) 3b Bark sloughing off in a different way. Late suberization. Leaves with stellate hairs. ICACINACEAE -METTENIUSACEAE e.g. Dendrobangia (AM) 3c ROUX Fig. 5.17c. (Inner bark with sclerenchymatous, orange-coloured inclusions Fig. 5.17d, spreading hairs: Okoubaka) ERYTHROPALACEAE+SANTALACEAE Octoknema, Okoubaka (AF) 3d AM. ROUX-MASSART. Leaves with stellate hairs. Leaf margin entire Fig. 5.17e. PERACEAE (see EUPHORBIACEAE) Pera (AM) 2b Neither stellate hairs or scaly hairs. (See 4a to 4c). 4a Petiole distally pulvinate (leaves ‘unifoliolate’ Fig. 5.17f). CONNARACEAE e.g. Hemandradenia, Manotes (AF), Ellipanthus (AF, AS) 4b Petiole different. Bark or leaves aromatic or giving out a curious smell. 5

Fig. 5.17

5a Tree. Leaves coriaceous. (Aromatic smell, ROUX).

LAURACEAE e.g. Cryptocarya (AS) 5b Shrub. Leaves not coriaceous. Stem apices or buds pubescent. (Crushed leaves giving out a curious smell). SOLANACEAE e.g. Cestrum (AM) 4c Petiole different. Bark and leaves not aromatic. 6 6a Young leaves or buds with appressed hairs (Fig. 5.18a). 7 7a Leaf margin entire. (See 9a to 9c). 8 8a Domatia in vein axils Fig. 5.18b. (TROLL). MELIACEAE Turraea (AF, MA, AS) 8b TROLL. Absence of domatia. (MANGENOT). ARISTOLOCHIACEAE Aristolochia arborea (AM-C) 8c MASSART. (Foliar glands really absent?). EBENACEAE Diospyros 6b Leaf margin toothed. (Young leaves involute). 9 9a ROUX. TROLL. Branching monopodial (Fig. 5.18c). Twigs lenticellate. PENTAPHYLACACEAE e.g. Freziera (AM), Adinandra (AS)

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9b Frequent abortion of apical meristems Fig. 5.18d. (TROLL). THEACEAE e.g. Camellia (AS, cult.) 7b Leaves more or less glabrous or hairs not appressed. 10 10a Venation scalariform. Leaves pubescent. V III and V IV more or less oriented in a same direction Fig. 5.18e. ICACINACEAE-METTENIUSACEAE. Emmotum (AM) 10b Venation not scalariform. 11 11a Periderm suberizing early (on the last or next to last GU). 12 12a GUs with frequent abortion of their apical meristems. Leaves coriaceous. Petiolar base delimited by an abscission zone (Fig. 5.18f). ERICACEAE e.g. Vaccinium 12b GUs without apical meristem abortion. (ROUX). Bark scaly (exceptional in Lauraceae!). Venation camptodromous. LAURACEAE e.g. Eusideroxylon (AS) 11b Stem periderm remaining green for several consecutive GUs. 13

Fig. 5.18

13a Stem apices do not abort. (ROUX: Discophora, Stemonurus). ICACINACEAE-STEMONURACEAE e.g. Discophora (AM), Leptaulus (AF, MA), Stemonurus (AS) 13b Frequent abortion of stem apices. (Terminal flowering, PETIT Fig. 5.19a: Leptaulus). Leaves glabrous but stem apices or buds pubescent. ICACINACEAE-STEMONURACEAE e.g. Leptaulus (AF, MA), Gomphandra (AS, OC), Gonocaryum (AS) Group 13—Absence of a coloured or viscous exudate; bark without network of fibres; two-ranked leaf arrangement; leaves simple, not stipulate; leaves and stems without glands or glandular trichomes; stem apices and young leaves glabrous. ANISOPHYLLEACEAE, CANELLACEAE, CELASTRACEAE, CHRYSOBALANACEAE, CORNACEAE, ERICACEAE, ERYTHROPALACEAE, HUMIRIACEAE, ICACINACEAE, IXONANTHACEAE, LECYTHIDACEAE, MARCGRAVIACEAE, OLACACEAE (incl. Schoepfia), OPILIACEAE, PIPERACEAE, PUTRANJIVACEAE, SALICACEAE !!! Leaf arrangement looking 2-ranked because of a strong anisophylly where the larger leaves respect a two-ranked arrangement, smaller leaves disposed on the upper side of the twigs Fig. 5.19b. Tree or shrub. MASSART. (Leaves tripliveined). ANISOPHYLLEACEAE Anisophyllea (mainly Paleo) Shrub or herb. Plant aromatic. Twigs modular, consisting of a sympodial series of shorter modules, each of these bearing a terminal spike of flowers Fig. 5.19c. PIPERACEAE Piper 1a Venation densely reticulate Fig. 5.19d. 2 2a Young leaves folding involute (teeth minute or vestigial). 3

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3a Bark that can be pulled off in fibrous strips Fig. 5.19e. LECYTHIDACEAE e.g. Couratari, Eschweilera, Lecythis (AM) 3b Bark different. (L. with minute teeth). 4 4a AM, AF. Twigs brittle. Petiole very short. HUMIRIACEAE-IXONANTHACEAE e.g. Sacoglottis (AM), Vantanea (AM), Phyllocosmus (AF) 4b AF. Teeth tiny or leaf margin entire. (Base of lateral twigs with short internodes and scale-leaves). LECYTHIDACEAE e.g. Brazzeia, Scytopetalum (AF) 2b Leaf-folding other or not visible at the time. 5 5a Lamina with a hyaline margin Fig. 5.19f. 6

Fig. 5.19

6a Twigs lenticellate, supple. Venation densely reticulate. 6b Twigs lenticellate, brittle. (L. toothed). 5b Leaf margin not hyaline. 7a Leaf venation pinnate Fig. 5.20a. (See 8a to 8c). 8a TROLL. Twigs lenticellate. (Mainly AM).

PUTRANJIVACEAE e.g. Putranjiva (AS) SALICACEAE e.g. Scolopia (Paleo) 7

CHRYSOBALANACEAE e.g. Licania (AM) 8b Twigs scarcely or not at all lenticellate. Leaves coriaceous. (NOZERAN: Citronella). ICACINACEAE-STEMONURACEAE e.g. Lasianthera (AF), Citronella (AM, AS, OC) 8c Leaves not coriaceous. (Venation scalariform: Minquartia). OLACACEAE + ERYTHROPALACEAE e.g. Cathedra, Heisteria, Minquartia (AM), Ochanostachys (AM) 7b Venation palmate or leaves tripliveined Fig. 5.20b. CORNACEAE Alangium (Paleo, New Caledonia) 1b Venation more or less indistinct (Veinlets IV-V not visible) Fig. 5.20c. 9 9a Bark or leaves aromatic. 10 10a Plant somewhat herbaceous. Nodes swollen Fig. 5.20d. (Terminal spikes of flowers). PIPERACEAE e.g. Piper 10b Tree. Nodes not swollen. Upper side of the lamina with sunken VI Fig. 5.20e. (Chewed leaves have a sharp taste: Capsicodendron). CANELLACEAE Cinnamodendron (AM), Cinnamosma (MA) 9b Bark and leaves not aromatic. 11 11a Stem periderm remaining green for several consecutive GUs. 12 12a Leaves somewhat coriaceous. ERYTHROPALACEAE e.g. Heisteria (AM, AF), Strombosia (AF, AS) 12b Leaves soft, shortly petiolate. OPILIACEAE e.g. Lepionurus (AS), Opilia (Paleo)

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11b Periderm suberizing early (i.e. on the last or next to last GU). 13a AM. Lenticels forming longitudinal rows Fig. 5.20f. Leaves subsessile.

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Fig. 5.20

Plant climbing by means of clamp-roots Fig. 5.21a, acquiring, in aerial flowering branches, a spiral arrangement of leaves. MARCGRAVIACEAE e.g. Marcgravia, Norantea (AM) 13b Characters different. (See 14a to 14d). 14a Bark sloughing off in small scales Fig. 5.21b. Abscission zone between petiole and stem. (Epiphyte). ERICACEAE e.g. Cavendishia (AM) 14b (ROUX). Leaves glabrous, toothed. CELASTRACEAE e.g. Siphonodon (AS, AU) 14c Lamina base somewhat cuneiform extending into a grooved petiole. Leaf margin entire. OLACACEAE+OPILIACEAE +SCHOEPFIACEAE e.g. Dulacia (AM), Olax (AF), Agonandra (AM), Shoepfia (AM, AS) 14d AM. Twigs brittle. Leaves slightly toothed, glabrous. Petiole very short. HUMIRIACEAE e.g. Sacoglottis (AM), Vantanea (AM) Group 14—Absence of a coloured or viscous exudate; spiral arrangement of leaves; annular stipular scars or petiolar base sheathing the stem; leaves simple, stipulate. DILLENIACEAE, EUPHORBIACEAE, MAGNOLIACEAE, MALVACEAE, POLYGONACEAE, URTICACEAE, VITACEAE 1a Stipule cylindrical, s.t. truncate and very short (ochrea) Fig. 5.21c.

POLYGONACEAE e.g. Coccoloba (AM) 1b Stipules different, cylindrical-conical 2 2a Petiolar base sheathing the stem Fig. 5.21d. 3 3a Stipule adnate to petiole Fig. 5.21e. Tree or shrub. Stems angled at nodes. VITACEAE e.g. Leea (AF, MA, AS) 3b Small tree with periderm suberizing early. Petiolar expansions looking like stipules or petiole grooved Fig. 5.21f. DILLENIACEAE e.g. Dillenia (AS, OC, AU) 2b Petiolar base different. 4 4a Leaves tripliveined or palmate venation. 5 5a Inner bark with a network of fibres. (Leaves cordate or lobate). MALVACEAE e.g. Hibiscus (AM, AS)

Fig. 5.21

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5b Bark different. 6a Leaves with glands. (Glands on lamina base Fig. 5.22a).

6 EUPHORBIACEAE e.g. Macaranga (Paleo) 6b Leaves without glands. URTICACEAE e.g. Cecropia, Coussapoa (AM), Myrianthus (AF) 4b Leaf venation pinnate. Stipule hood-like Fig. 5.22b. (Bark with a network of fibres or bark or leaves aromatic). MAGNOLIACEAE Magnolia s.l. (AM, AS) Group 15—Absence of a coloured or viscous exudate; bark with a network of fibres; spiral arrangement of leaves; leaves simple, stipulate; stipular scars not annular and petiolar base not sheathing the stem. AHARIACEAE, BIXACEAE, DIPTEROCARPACEAE, ELAEOCARPACEAE, LECYTHIDACEAE, MALVACEAE, ROSACEAE, URTICACEAE 1a Petiole distally swollen (pulvinus) Fig. 5.22c or leaves tripliveined or with palmate venation Fig. 5.22d. 2 2a Underside of leaves with red-brownish scaly hairs. BIXACEAE Bixa (AM) 2b Indument different. (See 3a to 3c). 3a Leaves tripliveined or palmate venation. (Stellate hairs). MALVACEAE Ochroma (AM), Dombeya (AF, MA), Firmiana (Paleo) 3b Hairs simple. Leaf venation pinnate. ELAEOCARPACEAE Elaeocarpus (MA, AS, OC) 3c Hairs simple. Stipules of a same pair asymmetrically disposed, one larger and one smaller Fig. 5.22e. (Shrub or small tree). Leaves 3- or 5-veined. URTICACEAE e.g. Myriocarpa (AM) 1b Petioles not distally swollen and Leaf venation pinnate. 4 4a Branches plagiotropic by apposition Fig. 5.22f. 5 5a Leaves not cordate. (Leaves withering red–orange). ELAEOCARPACEAE e.g. Elaeocarpus (AS, OC) 5b Leaves cordate, toothed. ACHARIACEAE Pangium edule (AS) 4b Branches not plagiotropic by apposition. 6

Fig. 5.22

6a Leaf margin entire Fig. 5.23a. (See 7a to 7c). 7a Growth strongly rhythmic. (FAGERLIND). 7b AS. Growth not strongly rhythmic. 7c Growth weakly rhythmic. (Shrub).

ROSACEAE Eryobotrya (AS, s.t. invasive) DIPTEROCARPACEAE e.g. Vatica (AS) URTICACEAE Oreocnide (AS)

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6b Leaf margin toothed Fig. 5.23b. Mainly AS, OC. Growth strongly rhythmic. Stipules minute. (LEEUWENBERG, KORIBA, SCARRONE). LECYTHIDACEAE Barringtonia (MA, AS, OC) Group 16—Absence of a coloured or viscous exudate; bark without a network of fibres; spiral leaf arrangement; leaves simple, stipulate; stipular scars not annular and petiolar bases not sheathing the stem; petiole distally swollen. ACHARIACEAE, CAPPARACEAE, CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, MEDUSANDRACEAE 1a Branches plagiotropic by apposition Fig. 5.23c. 2 2a Young bark with thin longitudinal slits. Twigs supple. EUPHORBIACEAE e.g. Baccaurea (AS, OC), Richeria (AM) 2b Leaves crenulate, (withering red–orange). ELAEOCARPACEAE e.g. Elaeocarpus (MA, AS, OC) 1b Branches not plagiotropic by apposition. 3 3a Branches plagiotropic (ROUX Fig. 5.23d). 4 4a Series of sequential GUs without abortion of apical meristems. (ROUX). ACHARIACEAE e.g. Hydnocarpus (AS) 4b Frequent abortion of stem apices Fig. 5.23e. (Bark with thin longitudinal slits). EUPHORBIACEAE e.g. Aporosa (AS, OC) 3b All branches orthotropic or plant not ramified (CORNER Fig. 5.23f). 5 5a Series of sequential GUs without abortion of apical meristems. Leaves without glands. 6

Fig. 5.23

6a AS. Venation scalariform Fig. 5.24a.

CELASTRACEAE? CENTROPLACACEAE e.g. Bhesa (AS) 6b Venation not scalariform or not in Asia. (See 7a to 7d). 7a Twigs stiff (RAUH). ELAEOCARPACEAE e.g. Sloanea (AM) 7b Twigs supple. Young bark with thin longitudinal slits Fig. 5.24b. EUPHORBIACEAE e.g. Mareya (AF) 7c Twigs brittle. Young internodes grooved, Rhythmic growth. Sectioned bark smelling bitter almonds. ACHARIACEAE e.g. Lindackeria (AM, AF) 7d Underside of leaves covered with thick hairs. MEDUSANDRACEAE Medusandra (AF)

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5b Stem apical meristems frequently aborted or leaves with glands. 8 8a Young bark with thin longitudinal slits. EUPHORBIACEAE e.g. Macaranga, Mallotus (Paleo), Alchornea, Croton 8b Midrib ending in a mucro Fig. 5.24c. CAPPARACEAE e.g. Capparis spp., Steriphoma (AM) Group 17—Absence of a coloured or viscous exudate; bark without network of fibres; spiral arrangement of leaves; leaves simple, stipulate; petiole not distally swollen; stipular scars not annular and petiolar base not sheathing the stem; leaves with glands. EUPHORBIACEAE, MALPIGHIACEAE, PASSIFLORACEAE, RHAMNACEAE, SALICACEAE 1a Leaves (lamina or petiole) with glands. 2a Petiole with glands Fig. 5.24d. (Liana).

2 PASSIFLORACEAE Passiflora (AM, cult., invasive) 2b Lamina with glands Fig. 5.24e. 3 3a Young bark with thin longitudinal slits or plant aromatic. (Leaves tripliveined). EUPHORBIACEAE e.g. Croton 3b Bark different or plant not aromatic. 4 4a Glands on upper side of lamina base. (Leaves tripliveined Fig. 5.24f). SALICACEAE Hasseltia (AM)

Fig. 5.24

4b Glands disposed otherwise. 5 5a Young leaves, buds, or stem apices with appressed hairs Fig. 5.25a. Glands on underside of lamina base. MALPIGHIACEAE Acridocarpus (AF) 5b Stem apices, buds and leaves without appressed hairs. Glands on lamina margin (Fig. 5.25b). RHAMNACEAE e.g. Colubrina (AM) Group 18—Absence of a coloured or viscous exudate; bark without a network of fibres; spiral arrangement of leaves; leaves simple, stipulate; stipular scars not annular and petiolar base not sheathing the stem; petiole not distally swollen; leaves without glands. ACHARIACEAE, AQUIFOLIACEAE, CELASTRACEAE, CORYNOCARPACEAE, DIPTEROCARPACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, FAGACEAE, IXONANTHACEAE, LECYTHIDACEAE, LINACEAE, OCHNACEAE, RHAMNACEAE, SALICACEAE, VIOLACEAE 1a External wood furrowed Fig. 5.25c. (Leaf margin entire or toothed). FAGACEAE e.g. Castanopsis (AS), Quercus 1b External wood not furrowed. 2 2a Midrib and V II raised on the upper side of the lamina. Numerous VII or supplementary intercostals V II raised on upper side of lamina. Stipules intrapetiolar Fig. 5.25d. OCHNACEAE e.g. Ouratea, Poecilandra (AM)

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2b Characters different (stipules not intrapetiolar). 3 3a Plant monocaulous, not ramified. (CORNER Fig. 5.25e). Large, cuneate leaves, shortly petiolate. 4 4a AF. Understorey shrub. ACHARIACEAE Phyllobotryon (AF) 4b AS. Understorey shrub. EUPHORBIACEAE Agrostistachys (AS) 3b Plant ramified at adult stage. 5 5a Leaves with palmate venation or tripliveined Fig. 5.25f. 6 6a Petiolar scars almost as wide as the youngest internodes. (Base of lamina with glands, leaves peltate for some spp.). EUPHORBIACEAE e.g. Dalechampia (AM), Macaranga (Paleo)

Fig. 5.25

6b Petiolar scars different.

ACHARIACEAE e.g. Caloncoba (AF) 5b Leaves with pinnate venation or supra-tripliveined Fig. 5.26a. 7 7a Leaf margin toothed or crenulate. 8 8a Leaves with a hyaline margin Fig. 5.26b. SALICACEAE e.g. Flacourtia (Paleo, cult.) 8b Leaves without a hyaline margin. 9 9a Buds and young leaves glabrous. 10 10a Rhythmic growth with series of shorter internodes and long scale-leaves. Parallel V II: Roucheria. (Laticiferous threads in leaves: Ixonanthes; leaf-folding involute). LINACEAE-IXONANTHACEAE Hebepetalum, Roucheria (AM) - Ixonanthes (AS.), 10b (Shrub). Absence of long scale-leaves. CELASTRACEAE e.g. Mystroxylon (AF-E, MA) 9b Buds and young leaves pubescent. AS. (Leaf base cuneate, RAUH, AUBRÉVILLE). ELAEOCARPACEAE Elaeocarpus (MA, AS, OC) 7b Leaf margin entire. 11 11a Venation not densely reticulate, s.t. indistinct Fig. 5.26c. 12 12a Branches not plagiotropic by apposition. AQUIFOLIACEAE Ilex 12b NG, AU, OC. Branches plagiotropic by apposition Fig. 5.26d. (Bitter bark). CORYNOCARPACEAE Corynocarpus (NG, AU, OC) 11b Venation densely reticulate (N IV visible) Fig. 5.26e. 13 13a AM. Leaves somewhat grouped on shorter internodes. Stipules minute. Venation camptodromous. LECYTHIDACEAE Couroupita (AM, cult.)

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13b At least, at least one of these characters different. (L. entire or toothed). (See 14a to 14c). 14a Lamina with translucent dots Fig. 5.26f. SALICACEAE e.g. Casearia, Ryania (AM) 14b Underside of leaves with “satin finish”. VIOLACEAE e.g. Leonia, Payparola, Rinorea (AM)

Fig. 5.26

14c Underside of lamina different. 15 15a Branches plagiotropic by apposition Fig. 5.27a. ELAEOCARPACEAE Elaeocarpus (AS, OC). 15b Branches not plagiotropic by apposition. (See 16a to 16c). 16a Branches monopodial. Leaf margin entire. V II more or less parallel Fig. 5.27b. (MASSART). DIPTEROCARPACEAE e.g. Parashorea, Shorea (AS) 16b Young bark with thin longitudinal slits Fig. 5.27c. EUPHORBIACEAE e.g. Claoxylon (Paleo), Acalypha 16c AS. Leaves tripliveined. (RAUH?). RHAMNACEAE e.g. Colubrina (AS) Group 19—Absence of a coloured or viscous exudate; spiral arrangement of leaves; leaves simple, not stipulate; petiole distally swollen (pulvinus); venation palmate or leaves tripliveined. ARALIACEAE, EUPHORBIACEAE, HAMAMELIDACEAE, MENISPERMACEAE 1a Transversal cut of trunk or branches showing a succession of concentric layers Fig. 5.27d. (Liana, petiole pulvinate at both ends Fig. 5.27e). MENISPERMACEAE e.g. Abuta (AM), Cocculus 1b Trunk and branch sections different. 2 2a Underside of lamina with waxy, pale-rose, indumentum. Petiole slightly swollen distally. (Stipules could be present at some nodes). HAMAMELIDACEAE Rhodoleia (AS) 2b Leaves different. 3 3a Bark or leaves aromatic or spiced. Petiolar bases sheathing Fig. 5.27f. ARALIACEAE e.g. Dendropanax (AM, AS) 3b Young bark with thin longitudinal slits. Twigs supple. (Petiole distally pulvinate, e.g. Hieronyma, Mallotus). EUPHORBIACEAE e.g. Hieronyma (AM), Mallotus (Paleo), Croton Group 20—Absence of a coloured or viscous exudate; spiral arrangement of leaves; leaves simple, not stipulate; petiole not distally pulvinate; venation palmate or leaves tripliveined.

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Fig. 5.27

ACHARIACEAE, ANISOPHYLLEACEAE, BORAGINACEAE, CARICACEAE, ERICACEAE, EUPHORBIACEAE, HERNANDIACEAE, LAURACEAE, PIPERACEAE, TETRAMELACEAE 1a Bark or leaves aromatic. 2 2a Petiolar base sheathing the stem. Branches sympodial, modular (flowering terminal). PIPERACEAE Piper 2b Petiolar base not sheathing the stem. 3 3a Stem periderm remaining green for several consecutive GUs. Young internodes angular or grooved Fig. 5.28a. LAURACEAE e.g. Nectandra (AM), Cinnamomum (AS, AU) 3b Periderm suberizing early (i.e. on the last or next to last GU). EUPHORBIACEAE e.g. Croton 1b Bark or leaves not aromatic. 4 4a Inner bark with a network of fibres Fig. 5.28b. 5 5a Plant monocaulous, not ramified (CORNER). Leaves palmatilobate Fig. 5.28c. CARICACEAE e.g. Carica (AM, cult.) 5b Plant ramified or leaves not palmatilobate. (See 6a to 6c). 6a Branches erect. Branching monopodial Fig. 5.28d. (Large tree). (Peltate hairs: Octomeles). TETRAMELACEAE Octomeles, Tetrameles (AS) 6b Branches plagiotropic by apposition Fig. 5.28e. (Small tree). Leaves toothed with cordate base. Bark with smell of bitter almonds. ACHARIACEAE Pangium (AS) 6c Leaves pubescent on both faces, at least at their base. (Branches drooping). BORAGINACEAEE e.g. Cordia 4b Inner bark without a network of fibres. 7 7a Base of lamina with glands Fig. 5.28f. EUPHORBIACEAE e.g. Croton 7b Lamina without glands. 8 8a Upper side of leaves with erect hairs. (PRÉVOST, CHAMPAGNAT). BORAGINACEAE e.g. Cordia 8b Upper side of leaves different. 9

Fig. 5.28

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9a Leaf margin entire. 10 10a Petiole base with decurrent ridges or wings extending down the stem Fig. 5.29a. (Leaves peltate: Hernandia). HERNANDIACEAE Gyrocarpus, Hernandia 10b Petiole base different. (See 11a to 11d). 11a Young bark with thin longitudinal slits Fig. 5.29b. Twigs supple. EUPHORBIACEAE e.g. Strophioblachia (AS) 11b Trunk with eye marks. Serial buds Fig. 5.29c. ACHARIACEAE e.g. Kiggelaria (AF) 11c Shrub. Bark sloughing off in small scales Fig. 5.29d. (in montane or cloud forests). ERICACEAE e.g. Cavendishia, Semiramisia (AM-S) 11d Venation distinctly reticulate. (leaf arrangement appearing superficially tworanked, Large leaves alternating with smaller leaves). ANISOPHYLLEACEAE Anisophyllea (mainly Paleo) 9b Leaf margin toothed. Branches erect, monopodial. Petiole base with decurrent ridges or wings extending down the stem. Branches erect, monopodial. (Very large tree, peltate hairs: Octomeles). TETRAMELACEAE Tetrameles, Octomeles (AS) Group 21—Absence of a coloured or viscous exudate; lateral or terminal short shoots, s.t. modified into thorns, or spines epidermal; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or supratripliveined. ASTERACEAE, BIGNONIACEAE, BORAGINACEAE, CACTACEAE, CAPPARACEAE, CELASTRACEAE, COMBRETACEAE, CONNARACEAE, LAURACEAE, OLACACEAE, RUTACEAE, SOLANACEAE, SIMAROUBACEAE, SURIANACEAE 1a Presence of short shoots (Fig. 5.29e). (See also 1b and 1c). 2a Leaves with translucent dots (essential oils).

2 RUTACEAE Aeglopsis (AF), Atalantia (AS) 2b Leaves different. 3 3a Shrub, ROUX’s model (Fig. 5.29f). 4 4a Leaves soft, glabrous. Venation somewhat indistinct (ROUX). OLACACEAE Ximenia americana (AM, invasive)

Fig. 5.29

4b Leaves glabrous, coriaceous.

CELASTRACEAE e.g. Gymnosporia (AF, AS)

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3b Plant different. (Leaves small). 4a Venation clearly visible. (Branches plagiotropic by apposition). 5a Petiole not distally pulvinate. (L. with glands).

4 5 COMBRETACEAE e.g. Terminalia (AF) 5b AF. Petiole distally pulvinate (Fig. 5.30a). CONNARACEAE Burttia (AF-E) 4b Venation indistinct. (Shrub bearing small or medium-sized leaves). 6 6a Stem apices or buds pubescent. Lamina notched (Fig. 5.30b). BORAGINACEAE e.g. Rochefortia (AM) 6b Stem apices or buds pubescent. Lamina not notched. SOLANACEAE e.g. Lycium (AM, AF) 6c AM. Plant entirely glabrous. SIMAROUBACEAE Castela (AM) 1b Spines of epidermal origin or in stipular position (See 7a to 7c). 7a Spines in stipular position (Fig. 5.30c). L. notched or with apical mucro. CAPPARACEAE e.g. Capparis (Paleo) 7b Spines axillary (1–2-3 per axil). Venation indistinct. CACTACEAE Pereskia (AM, cult.) 7c Spines of epidermal origin. Leaves not notched. (LEEUWENBERG (Fig. 5.30d), KORIBA (Fig. 5.30e, PRÉVOST Fig. 5.30f). (Possibly an ASTERACEAE). SOLANACEAE e.g. Solanum 1c Plant without short shoots and not spiny. 8 8a Stem periderm remaining green for several consecutive GUs. Young internodes angular or grooved. (AUBRÉVILLE). LAURACEAE Aniba (AM) 8b Periderm suberizing early (i.e. on the last or next to last GU). Or young internodes (sub)cylindrical. 9 9a Upper side of leaves pubescent. BORAGINACEAE e.g. Bourreria (AM), Cordia 9b Leaves glabrous or only the underside pubescent. 10 10a Petiole short. (See 11a to 11c). 11

Fig. 5.30

11a Shrub or small tree. CHAMPAGNAT (Fig. 5.31a). Leaves grouped on short twigs. BIGNONIACEAE Crescentia (AM, cult.) 11b Venation camptodromous. (AUBRÉVILLE). (Leaves with glands). COMBRETACEAE e.g. Terminalia (AF) 11c Shrub of seashore bearing small leaves. Twigs lenticellate. SURIANACEAE Suriana

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Group 22—Leaves simple; no coloured or viscous exudate; plant not spiny; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or tripliveined; leaves with glands or stellate or peltate hairs. CAPPARACEAE, COMBRETACEAE, EUPHORBIACEAE, HUMIRIACEAE, PASSIFLORACEAE, SALICACEAE, STYRACACEAE 1a Leaf margin entire (Fig. 5.31b). (See 2a to 2d). 2 2a Base or margin of lamina with glands or glands in vein axils. COMBRETACEAE e.g. Terminalia (e.g. AM), Strephonema (AF), Conocarpus (mangrove) 2b Leaf margin entire or with scaly hairs. Midrib ending in a mucro or/and an indentation (Fig. 5.31c). CAPPARACEAE e.g. Capparis 2c Stellate hairs. Neither mucro nor indentation. (RAUH Fig. 5.31d). STYRACACEAE Styrax (AM) 2d Young bark with thin longitudinal slits. Twigs supple. (Stipules really absent ?). EUPHORBIACEAE e.g. Spondianthus (AF), Croton 1b Leaf margin toothed (Fig. 5.31e). 3 3a AM. Leaves glabrous, with a short petiole basally enlarged (pb), lamina with glands at base. HUMIRIACEAE e.g. Humiria (AM) 3b At least, at least one of these character different. (See 4a to 4c). 4a Leaves pubescent. Leaves crenulate, glands at lamina base (Fig. 5.31f). PASSIFLORACEAE e.g. Turnera (AM)

Fig. 5.31

4b One pair of glands at the junction of the petiole with the lamina. Lamina with a hyaline margin. Twigs lenticellate and brittle. (Trunk spiny: Scolopia). SALICACEAE e.g. Scolopia (Paleo) 4c Young bark with thin longitudinal slits (Fig. 5.32a). Twigs supple. (Stipules really absent ?). EUPHORBIACEAE e.g. Pausandra (AM), Alchornea Group 23—Absence of a coloured or viscous exudate; plant not spiny; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or supratripliveined; leaves without glands but with glandular trichomes (possibly a waxy indument beneath) or translucent dots. ASTERACEAE, LAURACEAE, MYRTACEAE, PENTAPHYLACACEAE, PRIMULACEAE, RHABDODENDRACEAE, RUTACEAE, SAPINDACEAE, SCHISANDRACEAE, VERBENACEAE, WINTERACEAE 1a Leaves with translucent dots or lines (Fig. 5.32b). 2 2a Bark sloughing off in scales or narrow strips (Fig. 5.32c). (ATTIMS). MYRTACEAE e.g. Eucalyptus, Tristania (AS, AU, cult.), Leptospermum (AS, OC, NZ)

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2b Bark different. 3 3a Leaves with an articulation between petiole and lamina (Plant aromatic). RUTACEAE e.g. Esenbeckia, Pilocarpus (AM), Toddalia (AS), Vepris (e.g. MA) 3b No such articulation. (See 4a to 4c). 4a Plant not aromatic. (CORNER: Oncostemum, RAUH, MASSART). (An oval-shaped scar is left after branch shedding Fig. 5.32d). PRIMULACEAE e.g. Ardisia (e.g. AS), Oncostemum (MA) 4b Young internodes angular or grooved (Fig. 5.32e). Periderm green on the last or next to last GU. (RAUH, MASSART, AUBRÉVILLE). LAURACEAE e.g. Beilschmiedia 4c Shrub. Leaves aromatic or with translucent dots. VERBENACEAE Aloysia (AM, cult.) 1b Leaves without translucent dots. 5 5a Underside of leaves with glandular trichomes or minute dark dots (Fig. 5.32f). 6 6a Trunk with continuous branches (ATTIMS). Young internodes angular. Twigs brittle. SAPINDACEAE Dodonaea 6b Plant different. 7

Fig. 5.32

7a External wood furrowed (Fig. 5.33a). Leaves with minute shiny trichomes. (Terminal flowering). ASTERACEAE e.g. Gochnatia (AM) 7b External wood not furrowed. (See 8a to 8d). 8a Trichomes shiny or dark. (CORNER, RAUH, MASSART). (Bark with a network of fibres, oval-shaped scars (bs) on trunk). PRIMULACEAE e.g. Myrsine (mainly AM), Tapeinosperma (NG, AU, OC) 8b Trichomes minute, not shiny. Growth strongly rhythmic (Fig. 5.33b). GUs with leaves not grouped distally. (RAUH). PENTAPHYLACACEAE Anneslea (AS), Ternstroemia (mainly AM-AS) 8c Growth units with leaves grouped distally. SCHISANDRACEAE Illicium (AM, AS) 8d AM. CORNER (Fig. 5.33c). Rain forest treelet. Leaves strechted, cuneate, entire. Petiole grooved. Venation indistinct. RHABDODENDRACEAE Rhabdodendron (AM) 5b Underside of leaves waxy (waxy plugged stomata, RAUH: Drimys, SCARRONE: Takhtajania). WINTERACEAE e.g. Drimys (AM, AS, OC), Takhtajania (MA), Zygogynum (NG, OC) Group 24—Absence of a coloured or viscous exudate; plant not spiny; branches plagiotropic by apposition; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or supratripliveined; leaves without glands or glandular trichomes or translucent dots. ACTINIDIACEAE, ANACARDIACEAE, BONNETIACEAE, BORAGINACEAE, COMBRETACEAE, GOODENIACEAE, LAURACEAE, MELIACEAE, RUTACEAE, SAPOTACEAE

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1a Bark or leaves giving out an aromatic or spicy smell. 2a Petiole distally swollen (or bearing an articulation) (Fig. 5.33d).

2 RUTACEAE e.g. Esenbeckia (AM) 2b Petiole different. 3 3a Young internodes angular or grooved (Fig. 5.33e). (Stem periderm remaining green for several consecutive GUs, if not then: Aniba, AM). LAURACEAE e.g. Aniba (AM), Dehaasia (AS, NG), Neolitsea (AS, AU), Ocotea 3b V II strongly curved near the lamina margin (Fig. 5.33f). ANACARDIACEAE e.g. Campnosperma (e.g. AS)

Fig. 5.33

1b Bark and leaves aromatic. 4 4a V II secant on lamina margin (Fig. 5.34a) or V II ending almost at lamina margin. Thick hairs, appressed on young parts. Venation secant. ACTINIDIACEAE e.g. Saurauia (AM, AS) 4b Venation not secant, forming loops clearly separate from lamina margin. 5 5a Venation IV-V visible (Fig. 5.34b). 6 6a Erect or oblique hairs (also on the upper side of the lamina). BORAGINACEAE e.g. Cordia (AM) 6b Indument different. (See 7a to 7d). 7a Leaves subsessile or shortly petiolate. (Glands really absent?). COMBRETACEAE e.g. Bucida, Buchenavia (AM), Terminalia 7b Young internodes angular or grooved (Fig. 5.34c). (Stem periderm remaining green for several consecutive GUs). LAURACEAE e.g. Aniba (AM), Actinodaphne, Phoebe (AS) 7c Twigs lenticellate. Petiole quite long. Venation camptodromous. (AUBRÉVILLE). MELIACEAE Vavaea (AS, OC) 7d Twigs not lenticellate. Petiole quite long. Venation camptodromous (Fig. 5.34d) (Latex really absent?). SAPOTACEAE e.g. Pradosia (AM) 5b Venation IV-V indistinct. 8 8a Leaves auriculate at its base. (AUBRÉVILLE). BONNETIACEAE Archytaea (AM), Ploiarium (AS) 8b Leaves not auriculate. 9 9a Leaves shortly petiolate or subsessile. Young leaf-folding revolute (Fig. 5.34e). (shrub of back-shore). GOODENIACEAE e.g. Scaevola (AS, AU, OC, cult.) 9b Petiole quite long. 10 Bark sloughing off in strips (Fig. 5.34f). (AUBRÉVILLE). COMBRETACEAE e.g. Terminalia (e.g. AF)

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Group 25—Absence of a coloured or viscous exudate; inner bark fibrous, fibres parallel or forming a network or external bark furrowed; plant not spiny; branches not plagiotropic by apposition; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or supratripliveined; leaves with no glands, no glandular trichomes and no translucent dots.

Fig. 5.34

BORAGINACEAE, THYMELAEACEAE, ICACINACEAE s.l., LECYTHIDACEAE, PROTEACEAE, SABIACEAE 1a External wood furrowed (Fig. 5.35a). 2 2a Venation brochidodromous Fig. 5.35b. Inner bark with a network of fibres. PROTEACEAE e.g. Roupala (AM), Faurea (AF, MA), Helicia (AS) 2b Venation camptodromous Fig. 5.35c and V III-IV somewhat parallel. Bark not fibrous. ICACINACEAE-METTENIUSACEAE e.g. Emmotum (AM) 1b External wood not furrowed. 3 3a Upper side of leaves pubescent. BORAGINACEAE e.g. Cordia (AM) 3b Leaves glabrous or only their underside pubescent. (See 4a to 4d). 4a Leaf margin entire. Stem apices pubescent. (KORIBA). BORAGINACEAE e.g. Bourreria (AM) 4b Leaf margin entire. (MASSART, bark difficult to tear and detaching in strips Fig. 5.35d). THYMELAEACEAE e.g. Daphnopsis (AM) 4c Large glabrous leaves, cuneate at base, bearing minute teeth. (CORNER: Gustavia, LEEUWENBERG, KORIBA). LECYTHIDACEAE e.g. Gustavia, Grias (AM), Barringtonia, Chydenanthus, Planchonia (mainly AS, OC) 4d Petiolar base distinctly swollen Fig. 5.35e. Leaves somewhat grouped. Lamina cuneate at base. SABIACEAE Meliosma (AS) Group 26—Absence of a coloured or viscous exudate; inner bark not fibrous; external bark not furrowed; plant not spiny; branches not plagiotropic by apposition; spiral arrangement of leaves; leaves simple, not stipulate; petiolar base sheathing the stem or petiole distinctly grooved; leaf venation pinnate or supratripliveined; leaves with no glands, no glandular trichomes and no translucent dots. ARALIACEAE, DILLENIACEAE, TORRICELLIACEAE 1a Bark or L. with an aromatic or distinctive smell. (Petiolar base sheathing the stem Fig. 5.35f). ARALIACEAE e.g. Dendropanax (AM)

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Fig. 5.35

1b Bark and leaves not aromatic. 2a Small tree bearing deeply incised leaves.

2 TORRICELLIACEAE e.g. Aralidium (AS) 2b Leaves different. 3 3a Long, grooved petiole Fig. 5.36a. Stems and leaves smooth. TORRICELLIACEAE Melanophylla (MA) 3b Venation secant Fig. 5.36b. Stems and leaves scabrous. (Lianas). DILLENIACEAE e.g. Curatella (AM), Dillenia (AS, AU)

Group 27—Absence of a coloured or viscous exudate; inner bark not fibrous; external bark not furrowed; plant not spiny; branches not plagiotropic by apposition; spiral arrangement of leaves; leaves simple, not stipulate; leaf venation pinnate or supratripliveined; petiolar base neither sheathing the stem nor grooved; petiole distally swollen; leaves without glands, no glandular trichomes and no translucent dots. ACHARIACEAE, CAPPARACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, LEPIDOBOTRYACEAE, OXALIDACEAE, PHYLLANTHACEAE 1a Branches plagiotropic (ROUX). Bark with orange-coloured sclerenchymatic inclusions (Ryparosa), (Fig. 5.36c) or with smell of bitter almonds: Lindackeria. ACHARIACEAE Lindackeria (AM), Ryparosa (AS) 1b Characters different. 2 2a Leaf margin entire with midrib ending in a mucro or stellate hairs. (Stellate hairs: Steriphoma, minute peltate hairs: Morisonia), Fig. 5.36d. CAPPARACEAE e.g. Morisonia, Steriphoma (AM) 2b Leaves different. Hairs neither stellate nor scaly. 3 3a Petiole with an articulation (leaves unifoliolate). 4 4a AS Venation camptodromous Fig. 5.36e. OXALIDACEAE Sarcotheca (Malaysia, Indonesia) 4b AF. Venation brochidodromous Fig. 5.36f. LEPIDOBOTRYACEAE Lepidobotrys (AF) 3b Petiole without an articulation. (See 5a to 5c). 5a Buds or young leaves covered by appressed hairs. (Stipules really absent ?). ELAEOCARPACEAE e.g. Elaeocarpus (AS, A U, OC), Sloanea (AM) 5b Young bark with thin longitudinal slits. Twigs supple. EUPHORBIACEAE e.g. Codiaeum (AS, cult.)

Fig. 5.36

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5c Petiolar scars almost as wide as the youngest internodes. Petiole quite long. (RAUH, leaves with stellate hairs ?). PHYLLANTHACEAE Uapaca (AF) Group 28—Leaves simple; no coloured or viscous exudate; spiral arrangement of leaves; leaves not stipulate; Leaf venation pinnate or tripliveined; plant not spiny; leaves with no glands, no glandular trichomes and no translucent dots; branches not plagiotropic by apposition; inner bark not fibrous; external bark not furrowed; petiolar base neither sheathing the stem nor grooved; petiole not distally swollen: venation II or III secant Fig. 5.37a. ACTINIDIACEAE, ANACARDIACEAE, CLETHRACEAE, DILLENIACEAE 1a Leaves perfectly entire Fig. 5.37b, smelling of resin.

ANACARDIACEAE e.g. Ozoroa (AF) 1b Leaf margin toothed (at least at lamina apex) or plant not resinous. 2 2a Stems rough. Petiole distinctly grooved Fig. 5.37c. DILLENIACEAE e.g. Curatella (AM) 2b Leaves and young internodes with thick hairs Fig. 5.37d. ACTINIDIACEAE Saurauia (AM, AS) 2c Indument different. Growth strongly rhythmic Fig. 5.37e. (Stellate or peltate hairs, leaf margin entire: American Clethra). CLETHRACEAE Clethra (AM, AS)

Group 29—Absence of a coloured or viscous exudate; inner bark not fibrous; external bark not furrowed; plant not spiny; branches not plagiotropic by apposition; strong rhythmic growth denoted by series of GUs without apical abortion; leaves spiral; leaves simple, not stipulate; petiolar base neither sheathing the stem nor grooved; petiole not distally swollen; leaf venation pinnate or supratripliveined; absence of glands, glandular trichomes, translucent dots; venation II-III not secant. ANACARDIACEAE, BONNETIACEAE, CANELLACEAE, CAPPARACEAE, CLETHRACEAE, CORNACEAE s.l., CYRILLACEAE, DAPHNIPHYLLACEAE, ERICACEAE, GOODENIACEAE, HAMAMELIDACEAE, IXONANTHACEAE, LAURACEAE, LECYTHIDACEAE, PENTAPHYLACACEAE, PITTOSPORACEAE, PRIMULACEAE, SIMAROUBACEAE, THEACEAE

Fig. 5.37

1a Crushed leaves or scratched bark giving out an aromatic or resinous smell. 2 2a V II abruptly curved near the lamina margin Fig. 5.38a. Leaves and bark with resinous smell. ANACARDIACEAE e.g. Anacardium (AM, cult.), Mangifera (AS, cult.) 2b Venation somewhat indistinct. Aromatic or spicy smell. CANELLACEAE Canella (West Indies)

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1b Leaves without aromatic smell. 3 3a Leaves very shortly petiolate, coriaceous or somewhat fleshy. 4 4a Lamina obtuse or acuminate. 5 5a Leaves grouped at the distal ends of the GUs Fig. 5.38b. (See 6a to 6c). 6a Paleo. Young leaves convolute Fig. 5.38c. LECYTHIDACEAE e.g. Petersianthus (AS) Foetidia (AF, MA, AS), Barringtonia (Paleo) 6b AS, OC. Leaves fleshy, entire or toothed. (RAUH). GOODENIACEAE Scaevola (AS, AU, OC, cult.) 6c AM. Mangrove. Young leaves convolute. Trunk fluted. TETRAMERISTACEAE Pelliciera (AM-C) 5b Leaves not grouped at ends of GUs. AM. Young leaves involute Fig. 5.38d. (RAUH). (Highlands). BONNETIACEAE Bonnetia (AM-S) 4b Leaves acute, coriaceous. PRIMULACEAE Jacquinia (AM) 3b Characters different. 7 7a Lamina cuneate at base and long red petiole. (RAUH). DAPHNIPHYLLACEAE Daphniphyllum (AS, AU, cult.) 7b Leaves different. 8 8a Adult plant monocaulous (CORNER Fig. 5.38e). Leaves large. PRIMULACEAE e.g. Clavija, Theophrasta (AM) 8b Adult plant ramified. 9 9a Leaves distinctly grouped at distal parts of the GU. (See 10a to 10e). 10a Bark fibrous. (Leaves large, s.t. toothed). CORNER: Grias. LECYTHIDACEAE e.g. Gustavia, Grias (AM), Careya (AS) 10b Leaves small or medium-sized, entire. Sympodial architecture. (KORIBA Fig. 5.38f). (Venation indistinct). PITTOSPORACEAE e.g. Pittosporum (Paleo)

Fig. 5.38

10c LEEUWENBERG, KORIBA. Abscission zone between lamina and petiole Fig. 5.39a. (Highlands). ERICACEAE e.g. Rhododendron (AS) 10d Leaf margin entire. Venation indistinct. (Underside of leaves with minute trichomes). PENTAPHYLACACEAE e.g. Ternstroemia (AM, AS) 10e Series of short internodes. Large leaves with long petioles. Numerous V II connected to a fimbrial vein. (Bark bitter). SIMAROUBACEAE Soulamea (AS, OC, NC) 9b Leaves not distinctly grouped. 11 11a Leaf margin toothed or crenulate Fig. 5.39b. (Asia). (See 12a to 12d). 12a Apical buds with several imbricate scale-leaves. HAMAMELIDACEAE Altingia (AS)

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12b Branches sympodial. Leaves large.

LECYTHIDACEAE Careya (AS) 12c Venation indistinct. (Branches monopodial, terminal flowering). THEACEAE e.g. Camellia, Gordonia (AS) 12d Venation distinctly reticulate. (Branches sympodial, terminal flowering). IXONANTHACEAE e.g. Ixonanthes (AS) 11b Leaf margin entire. 13 13a Buds or young leaves with appressed hairs Fig. 5.39c. 14 14a Young internodes angular or grooved. (Scaly buds f. s. spp.). LAURACEAE e.g. Licaria (AM), Persea (AM, cult.), Litsea (AS, AU) 14b Young internodes (sub)cylindrical. THEACEAE e.g. Schima (AS) 13b Buds or young leaves glabrous or without stellate hairs. 15 15a Venation indistinct Fig. 5.39d. Branching monopodial Fig. 5.39e. THEACEAE e.g. Gordonia (AS) 15b Venation reticulate. 16 16a Midrib raised on upper side Fig. 5.39f. Leaves notched or/and with an apical mucro. Stellate hairs. CAPPARACEAE e.g. Morisonia (AM), Buchholzia (AF), Capparis

Fig. 5.39

16b Midrib sunken at upper side Fig. 5.40a. 17 17a Venation brochidodromous Fig. 5.40b. (leaves glabrous). 18 18a Young leaves conduplicate Fig. 5.40c. CORNACEAE s.l. e.g. Mastixia (AS), Nyssa (AM-C, AS) 18b AM. Lamina with a hyaline margin. CYRILLACEAE Cyrilla (AM-C) 17b Venation camptodromous. AM. Leaves pubescent. CLETHRACEAE Clethra (AM) Group 30—Absence of a coloured or viscous exudate; inner bark not fibrous; external bark not furrowed; plant not spiny; branches not plagiotropic by apposition; growth weakly rhythmic or frequent abortion of apical meristems (discontinuous pith or terminal flowering); spiral arrangement of leaves; leaves simple, not stipulate; petiolar base neither sheathing the stem nor grooved; petiole not distally swollen; leaf venation pinnate or supratripliveined; venation II or II not secant; leaves with no glands, no glandular trichomes and no translucent dots. BORAGINACEAE, CANELLACEAE, CAPPARACEAE, CELASTRACEAE, COMBRETACEAE, ERICACEAE, HUMIRIACEAE, ICACINACEAE, IXONANTHACEAE, LAURACEAE, LEGUMINOSAE (phyllodized Australian Acacia), MYRICACEAE, OPILIACEAE, PENTAPHYLACACEAE, PHYSENACEAE, SABIACEAE, SALICACEAE, SANTALACEAE, SOLANACEAE, SYMPLOCACEAE, THEACEAE

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1a Leaf margin toothed or crenulate Fig. 5.40d. 2a MA. Large leaves with distinct hyaline margin and secant venation.

2 CELASTRACEAE Brexia (MA) 2b Not in MA or characters different. 3 3a Leaves or buds pubescent (hand lens !). 4 4a Branches monopodial Fig. 5.40e. RAUH. Buds and young leaves pubescent. (Leaves turning yellow when drying). SYMPLOCACEAE Symplocos (AM, AS) 4b Branches sympodial Fig. 5.40f. Buds and young leaves pubescent (s.t. scabrous). Petiole quite long. BORAGINACEAE e.g. Ehretia (Paleo) 3b Leaves glabrous. (See 5a to 5c). 5a Branches monopodial. RAUH. Underside of leaves with waxy indumentum. L. crenulate, shortly petiolate with a cuneate base. MYRICACEAE Morella

Fig. 5.40

5b Short shoots modified into spines Fig. 5.41a: Gymnosporia. CELASTRACEAE e.g. Gymnosporia (Paleo), Mystroxylon (AF, MA) 5c Northern AM-S. Leaves glabrous, somewhat crenulate. Midrib raised on the upper side of the lamina Fig. 5.41b. IXONANTHACEAE Ochthocosmus (AM, Guiana highlands) 1b Leaf margin entire. 6 6a Venation somewhat indistinct. 7 7a Mangrove tree. (ATTIMS). COMBRETACEAE e.g. Lumnitzera (Paleo) 7b Not a mangrove tree. 8 8a V II more or less parallel. SANTALACEAE e.g. Exocarpos (AS, AU) 8b Venation different (distinctly reticulate). 9 9a Bark aromatic. Midrib sunken on the upper side of the lamina Fig. 5.41c. (Bark lenticellate: Warburgia). CANELLACEAE e.g. Canella (AM), Warburgia (AF-E), Cinnamodendron (AU) 9b Bark not aromatic. (See 10a to 10d). 10a AM, AS. Leaves coriaceous, entire or toothed, glabrous. Trunk and branches monopodial, lateral flowering: Gordonia. Branches sympodial: Ternstroemia. THEACEAE + PENTAPHYLACACEAE e.g. Gordonia (AM, AS), Ternstroemia (AM, AS) 10b AS, AU. Leaves somewhat smooth. (Spines: Cansjera). OPILIACEAE e.g. Cansjera (AS, AU), Lepionurus, Melientha (AS) 10c AM. L. entire. Short petioles swollen at base Fig. 5.41d. HUMIRIACEAE e.g. Humiria (AM)

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10d MA. Leaf margin entire. Periderm remaining green.

PHYSENACEAE Physena (MA) 6b Venation reticulate with distinct V III Fig. 5.41e. 11 11a Bark sloughing off in strips or small scales. Abscission zone between petiole and lamina Fig. 5.41f. ERICACEAE e.g. Bejaria (AM), Craibiodendron (AS) 11b Bark or leaves different. 12 12a Petiole distally swollen or leaves with stellate hairs. (Midrib ending in an apical mucro). CAPPARACEAE e.g. Morisonia (AM), Capparis

Fig. 5.41

12b Petiole not pulvinate and leaves without stellate hairs. 13 13a Young internodes angular or grooved Fig. 5.42a. Stem periderm remaining green for several consecutive GU.s Branches monopodial. Lateral flowering. LAURACEAE e.g. Ocotea 13b Characters different. 14 14a Leaves more or less coriaceous. 15 15a Leaves and stem apices glabrous. 16 16a Twigs lenticellate Fig. 5.42b. Petioles short. SALICACEAE e.g. Ludia (AF, MA), Scolopia (AS) 16b Leaf margin entire. Terminal flowering. ICACINACEAE e.g. Apodytes (Paleo) 15b Leaves, buds or stem apices pubescent Fig. 5.42c. 17 17a Branches monopodial Fig. 5.42d. (RAUH). Leaves turning yellow when drying. Lateral flowering. SYMPLOCACEAE Symplocos (AM, AS) 17b AM. Plant slightly lianescent or with drooping branches. Underside of L. with rust-coloured indument. ICACINACEAE Leretia (AM-S) 14b Leaves not coriaceous. (See 18a to 18c). 18a Tree. Branches sympodial Fig. 5.42e. Leaves not coriaceous, glabrous, entire and longly petiolate. PHYTOLACCACEAE e.g. Phytolacca dioica (AM, cult.) 18b Shrub not very woody. Leaves pubescent or scabrous. Sympodial architecture (LEEUWENBERG) or branches drooping (CHAMPAGNAT). (Leaves without axillary buds due to fusion between petiole and stem): Solanum. SOLANACEAE e.g. Brunfelsia, Cestrum, Solanum (AM) 18c Liana. Leaves glabrous. Twigs not lenticellate. Base of petiole enlarged Fig. 5.42f. Venation embossed at upper side. SABIACEAE Sabia (AS) Note: Many phyllode-bearing Australian species of Acacia (LEGUMINOSAE) are also part of this group. These have twigs angular in cross-section and a somewhat parallel venation.

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Group 31—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; leaves simple, stipulate; venation palmate or leaves tripliveined. CANNABACEAE, EUPHORBIACEAE, RHAMNACEAE, URTICACEAE

Fig. 5.42

1a Leaves with glands (or without glands, but with venation camptodromous and midrib sunken on the upper side of lamina). RHAMNACEAE Colubrina (e.g. AM) 1b Leaves without glands. 2 2a Bark fibrous Fig. 5.43a. (Shrub or small tree). URTICACEAE e.g. Debregeasia (Paleo), Boehmeria 2b Bark not fibrous. 3 3a Venation densely reticulate (V IV-V visible), (Fig. 5.43b). Leaf margin toothed. CANNABACEAE (see ULMACEAE s.l.) Lozanella (AM) 3b Venation different. (Young bark with thin longitudinal slits Fig. 5.43c). (Anisophylly). (Petiole distally swollen for some species). EUPHORBIACEAE e.g. Hancea (MA, AS), Mallotus (Paleo) Group 32—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; petiolar bases connected by stipular scars or stipules intrapetiolar; leaves simple, stipulate; leaf venation pinnate. CHLORANTHACEAE, CTENOLOPHONACEAE, CUNONIACEAE, ERYTHROXYLACEAE, LOGANIACEAE s.l., RHAMNACEAE, RHIZOPHORACEAE, RUBIACEAE, TRIGONIACEAE 1a Leaves distinctly toothed or crenulate. 2a Venation scalariform Fig. 5.43d. (ROUX).

2 RHAMNACEAE Lasiodiscus (AF, MA)

2b Venation not scalariform. (See 3a to 3c). 3a Plant aromatic. Leaf margin toothed Fig. 5.43e. (ATTIMS). CHLORANTHACEAE e.g. Hedyosmum (AM, AS), Ascarina (MA, AS, OC), Sarcandra (AS) 3b Plant not aromatic. (Leaves crenulate, stipules inter- or intrapetiolar). CUNONIACEAE e.g. Weinmannia (AM), Geissois (AU, OC) 3c Plant not aromatic. Twigs lenticellate Fig. 5.43f. (ROUX-ATTIMS). RHIZOPHORACEAE e.g. Cassipourea

Fig. 5.43

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1b Leaf margin entire or with minute teeth. 4 4a Venation camptodromous and twigs not lenticellate. (Petiole bearing one solitary gland or a pair of glands: Henriquezia, Platycarpum, Northern AM-S). RUBIACEAE Most species in this family Coffea (AF, cult.), Mussaenda, Pavetta (Paleo), Psychotria, etc. 4b Venation brochidodromous Fig. 5.44a. or twigs lenticellate. 5 5a Underside of leaves whitish. Twigs densely lenticellate. TRIGONIACEAE Trigonia (AM) 5b Plant different. 6 6a Mangrove. (Underside of lamina with trichomes). RHIZOPHORACEAE e.g. Bruguiera (Paleo), Rhizophora 6b Vegetation not a mangrove. 7 7a Large, intrapetiolar stipules Fig. 5.44b sheathing the apical bud. CUNONIACEAE e.g. Lamanonia (AM), Schizomeria (AS, NG, AU) 7b Stipules interpetiolar Fig. 5.44c (s.t. reduced to an interpetiolar ring). 8 8a Twigs lenticellate. L. entire. (Branches erect). RUBIACEAE e.g. Guettarda spp, Simira spp (AM), Gardenia 8b Branches decumbent. GELSEMIACEAE (see LOGANIACEAE) Mostuea (AF, MA) 8c AF. (Plant glabrous: Aneulophus; stellate hairs: Ctenolophon). ERYTHROXYLACEAE (Aneulophus, AF-W) CTENOLOPHONACEAE (Ctenolophon, AF-W), Group 33—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; leaves simple, entire, stipulate; petiolar bases not connected by stipular scars; leaf venation pinnate. CELASTRACEAE, CLUSIACEAE, LOGANIACEAE s.l., LYTHRACEAE, MALPIGHIACEAE, PICRODENDRACEAE, RHAMNACEAE, RHIZOPHORACEAE, VIOLACEAE, VOCHYSIACEAE 1a Stipule large, adnate to the petiole Fig. 5.44d.

LOGANIACEAE Neuburgia (AS, OC) 1b Stipule different (s.t. small and inserted on petiole). 2 2a Leaves with glands or glands in stipular position. 3 3a AM, AF. Leaves with glands in stipular position Fig. 5.44e. VOCHYSIACEAE e.g. Qualea, Vochysia (AM), Erismadelphus (AF-W) 3b Leaves with glands (lamina and petiole have to be examined). Buds or young leaves with appressed hairs Fig. 5.44f. (Liana). MALPIGHIACEAE Most species in this family

Fig. 5.44

2b Leaves and stems without glands. 4 4a Buds and young leaves with appressed hairs. (Liana). AUBRÉVILLE or FAGERLIND and stipules intrapetiolar Fig. 5.45a: Byrsonima. MALPIGHIACEAE e.g. Banisteriopsis, Byrsonima, Stigmaphyllon (AM), Heteropterys (AM, AF-W) 4b Buds and young leaves without appressed hairs. 5

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5a Bark sloughing off in scales. Venation camptodromous. (Stipules minute or vestigial). LYTHRACEAE e.g. Lagerstroemia (AS) 5b Bark or venation different. 6 6a Tree bearing horizontal branches (MASSART Fig. 5.45b) and large leaves. Internodes distinctly angular. LYTHRACEAE Duabanga (AS) 6b Plant different. 7 7a Branches regularly spaced on trunk. ROUX-ATTIMS. (See 8a to 8c). 8a Leaves with resiniferous ducts (rd), (Fig. 5.45c). (Latex really absent?). CLUSIACEAE e.g. Garcinia (AS) 8b Liana bearing short twigs modified into spines (Scutia) or tree (Berchemia). RHAMNACEAE e.g. Berchemia (Paleo), Scutia 8c Twigs lenticellate. (External wood furrowed for some species Fig. 5.45d, leaves with vestigial teeth ?). (ATTIMS). RHIZOPHORACEAE e.g. Sterigmapetalum (AM), Anopyxis (AF), Blepharistemma (India), Gynotroches (AS, OC), Pellacalyx (AS), Cassipourea 7b Branches disposed otherwise or other model. (See 9a to 9c). 9a Underside of leaves with “satin finish”. Stipules distinct, caducous, leaving distinct scars. VIOLACEAE Rinorea (AM) 9b Leaves different, glabrous and coriaceous. (possibly lianas, orange-coloured layer between bark and inner bark, leaves with laticiferous threads Fig. 5.45e). CELASTRACEAE e.g. Pleurostylia, Zinowiewia (AM), Lophopetalum (AS, AU), Euonymus (Paleo) 9c AS. Whorled leaf arrangement Fig. 5.45f. PICRODENDRACEAE Mischodon (India, Ceylan)

Fig. 5.45

Group 34—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; leaves simple, stipulate; leaf margin toothed; petiolar bases not connected by stipular scars; leaf venation pinnate. CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, OCHNACEAE, RHIZOPHORACEAE, VIOLACEAE 1a Lamina with secant venation Fig. 5.46a. Stipules long and thin. Leaves glabrous and large.

OCHNACEAE e.g. Quiina (AM) 1b Venation not secant. 2 2a Buds and young leaves with appressed hairs Fig. 5.46b. ELAEOCARPACEAE e.g. Aceratium (AS, AU, OC), Sericolea (NG)

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2b Buds and young leaves without appressed hairs. 3a Leaves with glands. (PETIT Fig. 5.46c).

3 EUPHORBIACEAE e.g. Tetrorchidium (AF) 3b Leaves without glands. 4 4a Subopposite arrangement of leaves, these differing slightly in size. VIOLACEAE e.g. Rinorea (AM) 4b Leaves of a same pair opposite and equal-sized Fig. 5.46d. 5a Tree. Twigs distinctly lenticellate Fig. 5.46e. Stipules conspicuous. (ROUX, ATTIMS). RHIZOPHORACEAE e.g. Blepharistemma (Inde), Cassipourea 5b Shrubs. Twigs different. Stipules minute. (RAUH). CELASTRACEAE e.g. Crossopetalum (AM), Denhamia (AU), Elaeodendron, Reissantia (Paleo)

Group 35—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; nodes with interpetiolar ridge; leaves simple, stipulate; venation tripliveined, palmate or supra-tripliveined. LAMIACEAE, LOGANIACEAE s.l., MELASTOMATACEAE, OLEACEAE, VERBENACEAE, VIBURNACEAE 1a Venation scalariform and with main secondary veins extending to the lamina apex Fig. 5.46f. (Leaves pubescent). MELASTOMATACEAE Very numerous species: Miconia (AM), Dichaeanthera (AF, MA), Medinilla (Paleo) 1b Venation different (in particular not scalariform). 2

Fig. 5.46

2a Young twigs more or less quadrangular. (See 3a to 3c). 3 3a Plant pubescent. (Base of lamina with glands Fig. 5.47a: Citharexylum, Gmelina). LAMIACEAE - VERBENACEAE e.g. Gmelina (AS) - Citharexylum (AM) 3b AS. Plant pubescent. Leaves without glands. Upper side of lamina with hairs inserted on epidermic protuberences. OLEACEAE Nyctanthes (AS) 3c AS. Liana. Plant glabrous. OLEACEAE Myxopyrum (AS) 2b Young twigs not quadrangular. 4 4a Branches plagiotropic or drooping. Leaves 3-veined or supra-3-veined Fig. 5.47b, glabrous. (Liana bearing hooks). LOGANIACEAE Strychnos 4b Young leaves involute Fig. 5.47c. (Leaves pubescent). VIBURNACEAE e.g. Viburnum (AM, EU, AS) Group 36—Leaves simple; no coloured or viscous exudate; opposite or whorled leaf arrangement; nodes or internodes swollen; leaves not stipulate; nodes with interpetiolar ridge; leaf venation pinnate or leaves not 3-veined or venation somewhat indistinct. ACANTHACEAE, GNETACEAE, NYCTAGINACEAE (Mirabilis), AMARANTHACEAE

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1a Herb or shrub. Venation camptodromous Fig. 5.47d. (mainly understorey shrubs or herbs). ACANTHACEAE Numerous species 1b Small tree or liana. Venation without blind veinlets Fig. 5.47e. (Really no latex?). GNETACEAE Gnetum 1c Herb. Internodes cylindrical. (Anisophylly Fig. 5.47f). NYCTAGINACEAE e.g. Mirabilis (AM, cult.), Boerhavia Group 37—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; nodes with interpetiolar ridge; nodes and internodes not swollen; leaves simple, not stipulate; venation indistinct or leaves like small scales or needles. ACANTHACEAE, CASUARINACEAE, CUPRESSACEAE, GENTIANACEAE, LORANTHACEAE, LYTHRACEAE, MELASTOMATACEAE, PINACEAE

Fig. 5.47

1a Mangrove. 2 2a Stems with glands near base of petiole Fig. 5.48a. Erect aerial roots (pneumatophores Fig. 5.48b. LYTHRACEAE Sonneratia (Paleo) 2b Stems without glands. Venation indistinct. No pneumatophores. ACANTHACEAE Avicennia 1b Plant not of mangroves. 3 3a Leaves scale-like Fig. 5.48c or needle-like Fig. 5.48d. (See 4a to 4c). 4a Tree abundantly ramified with slender twigs bearing very numerous small scaly leaves (s.t. only 4 per whorl through). (RAUH). CASUARINACEAE Casuarina (AS, OC, cult.) 4b Scale-leaves present. Periderm remaining green on three successive orders of branches. CUPRESSACEAE e.g. Callitris (AU, NC) 4c Leaves needle-like. Needles in groups of 2, 3 ou 5. PINACEAE Pinus 3b Leaves different. (See 5a to 5c). 5a Tree or shrub. Venation indistinct. MELASTOMATACEAE e.g. Mouriri (AM), Memecylon (Paleo) 5b AM. Venation V II-III visible. (ROUX). GENTIANACEAE e.g. Lisianthus, Tachia (AM) 5c Epiphyte (hemiparasite). Venation indistinct. Leaves not coriaceous. LORANTHACEAE e.g. Phthirusa (AM), Dendrophthoe (Paleo)

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Group 38—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; leaves simple, not stipulate; nodes with interpetiolar ridge; nodes and internodes not swollen; leaf venation pinnate or leaves not tripliveined; venation reticulate or leaves standard-sized. ACANTHACEAE, ASTERACEAE, CLUSIACEAE, CRYPTERONIACEAE, GENTIANACEAE, LAMIACEAE, LOGANIACEAE s.l., LYTHRACEAE, VERBENACEAE 1a Leaves with glands or underside of leaves with dark dots or lines, or resiniferous ducts (rd) Fig. 5.48e, visible against the light. 2 2a Leaves with glands (on lamina base Fig. 5.48f). Internodes angular. VERBENACEAE e.g. Citharexylum (AM)

Fig. 5.48

2b Leaves without glands. Lamina with dark dots or lines, or resiniferous ducts (rd) visble against light. CLUSIACEAE + HYPERICACEAE e.g. Tovomita, Vismia (AM) 1b Leaves different, (s.t. translucent dots). 3 3a Petiolar scars united Fig. 5.49a, (forming a ridge). 4 4a Tree or shrub. GENTIANACEAE e.g. Anthocleista (AF, MA), Coutoubea (AM), Fagraea (AS, AU, OC) 4b AM. STONE’s model Fig. 5.49b. Small tree. Twigs lenticellate. LOGANIACEAE Antonia (AM) 3b Petiolar scars different. 5 5a Bark sloughing off in small scales or fibrous strips Fig. 5.49c. 6 6a Leaf margin toothed. (Extratropical). HYDRANGEACEAE Hydrangea (AS, cult.) 6b Leaf margin entire. 7 7a Branches monopodial. (Venation camptodromous). LYTHRACEAE e.g. Woodfordia (Paleo) 7b Branches sympodial. (Venation camptodromous). CRYPTERONIACEAE Crypteronia (AS) 5b Bark different. 8 8a External wood furrowed Fig. 5.49d. ASTERACEAE e.g. Calea, Verbesina (AM) 8b External wood not furrowed. 9 9a Leaves of the same pair slightly different in size (anisophylly) Fig. 5.49e. ACANTHACEAE e.g. Bravaisia (AM, cult.) 9b Leaves of the same pair equal in size. (See 10a to 10c). 10a Young leaves involute Fig. 5.49f. VIBURNACEAE Viburnum (AM, AS)

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10b Leaf-folding not involute. Shrub with drooping branches: Congea; large leaves: Tectona. LAMIACEAE-VERBENACEAE e.g. Congea, Tectona (AS, cult.), Callicarpa 10c Shrub. Leaf margin entire with leaf-folding not involute. Venation camptodromous. (LOGANIACEAE)-GELSEMIACEAE Mostuea (Paleo)

Fig. 5.49

Group 39—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; nodes with interpetiolar ridge; leaves simple, not stipulate; venation tripliveined or supra-3–5-veined. BUXACEAE, GNETACEAE, LAMIACEAE, LAURACEAE, MELASTOMATACEAE, MYRTACEAE, SAPINDACEAE, SCROPHULARIACEAE 1a One or two pairs of lateral veins extend from leaf base to leaf apex Fig. 5.50a. Twigs angular. Leaves pubescent. MELASTOMATACEAE e.g. Miconia (AM), Melastoma, Medinilla (AS) 1b Plant different. 2 2a Bark or leaves aromatic. (Leaves supra-tripliveined). LAURACEAE Cinnamomum (AS, AU, OC) 2b Bark or leaves not aromatic. 3 3a Venation polygonal, without blind veins Fig. 5.50b. Base of internodes swollen. Small tree or liana. (ROUX). GNETACEAE Gnetum 3b Plant different. 4 4a Growth strongly rhythmic (series of shorter internodes) Fig. 5.50c. 5 5a Early suberization of the periderm patchy. Internodes quadrangular. Leaves tripliveined. BUXACEAE e.g. Buxus 5b Suberization different. Twigs lenticellate. SAPINDACEAE Acer (Northern Thailand and Laos.) 4b Growth weakly rhythmic Fig. 5.50d. 6 6a Leaves without translucent dots. 7 7a Leaves with stellate hairs: Premna. Base of lamina with glands: Clerodendron pp, Gmelina. LAMIACEAE e.g. Gmelina (AS, AU, NC), Premna (AF, AS) 7b Shrub. Sunken venation in leaves Fig. 5.50e. Discolorous leaves. (ROUX, CHAMPAGNAT). SCROPHULARIACEAE Buddleja (Andes, AS) 6b Leaves with translucent dots Fig. 5.50f. MYRTACEAE e.g. Rhodamnia (AS, OC)

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Group 40—Absence of a coloured or viscous exudate; opposite or whorled leaf arrangement; leaves simple, not stipulate; nodes without interpetiolar ridge; leaf venation pinnate; leaves with translucent dots or glandular trichomes, or stellate or peltate hairs.

Fig. 5.50

ASTERACEAE, BIGNONIACEAE, COMBRETACEAE, EBENACEAE, MALPIGHIACEAE, MONIMIACEAE, MYRTACEAE, OLEACEAE, RUTACEAE, TRIMENIACEAE, VERBENACEAE 1a Petiole or lamina with glands. 2 2a Young leaves or buds with appressed hairs Fig. 5.51a. (Liana). (Leaves with glands). MALPIGHIACEAE e.g. Bunchosia (AM) 2b No appressed hairs. 3 3a Leaf arrangement in whorls of three. Base of upper side of lamina with 1–2 cupuliform glands. BIGNONIACEAE Deplanchea (AM) 3b Plant different. 4 4a Young internodes angular Fig. 5.51b. Base of lamina distal part of petiole with glands. VERBENACEAE e.g. Citharexylum (AM) 4b Young internodes subcylindrical. (MASSART, ATTIMS). COMBRETACEAE e.g. Laguncularia (mangrove AM, AF), Combretum, Terminalia 1b Leaves without glands. 5 5a Petiole distally swollen Fig. 5.51c. Lamina with translucent dots. RUTACEAE e.g. Metrodorea (AM), Euodia (Paleo) 5b Petiole not distally swollen or lamina without translucent dots. 6 6a Leaves with translucent dots Fig. 5.51d. (See 7a to 7c). 7a Shrub of open sites. External wood furrowed Fig. 5.51e. ASTERACEAE e.g. Porophyllum (AM) 7b External wood not furrowed. Young bark rust-coloured. (Section of inner bark turning violet: polyphenols). MYRTACEAE Species very numerous (Eugenia, Syzygium) 7c ROUX. (Shrub or liana). TRIMENIACEAE Trimenia (Malaysia, AU, OC) 6b Leaves without translucent dots. 8 8a Underside of leaves with scaly hairs Fig. 5.51f. EBENACEAE Euclea (AF) 8b Leaves without scaly hairs (s.t. minute glands or peltate or spherical trichomes). (9a-9d).

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Fig. 5.51

9a Architecture sympodial. Some modules erect (trunk) and other modules oblique (branches) Fig. 5.52a. Sectioned bark turning violet (polyphenols). MYRTACEAE e.g. Calyptranthes (AM), Eugenia 9b Architecture not sympodial. (Venation camptodromous). (Peltate hairs, glandular trichomes, or petioles with glands). COMBRETACEAE e.g. Thiloa (AM-N), Combretum 9c Stem periderm remaining green for several consecutive GUs. (Plant aromatic, leaves with stellate hairs: Siparuna). MONIMIACEAE e.g. Siparuna (AM), Xymalos (AF) 9d Underside of leaves with minute peltate or spherical trichomes. Lenticels raised Fig. 5.52b. Periderm suberizing early. OLEACEAE e.g. Olea (Paleo) Group 41—Absence of a coloured or viscous exudate; bark sloughing off in strips or scales; opposite or whorled leaf arrangement; leaves simple, not stipulate; nodes without interpetiolar ridge; leaf venation pinnate; leaves without translucent dots, glandular trichomes, stellate or peltate hairs. CLUSIACEAE, COMBRETACEAE, LYTHRACEAE, MELASTOMATACEAE, MYRTACEAE, NYCTAGINACEAE, ONAGRACEAE, VERBENACEAE 1a Distal part of twigs with internodes twisting alternatively clockwise and counterclockwise Fig. 5.52c. 2 2a Venation III-IV indistinct. 3 3a Young leaves revolute. MASSART. MANGENOT MELASTOMATACEAE e.g. Mouriri (AM), Memecylon (AF, AS) 3b ROUX. ATTIMS. Leaves subsessile. LYTHRACEAE e.g. Pemphis (AS) 2b Venation reticulate (V III visible). 4 4a Young internodes angular. (See 5a to 5c). 5a Venation camptodromous Fig. 5.52d. (TROLL). LYTHRACEAE e.g. Lagerstroemia 5b Apical buds scaly. Bark astringent. KORIBA. HYPERICACEAE (see CLUSIACEAE) Cratoxylum (AS) 5c Bark sloughing off in scales or thin strips Fig. 5.52e. Venation brochidodromous Fig. 5.52f. (MASSART). LYTHRACEAE Duabanga (AS)

Fig. 5.52

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4b Young internodes not angular (MASSART, leaves with glands, venation camptodromous). COMBRETACEAE e.g. Anogeissus (AF, AS), Terminalia (AS), Combretum 1b All branches erect. No such alternating twisting of the internodes, but possibly twisting of petioles in horizontal branches. 6 6a Venation camptodromous. 7 7a Leaves with stellate hairs. Fig. 5.53a. VERBENACEAE e.g. Callicarpa 7b Leaves without stellate hairs.

LYTHRACEAE e.g. Physocalymma (Brazil), Lagerstroemia (AS) 6b Venation brochidodromous or indistinct. 8 8a Short twigs modified into spines Fig. 5.53b. NYCTAGINACEAE Pisonia 8b Short twigs not modified into spines. (See 9a to 9c). 9 9a Branches modular, sympodial or lamina with intramarginal veins Fig. 5.53c. MYRTACEAE e.g. Calyptranthes 9b Branches monopodial Fig. 5.53d. Crushed leaves aromatic. MYRTACEAE e.g. Pimenta 9c Branches erect: e.g. Lafoensia; small marsh plant: Ludwigia; (ATTIMS). LYTHRACEAE-ONAGRACEAE e.g. Lafoensia (AM), Fuchsia (AM), Ludwigia Group 42—Absence of a coloured or viscous exudate; bark not sloughing off in small strips or scales; opposite or whorled leaf arrangement; leaves simple, not stipulate; nodes without interpetiolar ridge; leaf venation pinnate; leaves with intramarginal or fimbrial veins, or venation parallel; leaves without translucent dots, glandular trichomes, stellate or peltate hairs. ARAUCARIACEAE, CALOPHYLLACEAE, MYRTACEAE, VOCHYSIACEAE 1a Lamina with intramarginal veins Fig. 5.53e or numerous parallel V II (Fig. 5.53f). 2 2a Branches erect. VOCHYSIACEAE Ruizterania (AM-S) 2b Branches plagiotropic. 3 3a Sectioned bark turning violet (polyphenols). MYRTACEAE e.g. Eugenia (AM), Syzygium (Paleo)

Fig. 5.53

3b Bark different. Numerous parallel V (KORIBA Fig. 5.54a). Underside of lamina waxy. CALOPHYLLACEAE (See CLUSIACEAE) Mesua (AS)

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1b Venation different. 4 4a AS, NG, AU. Lamina developed. Venation parallel. Petioles decurrent on stems Fig. 5.54b. ARAUCARIACEAE Agathis (AS, NG, OC) 4b Leaves coriaceous, needle-like. (Plant aromatic). MYRTACEAE Baeckea (AS, AU, OC) Group 43—Absence of a coloured or viscous exudate; bark not sloughing off in small strips or scales; opposite or whorled leaf arrangement; nodes without interpetiolar ridge; leaves simple, not stipulate; leaves standard-sized, without intramarginal or fimbrial veins; petiole distally swollen; venation pinnate; leaves without translucent dots, glandular trichomes, stellate or peltate hairs. LAMIACEAE, BIGNONIACEAE 1a Young internodes quadrangular Fig. 5.54c or lenticels not raised.

LAMIACEAE Teijsmanniodendron (AS) 1b Young internodes not quadrangular. (Lenticels s.t. raised Fig. 5.54d). BIGNONIACEAE e.g. Tabebuia spp (AM) Group 44—Absence of a coloured or viscous exudate; bark not sloughing off in small strips or scales; opposite or whorled leaf arrangement; nodes without interpetiolar ridge; leaves simple, not stipulate; petiole not distally swollen; leaves standard-sized, without intramarginal or fimbrial veins; leaf margin entire; leaf venation pinnate; leaves without translucent dots, glandular trichomes, stellate or peltate hairs. ANACARDIACEAE, BUXACEAE, ICACINACEAE, LAMIACEAE, LAURACEAE, MALPIGHIACEAE, MELASTOMATACEAE, MONIMIACEAE, MONTINIACEAE, MYRTACEAE, NYCTAGINACEAE, OLEACEAE, PICRODENDRACEAE, PROTEACEAE, SANTALACEAE, THYMELAEACEAE 1a Inner bark with a network of fibres or external wood furrowed. (See 2a to 2c). 2a Inner bark fibrous Fig. 5.54e. Attempts to break the stem often result in a strip of bark peeling down the side. THYMELAEACEAE e.g. Lophostoma (AM), Craterosiphon (AF), Phaleria (AS, OC)

Fig. 5.54

2b External wood furrowed Fig. 5.54f.

PROTEACEAE e.g. Panopsis (AM) 2c Bark not fibrous. Anisophylly (Fig. 5.55a). NYCTAGINACEAE e.g. Neea (AM) 1b Inner bark or external wood different. 3 3a Bark or leaves giving out an aromatic, spicy or resinous smell. (See 4a to 4c). 4a Secondary veins abruptly curved near lamina margin Fig. 5.55b. Bark or leaves with smell of resin. ANACARDIACEAE Bouea (AS) 4b Leaves with aromatic, not resinous smell. Venation different. Young internodes angular Fig. 5.55c. Stem periderm remaining green on several consecutive GUs. (Venation camptodromous). LAURACEAE e.g. Beilschmiedia (AM), Caryodaphnopsis (AM, AS)

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4c AF-S. MA. Spicy smell. Leaf margin entire, not coriaceous. Modular and sympodial architecture. MONTINIACEAE Grevea (MA) 3b Bark and leaves without such smell. 5 5a Branches sympodial, each module consisting of a single GU. 6 6a Venation brochidodromous Fig. 5.55d. (See 7a to 7c). 7a Mainly AM. Stem apices or buds with rust-coloured indument. NYCTAGINACEAE e.g. Neea (AM), Pisonia 7b Paleo. Leaves coriaceous. Twigs not lenticellate. (Young bark rust-coloured, venation indistinct). MELASTOMATACEAE Memecylon (AF, MA) 7c Paleo. Twigs lenticellate, lenticels raised (Fig. 5.55e). OLEACEAE e.g. Chionanthus, Schrebera (AF, AS) 6b If branches sympodial, then some GUs not aborting, or branches monopodial. 8 8a Venation densely reticulate (V IV visible) Fig. 5.55f. 9 9a Young internodes angular. (See 10a to 10c). 10 10a Periderm with early suberization. (Serial buds or stellate hairs, leaf margin entire). LAMIACEAE-VERBENACEAE e.g. Callicarpa (AS), Clerodendron 10b AS, AU. Distal part of the last internodes somewhat flattened. Leaf margin entire. SANTALACEAE Santalum (AS, AU) 10c AS. Leaf margin entire. Leaves 3-whorled.

Fig. 5.55

PICRODENDRACEAE e.g. Mischodon (AS) 9b Young internodes subcylindrical. (See 11a to 11c). 11a Twigs with raised lenticels Fig. 5.56a. (Underside of leaves with dense, very short hairs). OLEACEAE e.g. Noronhia (MA), Linociera 11b Twigs lenticellate. Young leaves or buds with appressed hairs Fig. 5.56b. (Twining liana). MALPIGHIACEAE e.g. Flabellaria (AF), Tetrapterys (AM) 11c Stem periderm remaining green for several consecutive GUs. Venation brochidodromous Fig. 5.56c. Stem apex somewhat flattened Fig. 5.56d. (Leaves with vestigial teeth?). MONIMIACEAE e.g. Mollinedia (AM) 8b Venation different. 12 12a Stem apices or buds or underside of leaves with appressed hairs. (Twining liana). MALPIGHIACEAE Many lianas species 12b Underside of leaves and buds without appressed hairs. (See 13a to 13d).

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13a Underside of leaves with very numerous, densely disposed minute epidermal dots (trichomes?). OLEACEAE e.g. Comoranthus (MA), Linociera, Olea 13b Leaves coriaceous. Lamina supra-tripliveined (Fig. 5.56e). Petiole base with decurrent ridges down the stem. BUXACEAE e.g. Buxus (AM, AS) 13c Two opposite faces of young internodes grooved, but the two other faces not (Fig. 5.56f). MYRTACEAE e.g. Campomanesia (AM-S) 13d AF, MA. (ATTIMS). ICACINACEAE Cassinopsis (AF, MA)

Fig. 5.56

Group 45—Absence of a coloured or viscous exudate; bark not sloughing off in small strips or scales; opposite or whorled leaf arrangement; nodes without interpetiolar ridge; leaves simple, not stipulate; leaf venation pinnate; leaves standard-sized, without intramarginal or fimbrial veins; petiole not distally swollen; leaf margin toothed, crenulate, or lobate; leaves without translucent dots, glandular trichomes, stellate or peltate hairs. CELASTRACEAE, LAMIACEAE, MONIMIACEAE, OLEACEAE, PROTEACEAE 1a Inner bark with a network of fibres. External wood furrowed Fig. 5.57a.

PROTEACEAE e.g. Panopsis (AM) 1b Bark or wood different. 2 2a Stem periderm remaining green on several consecutive GUs. Apical internode somewhat flattened Fig. 5.57b. MONIMIACEAE e.g. Glossocalyx (AF), Mollinedia, Siparuna (AM) 2b Periderm different. (See 3a to 3c). 3a Young internodes more or less quadrangular Fig. 5.57c. LAMIACEAE e.g. Premna (AF, AS) 3b Internodes subcylindrical. Twigs with raised lenticels Fig. 5.57d. OLEACEAE e.g. Chionanthus (e.g. AM), Forsythia (AS, cult.) 3c Young internodes subangular. Leaves shortly petiolate. CELASTRACEAE e.g. Crossopetalum (AM)

Group 46—Leaves compound: alternate leaf arrangement; leaves stipulate; leaves pinnate. BURSERACEAE, LEGUMINOSAE, MELIANTHACEAE, ROSACEAE, RUTACEAE, SIMAROUBACEAE, SAPINDACEAE, STAPHYLEACEAE, VITACEAE 1a Shrub or small tree. Stipules of the same pair appressed one to another Fig. 5.57e. (Venation more or less scalariform). VITACEAE Leea (AF, MA, AS)

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1b Stipules different (often very small). 2 2a Petiolar base pulvinate. Abscission zone (abs) between leaflet and rachis Fig. 5.57f. Leaves not aromatic. Leaflets entire. (Lenticels transversely elongate, leaflets capable of autonomous motion). LEGUMINOSAE Very numerous species, see key D 2b Plant different. 3

Fig. 5.57

3a Spines in stipular position Fig. 5.58a.

RUTACEAE Harrisonia (AF-E, AS) 3b No spines or spines different. 4 4a Bark or leaves aromatic. 5 5a Petiolules (p) distally swollen Fig. 5.58b. BURSERACEAE Canarium pp. (AS) 5b Petiolules not distally swollen. RUTACEAE Zanthoxylum 4b Plant not aromatic. 6 6a Leaflets toothed. 7 7a AF. Leaflets sessile with numerous parallel veins. Stipule adnate to the petiole. ROSACEAE Hagenia (AF-E) 7b Characters different. 8 8a Leaflets with alternate or subopposite arrangement Fig. 5.58c. (Base of petiolule delimited by an abscission zone). Bark bitter. SIMAROUBACEAE Picrasma (AS) 8b Leaflets opposite Fig. 5.58d. 9 9a Stipules intrapetiolar. (Rachis winged: Bersama). MELIANTHACEAE Bersama, Melianthus (AF) 9b Stipules different. (Petiolules without abscission zone Fig. 5.58e). 10 10a AM. Tree. STAPHYLEACEAE-TAPISCIACEAE Huertea (Andes, West Indies) 10b AM. Shrub or liana. SAPINDACEAE Paullinia (AM) 6b Leaflets entire. Young internodes angular or grooved Fig. 5.58f. Twigs brittle. (Liana). SAPINDACEAE e.g. Paullinia (AM)

Group 47—Leaves compound, palmate or 3-foliolate, stipulate; alternate leaf arrangement. ARALIACEAE, BIXACEAE, CAPPARACEAE, CARYOCARACEAE, EUPHORBIACEAE, LEGUMINOSAE, MALVACEAE, PHYLLANTHACEAE, URTICACEAE, VITACEAE

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Fig. 5.58

Caution! «compound palmate» leaves of Cecropia spp. (URTICACEAE) are in fact simple, deeply incised leaves. 1a Inner bark with a network of fibres Fig. 5.59a. (See 2a to 2c). 2a Petiole distally swollen Fig. 5.59b. Leaflets entire. Bark without orange-coloured exudate. (RAUH). MALVACEAE e.g. Bombax (AS, cult.), Cola (AF), Ceiba (AM, cult.) 2b Petiole not distally swollen, or bark with aqueous orange-coloured exudate. BIXACEAE Cochlospermum 2c Petiole not distally swollen. Stipules hood-like. Leaflets entire: Musanga, toothed: Myrianthus. URTICACEAE Musanga, Myrianthus (AF) 1b Bark without a network of fibres. 3 3a Latex. (Leaves 3- or 5-foliolate). EUPHORBIACEAE e.g. Hevea (AM, cult.), Joannesia (AM-S) 3b Bark and leaves without latex. 4 4a Leaflets entire. 5 5a Petiolar base pulvinate Fig. 5.59c. Leaves 3-foliolate. LEGUMINOSAE e.g. Dioclea (AM), Erythrina 5b Petioles different. 6 6a Petiolar base sheathing Fig. 5.59d. (Stipules intrapetiolar). ARALIACEAE e.g. Osmoxylon (AS, cult.), Steganotaenia (APIACEAE, AF-E). 6b Petiolar base different. Midrib ending in a mucro or a small indentation Fig. 5.59e. CAPPARACEAE e.g. Euadenia (AF), Crateva 4b Leaflets toothed or crenulate. (See 7a to 7d). 7a AM. Leaflets crenulate. (Bark resinous, translucent dots). CARYOCARACEAE Anthodiscus (AM) 7b AF. Leaves compound palmate. (Bark scaly). EUPHORBIACEAE e.g. Ricinodendron (AF) 7c AS. Bark with reddish exudate. (L. 3-foliolate). PHYLLANTHACEAE Bischofia (AS) 7d Liana bearing oppositifoliate tendrils Fig. 5.59f. VITACEAE e.g. Tetrastigma (AS, AU)

Fig. 5.59

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Group 48—Leaves compound, pinnate or bifoliolate, not stipulate; alternate leaf arrangement. ARALIACEAE, BERBERIDACEAE, CONNARACEAE, JUGLANDACEAE, KIRKIACEAE, LEGUMINOSAE, MORINGACEAE, OXALIDACEAE, PICRAMNIACEAE, PROTEACEAE, SABIACEAE, SAPINDALES (MELIACEAE, RUTACEAE, etc.), ZYGOPHYLLACEAE Leaves really compound? or phyllomorphic branches bearing stipulate simple leaves ? (See group 7: Phyllanthus). 1a Petiole base sheathing the stem Fig. 5.60a. 2 2a Bark or leaves aromatic. ARALIACEAE e.g. Sciadodendron (AM), Polyscias (Paleo) 2b Wood yellow (cut !). Plant not aromatic. BERBERIDACEAE Berberis sect. Mahonia (AM-C, AS) 1b Petiolar base not sheathing. (See 3a to 3d). 3a Leaves bifoliolate. CHAMPAGNAT. ZYGOPHYLLACEAE s.l. Balanites (AF, AS, India, Myanmar) 3b Leaves 2- or 3-pinnate Fig. 5.60b. Bark with peculiar smell (myrosine). (Inner bark with a network of fibres). MORINGACEAE Moringa (AF, AS, cult.) 3c Leaves bipinnate. No smell of myrosine. MELIACEAE Melia (AF, AS, cult.) 3d Plant not as in 3a, 3b or 3c (palmately compound Fig. 5.60c or once pinnate). 4 4a Abscission zone between leaflet and rachis Fig. 5.60d. 5 5a Bark aromatic or leaves with translucent dots. RUTACEAE e.g. Zanthoxylum 5b Bark not aromatic. No translucent dots. 6 6a Bark bitter. (See 7a to 7c). 7a Leaflets alternate Fig. 5.60e. PICRAMNIACEAE Picramnia (AM) 7b Leaves glabrous, imparipinnate with leaflets subopposite. (Inner bark with a network of fibres). SIMAROUBACEAE Picrasma (AM) 7c Leaves glabrous, paripinnate Fig. 5.60f. MELIACEAE Xylocarpus (AS, OC, mangrove) 6b Bark not bitter. (See 8a to 8d).

Fig. 5.60

8a Leaflets not coriaceous. Midrib sunken on the upper side of the lamina. Shrub or small tree (TROLL). OXALIDACEAE Averrhoa (AS, cult.) 8b Leaflets coriaceous. Midrib sunken Fig. 5.61a on the upper side of the leaflet. Tree or liana. (Young leaves with a rust-coloured indument). CONNARACEAE e.g. Agelaea (Paleo), Connarus

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8c Tree. Leaves large, coriaceous.

LEGUMINOSAE e.g. Alexa, Dipteryx (AM) 8d AF-S. Growth strongly rhythmic with series of shorter internodes. Leaves pinnate. Leaflets small, thin and toothed. KIRKIACEAE, Kirkia (AF-S) 4b Petiolules without an abscission zone Fig. 5.61b. 9 9a Inner bark with a network of fibres. External wood furrowed Fig. 5.61c. PROTEACEAE e.g. Roupala (AM), Grevillea (AS, AU, OC, cult.) 9b Inner bark or wood different. (See 10a to 10c). 10a Leaflets entire. Bark with a network of fibres. JUGLANDACEAE e.g. Juglans (AM, AS), Engelhardia (AS) 10b Leaflets entire, rarely toothed (e.g. Allophylus, Azadirachta, Garuga). SAPINDALES (see key C) ANACARDIACEAE, BURSERACEAE, MELIACEAE, RUTACEAE, etc. 10c AS. Petiolar base swollen Fig. 5.61d. (Leaflets toothed). SABIACEAE e.g. Ophiocaryon (AM), Meliosma (AS) Group 49—Leaves compound; alternate leaf arrangement; leaves not stipulate; leaves palmate or 3foliolate. ARALIACEAE, BIGNONIACEAE, CARICACEAE, CONNARACEAE, OXALIDACEAE, PICRODENDRACEAE, RUTACEAE, SAPINDALES (SAPINDACEAE) 1a Petiolar base sheathing Fig. 5.61e.

ARALIACEAE e.g. Cussonia (AF, MA, cult.), Schefflera 1b Petiole different. 2 2a Leaves with translucent dots Fig. 5.61f. (Aromatic smell). RUTACEAE e.g. Cusparia (AM), Vepris (AF, MA), Aegle (AS)

Fig. 5.61

2b Leaves without translucent dots. 3 3a Short shoots present (leaves clustered) Fig. 5.62a. Rachis or petiole winged Fig. 5.62b. (Some simple leaves also present). BIGNONIACEAE Crescentia (AM, cult.) 3b Short shoots absent or rachis/petiole not winged. 4 4a Two-ranked leaf arrangement. OXALIDACEAE Sarcotheca (AS) 4b All stems with spiral leaf arrangement. 5 5a Abscission zone between leaflets and rachis Fig. 5.62c. (See 6a to 6c). 6a Tree or potential liana. Twigs lenticellate Fig. 5.62d. Leaves 3-foliolate Fig. 5.62e. Upper side of lamina with sunken venation. CONNARACEAE Agelaea (Paleo) 6b AM. Twigs not lenticellate. Leaves 3-foliolate. (Bitter bark). PICRODENDRACEAE Picrodendron (AM: West Indies)

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6c Shrub. Rhythmic growth. Short twigs. Leaves 3-foliolate.

OXALIDACEAE Oxalis (e.g. AM) 5b Base of leaflets without an abscission zone. 7 7a Leaves palmately compound. Stems prickly Fig. 5.62f. (Latex). CARICACEAE Jacaratia (AM-C) 7b Leaves not palmately compound (s.t. trifoliolate). SAPINDALES e.g. SAPINDACEAE: Allophylus Group 50—Leaves compound, stipulate; opposite or whorled leaf arrangement. BRUNELLIACEAE, CARYOCARACEAE, CUNONIACEAE, LEGUMINOSAE, OCHNACEAE, STAPHYLEACEAE, ZYGOPHYLLACEAE 1a Leaflets toothed or crenulate, or venation secant. 2a Leaflets toothed with its V II ending in a tooth. 3a Large interpetiolar stipular scars. 3b Stipules different (thin). 4a AM. Leaves pubescent.

2 3 STAPHYLEACEAE e.g. Turpinia (AM, AS) 4 BRUNELLIACEAE Brunellia (AM)

Fig. 5.62

4b AM. Leaves glabrous. “Finger-print” venation Fig. 5.63a.

OCHNACEAE Froesia, Touroulia (AM-S) 2b Leaves or V II different. 5 5a Stipules interpetiolar Fig. 5.63b. CUNONIACEAE e.g. Weinmannia (AM, MA, AS, NZ) 5b Stipules not interpetiolar. Leaves with three crenulate leaflets. CARYOCARACEAE Caryocar (AM) 1b Leaflets entire and venation different. 6 6a AM. Plant of savanna or dry forest. Sympodial branches consisting of short modules. ZYGOPHYLLACEAE e.g. Guaiacum, Bulnesia (AM) 6b AM. Architecture different. LEGUMINOSAE e.g. Taralea, Platymiscium (AM) Group 51—Leaves compound pinnate with four or more leaflets, not stipulate; opposite or whorled leaf arrangement. BIGNONIACEAE, LAMIACEAE, MELIACEAE, OLEACEAE, RUTACEAE, SAPINDACEAE, VIBURNACEAE 1a Bark or leaves aromatic, or leaves with translucent dots Fig. 5.63c. RUTACEAE e.g. Flindersia (AS, AU, OC)

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1b Plant not aromatic. Leaves without translucent dots. 2a Young internodes quadrangular. Rachis winged (w) Fig. 5.63d.

2 LAMIACEAE Peronema (AS) 2b Young internodes not quadrangular or rachis not winged. 3 3a Base of leaflets with an abscission zone Fig. 5.63e. 4 4a Leaflets toothed. Young leaflet folding involute. VIBURNACEAE Sambucus (AM, AS) 4b Leaflets different. MELIACEAE e.g. Dysoxylum (AS) SAPINDACEAE e.g. Matayba oppositifolia (AM), Acer, Paranephelium (AS) 3b Base of leaflets without abscission zone Fig. 5.63f. (Lenticels raised). 5

Fig. 5.63

5a Leaves simply Fig. 5.64a or 2  compound Fig. 5.64b: Jacaranda, Millingtonia, CORNER: Colea. (Tree, shrub, or see keys to lianas). BIGNONIACEAE e.g. Markhamia (AF, AS), Kigelia, Spathodea (AF, cult), Tecoma (AM) Jacaranda (AM, cult.), Colea (MA), Millingtonia (AS) 5b AM, AS. (Buds distinctly visible). OLEACEAE Fraxinus (AM, AS), Schrebera (AF, AS) Group 52—Leaves compound palmate or 3-foliolate, not stipulate; opposite or whorled leaf arrangement. BIGNONIACEAE, LAMIACEAE, PICRODENDRACEAE, RUTACEAE, SAPINDACEAE 1a Leaves aromatic or with translucent dots.

RUTACEAE Raputia (AM-S) 1b Leaves not aromatic and without translucent dots. 2 2a Petiolules slightly swollen distally. BIGNONIACEAE e.g. Tabebuia (AM) 2b Petiolules different. 3 3a Twigs with raised lenticels Fig. 5.64c. (OLEACEAE: Schrebera, Paleo). BIGNONIACEAE e.g. Cybistax, Godmania, Parmentiera, Tabebuia (AM) 3b No such lenticels. (See 4a to 4c). 4a Suberized petiolar base or young internodes more or less quadrangular Fig. 5.64d, or leaves 3-whorled. LAMIACEAE e.g. Teijsmanniodendron (AS), Vitex 4b AM. EU. AS. Leaflets without an abscission zone. SAPINDACEAE e.g. Aesculus (AM, EU, AS, cult.), Billia (AM)

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4c AF. Leaves palmately compound or trifoliolate Fig. 5.64e. Leaflets with an abscission zone. PICRODENDRACEAE Oldfieldia (AF)

Fig. 5.64

6

Supplementary Keys

6.1

Key A—Tribes and Main Genera of the Euphorbiaceae s.l.

PHY—PHYLLANTHACEAE, ACA—Acalyphoideae, CRO—Crotonoideae, EUP—Euphorbioideae. Circumscription of tribes according to Webster [1], however Pandaceae, Phyllantaceae, Picrodendraceae and Putranjivaceae are here families of their own. 1a No white or opalescent latex, and no coloured exudate. 2 2a Some twigs with two-ranked leaf arrangement Fig. 6.1a. (Phyllomorphous twigs: Phyllanthus). 3 3a Branches with GU in monopodial series. Leaves without glands. 4 4a Venation pinnate Fig. 6.1b. 5 5a ROUX. V I ending in a minute apical mucro. Stipules minute. Underside of leaves glabrous, not glaucous beneath. PANDACEAE e.g. Galearia (AS), Microdesmis (AF, AS), Panda (AF) 5b One, or more, of these characters different. 6 6a Young internodes angular Fig. 6.1c. 7 7a ROUX. TROLL. Growth of plagiotropic branches weakly rhythmic. (Annular stipules: Suregada, underside of leaves glaucous f. s. spp.). PHY—Phyllantheae e.g. Securinega (AF, MA), Sauropus (AS) Glochidion (mainly AS, OC), Suregada (Paleo), Phyllanthus 7b Trunk with rhythmic growth. PHY—Amanoeae e.g. Amanoa (AM) 6b Young internodes subcylindrical. TROLL. V I not ending in an apical mucro, underside of leaves hairy f. s. spp.). (Usually V I protruding at upper side, but embossed in Cleistanthus). PHY—Antidesmeae e.g. Antidesma (Paleo), Thecacoris (mainly AF, MA) 4b Leaves tripliveined Fig. 6.1d. Leaves pubescent. TROLL. PHY—Antidesmeae e.g. Antidesma (AS) 3b Apical meristem abortion in each GU Fig. 6.1e or leaves with glands. 8 8a Venation pinnate. (exceptionally leaves with glands in Phyllanthus). 9

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 R. Keller, A Field Guide to Tropical Plant Families, https://doi.org/10.1007/978-3-031-05942-1_6

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9a Petiole distally pulvinate Fig. 6.1f.

Supplementary Keys

PHY—Aporuseae e.g. Aporosa (AS)

9b Petiole not distally pulvinate.

Fig. 6.1

10a Leaf margin entire (young internodes angular). (See 11a to 11d). 11a Stipules small. No peltate hairs. L. glabrous, coriaceous f. s. spp. (Phyllomorphic twigs: Phyllanthus, fimbrial vein Fig. 6.2a, TROLL, Bridelia, Cleistanthus, ROUX: Drypetes, Phyllanthus). PHY—Wielandieae, e.g. Savia (AM, AF, MA), Phyllantheae, e.g. Phyllanthus PHY—Antidesmeae, e.g. Antidesma (Paleo) PHY—Bridelieae, e.g. Bridelia, Cleistanthus (Paleo) PUTRANJIVACEAE, e.g. Drypetes 11b Stipules large. No peltate hairs. (TROLL). PERACEAE e.g. Chaetocarpus 11c Stipules annular. No peltate hairs. (PETIT ?) CRO—Gelonieae Suregada (AS) 11d Small peltate hairs. (MASSART). ACA—Pereae e.g. Pera (AM) 10b Leaves serrulate. (Young internodes angular.). Inflorescence oppositifoliate. (Leaves pellucid-punctate, Fig. 6.2b). PHY—Phyllantheae Suregada (AF, MA, AS) PUTRANJIVACEAE e.g. Putranjiva (AS) 8b Leaves tripliveined Fig. 6.2c. 12 12a Glands on the underside of the lamina. PHY—Phyllantheae e.g. Phyllanthus 12b No glands. Peltate hairs. Foliaceous stipules. CRO—Crotoneae Crotonoides? (AM, Guianas) 2b All stems with spiral phyllotaxy or leaves opposite Fig. 6.2d. 13 13a Leaves simple. Venation pinnate Fig. 6.2e. 14 14a Trunk and branches with strong rhythmic growth (scale-leaves) and ramification, i.e. RAUH’s, or AUBRÉVILLE’s model for Baccaurea. 15 15a Absence of stellate or peltate hairs. 16 16a Petiole distally pulvinate. (RAUH, AUBRÉVILLE). Two glands at base and upperside of the lamina Fig. 6.2f: Protomegabaria). PHY—Aporuseae e.g. Baccaurea (AS, OC), Maesobotrya, Protomegabaria (AF) 16b Petiole not distally pulvinate. (RAUH). PHY—Aporuseae e.g. Richeria (AM) PHY—Acalypheae e.g. Cleidion (AS)

6.1 Key A—Tribes and Main Genera of the Euphorbiaceae s.l.

97

Fig. 6.2

15b Minute peltate or stellate hairs. (See 17a to 17c). 17a Petiole distally pulvinate (Stilt-roots in forest species Fig. 6.3a). Branching monopodial (RAUH). PHY—Uapaceae, e.g. Uapaca (AF, MA) 17b Some GUs without apical meristem abortion. L. glabrous: Pycnocoma, L. hairy, petiole short: Ptychopyxis. PHY—Pycnocomeae e.g. Pycnocoma (AF, MA), Ptychopyxis (AS) 17c Petiole distally glandular. CRO-Crotoneae e.g. Croton (AS) 14b Absence of strong rhythmic growth. 18 18a Petiole distally pulvinate Fig. 6.3b, long enough for this character to be observed. 19 19a Leaves with glands Fig. 6.3c. (See 20a to 20c). 20a RAUH. L. entire. Underside of L. with glands at the junction between petiole and lamina. PHY—Spondiantheae Spondianthus (AF) 20b Base of lamina with two glands at its upper side. (See 21a to 21c). 21a Leaves entire. ACA—Caryodendreae e.g. Caryodendron (AM). 21b KORIBA. L. serrulate. Translucent dots. ACA—Acalypheae e.g. Mareya (AF) 21c RAUH. Red exudate Leaves serrulate. 2–4 glands at the junction between lamina and petiole Fig. 6.3d. CRO—Codieae e.g. Pausandra (AM) 20c Base of lamina with two glands on its underside. 22 22a Leaves toothed or crenulate Fig. 6.3e. ACA—Alchorneae e.g. Alchornea 22b Leaves entire. Shrub or small tree. Leaf margin entire or lobed (aromatic f.s.spp.). glands at the base of the lamina. Hairs simple, stellate or peltate Fig. 6.3f. CRO Crotoneae e.g. Croton 19b Leaves without glands. 23 23a Stellate hairs present. 24 24a Underside of L. almost white. ACA—Chrozophoreae e.g. Sumbaviopsis (AS) 24b Underside of L. different. ACA—Epirineae e.g. Cephalomappa (AS)

Fig. 6.3

98

6

Supplementary Keys

23b Leaves more or less glabrous, no stellate hairs. 25 25a Leaf margin entire. Petiole grooved Fig. 6.4a. V. camptodromous. PERACEAE e.g. Pogonophora (AM, AF) 25b Leaf margin entire or serrulate. ACA—Cheiloseae e.g. Neoscortechinia (AS) 18b Petiole not distally pulvinate or pulvinus very short. 26 26a No peltate-stellate hairs. (See 27a to 27c). 27a Petiole long. (RAUH). ACA—Acalypheae e.g. Claoxylon (Paleo), Acalypha 27b Petiole short. L. entire. Thorny shrub. (CHAMPAGNAT). PHY—Phyllantheae e.g. Flueggea (Paleo) 27c Opposite phyllotaxy Fig. 6.4b. Leaf margin entire. PICRODENDRACEAE Mischodon (AS) 26b Peltate or stellate hairs Fig. 6.4c. 28 28a AM. Young internodes angular. Indumentum not orange coloured. 29 29a L. entire. No glands. ACA—Pereae Pera (AM) 29b L. serrulate. Glands on the underside. ACA—Alchorneae e.g. Alchornea 28b AS. Young internodes subcylindrical. Orange peltate hairs PHY—Hymenocardieae Hymenocardia (AS) 13b Venation palmate Fig. 6.4d, or leaves (faintly) trinerved or compound. 30 30a Twining lianas (glabrous, pubescent or hairy). ACA—Plukenetieae e.g. Dalechampia, Plukenetia (AM), Cnesmone (AS), Tragia 30b Plant not twining. 31 31a Petiole distally pulvinate Fig. 6.4e. 32 32a Leaves with glands Fig. 6.4f. 33 33a Glands at the base and the upper side of the lamina. (See 34a to 34c). 34a Stellate hairs. ACA—Caryodendreae e.g. Discoglypremna (AF) 34b Small peltate hairs. (KORIBA). ACA—Acalypheae e.g. Mallotus (AS)

Fig. 6.4

34c No stellate hairs. (KORIBA).

ACA—Alchorneae e.g. Conceveiba (AM)

6.1 Key A—Tribes and Main Genera of the Euphorbiaceae s.l.

99

33b Glands at the base and the underside of the lamina (glands disposed between the veins) Fig. 6.5a. ACA—Alchorneae e.g. Alchornea 32b Leaves without glands. 35 35a Phyllotaxy opposite (s.t. slight anisophylly), (Fig. 6.5b). Minute stellate-or peltate hairs (CHAMPAGNAT). ACA—Acalypheae e.g. Hancea (MA, AS), Mallotus (Paleo) 35b Phyllotaxy alternate. PHY—Antidesmeae e.g. Hieronyma (AM) 31b Petiole not distally pulvinate. 36 36a Leaves simple. 37a Leaves with glands. 38 38a L. serrulate or entire, more or less peltate in Macaranga and Mallotus. ACA—Acalypheae e.g. Macaranga, Mallotus (Paleo) 38b L. entire. (Orange peltate hairs f. s. spp., really no opalescent latex?). CRO—Crotonodeae e.g. Croton 37b Leaves without glands. (See 39a to 39c). 39a Leaf margin entire. RAUH (Fig. 6.5c). PHY—Antidesmeae Hieronyma (AM) 39b Leaves serrulate or entire, more or less peltate in Macaranga and Hancea. Peltate hairs: e.g. Hancea. Giant herb: Ricinus. ACA—Acalypheae e.g. Macaranga, Ricinus (Paleo), Acalypha 39c Leaves serrulate, whitish beneath, stellate hairs. (RAUH). ACA—Epiprineae Cladogynos (AS) 36b Leaves trifoliolate Fig. 6.5d. (RAUH). PICRODENDRACEAE Piranhea (AM) 1b Latex white or opalescent, or coloured aqueous or latex-like exudate. 40 40a Latex or exudate other than white (s.t. whitish-opalescent), or petiole distally pulvinate (Fig. 6.5e), or leaves tri- or palmately nerved (Fig. 6.5f). 41 41a Leaves simple. Venation pinnate. 42

Fig. 6.5

42a Latex clear, uncoloured. Petiole not distally pulvinate. (See 43a to 43c). 43a (No stipules f. s. spp.). CRO—Codieae e.g. Blachia (AS), Codiaeum (AS, AU, OC), Ostodes (AS) 43b Leaves serrulate, shortly petiolate. (with glands at base: Gymnanthes, short shoots modified into spines f. s. spp. Fig. 6.6a). EUP—Hippomaneae e.g. Sebastiania (AM), Gymnanthes (e.g. AM)

100

6

Supplementary Keys

43c Leaf margin entire, long-petiolate.

CRO—Jatropheae e.g. Jatropha 42b Coloured exudate or white latex, or petiole distally pulvinate. 44 44a Leaves without glands. 45 45a Latex white. Petiole distally pulvinate with two distal glands Fig. 6.6b. CRO—Elateriospermeae e.g. Elateriospermum (AS) 45b Latex red. KORIBA, LEEUWENB. Petiole distally pulvinate. CRO—Codieae e.g. Sagotia (AM), Baloghia (AS, OC) 44b Leaves with glands Fig. 6.6c. 46 46a Two glands on the upper side and the base of the lamina Fig. 6.6d. 47 47a Exudate redddish. KORIBA. CRO—Trigonostemoneae e.g. Trigonostemon (AS) 47b Latex-like exudate? KORIBA. (L. glabrous: Grossera, peltate hairs: Crotonogyne). CRO—Aleuritideae e.g. Crotonogyne (AF), Grossera (AF, MA) 46b Glands disposed otherwise. 48 48a Two or four glands at the junction between petiole and lamina Fig. 6.6e. (RAUH). CRO—Codieae e.g. Pausandra (AM) 48b Petiole distally pulvinate. One gland at the base of the main vein. CRO—Jatropheae Vaupesia (AM) 41b Leaves compound (e.g. Hevea) or palmate venation or L. tripliveined Fig. 6.6f. 49 49a Twining liana. Latex opalescent or red. 50 50a Two glands at the base of the lamina or at the distal end of the petiole. ACA—Omphaleae Omphalea

Fig. 6.6

50b Glands disposed on lamina margin Fig. 6.7a. Petiole not distally pulvinate? (L. lobed). CRO—Aleuritideae Manniophyton (AF) 49b Shrub or tree. 51 51a Leaves without glands. 52 52a Tree with monopodial rhythmic growth, RAUH. Leaves palmately compound, its underside with stellate or peltate hairs. CRO—Ricinodendreae Ricinodendron (AF) 52b Characters different. (GUs with apical meristem abortion). 53

6.1 Key A—Tribes and Main Genera of the Euphorbiaceae s.l.

101

53a Branches orthotropic. (Petiole not distally pulvinate. 54 54a Stipules modified into ramified hairs. (LEEUWENBERG). CRO—Jatropheaee Jatropha (AM) 54b Stipules different. 55 55a Leaves more or less lobate Fig. 6.7b, margin entire. (LEEUWENBERG). CRO—Manihoteae Manihot (AM) 55b Leaves cordate Fig. 6.7c, entire. 56 56a Stellate-peltate hairs on both sides of the lamina. CRO—Aleuritideae e.g. Neoboutonia (AF) 56b No stellate-peltate hairs. CRO—Codieae e.g. Strophioblachia (AS). 53b Branches plagiotropic. (TROLL). Leaves glabrous. CRO—Aleuritideae e.g. Domohinea (MA) 51b Leaves with glands. 57 57a Glands disposed on the upper side of the leaf or on its margin. 58 58a Petiole (weakly) distally pulvinate. (RAUH). Two glands near the junction between lamina and petiole Fig. 6.7d. Leaves trifoliolate Fig. 6.7e: Hevea. CRO Micrandreae e.g. Hevea, Micrandra (AM). 58b Petiole not distally pulvinate. 59 59a Stipules modified into ramified hairs. Petiole with distal gland(s) Fig. 6.7f. CRO—Jatropheae e.g. Jatropha (AM) 59b Stipules different. (See 60a to 60c). 60a Two glands on the base and the margin of the lamina. CRO—Manihoteae e.g. Endospermum (AS, OC)

Fig. 6.7

60b Two glands at the base of the lamina. (Leaves lobed). CRO—Aleuritideae e.g. Aleurites (AS, OC) 60c Leaves palmatilobate with glandular teeth. Stellate hairs. CRO—Ricinodendreae e.g. Givotia (MA) 57b Glands disposed on the underside of the leaf. 61

102

6

Supplementary Keys

61a Hairs simple. Shrub or small tree with sympodial architecture Fig. 6.8a. Leaf margin entire, round shaped, diversely lobed or peltate. CRO—Jatropheae e.g. Jatropha 61b Stellate or peltate hairs, also with simple hairs f. s. spp. Shrub or small tree. RAUH, SCARRONE, LEEUWENBERG. Orange or red exudate. L. entire, serrulate, or lobed (aromatic f. s. spp.). Glands at the base of the lamina. CRO—Crotoneae e.g. Croton 40b Latex white Fig. 6.8b. Simple leaves with pinnate venation Fig. 6.8c. Petiole not distally pulvinate (all EUPHORBIOIDEAE and a few CROTONOIDEAE). 62 62a Venation camptodromous (but with an intramarginal vein f. s. spp.). 63 63a Two glands on the upper side of the main vein or two glands on the distal end of the petiole: AM. Tetrorchidium, or two glands on the upper side and the base of the lamina: AF. Tetrorchidium. CRO—Adenoclineae e.g. Tetrorchidium (AM, AF) 63b Two glands on the underside and the base of the lamina. EUP—Stomatocalyceae e.g. Pimelodendron (AS) 62b Venation brochidodromous Fig. 6.8d. 64 64a Leaves with glands Fig. 6.8e. 65 65a Leaf margin entire Fig. 6.8f. 66 66a Branches plagiotropic. 67 67a TROLL. Two glands at the base and the underside of the lamina. Branching not very rhythmic. EUP—Hippomaneae e.g. Maprounea (AM) 67b Two glands at the base and the underside of the lamina. Branching distinctly rhythmic. EUP—Hippomaneae e.g. Actinostemon (AM) 66b Branches orthotropic.

Fig. 6.8

RAUH, KORIBA. Two glands at the distal end of the petiole: Sapium, leaf roundshaped, two glands at the base and the underside of the lamina (glands disposed between the veins): Homalanthus. Glands appressed to basal veins: Balakata. EUP—Hippomaneae e.g. Balakata (AS), Homalanthus (AS, AU), Sapium 65b Leaves serrulate Fig. 6.9a. 68 68a Branches orthotropic. Two glands at the distal end of petiole Fig. 6.9b. 69 69a Latex clear. KORIBA (Fig. 6.9c), (trunk spiny). EUP—Hippomaneae e.g. Hura (AM) 69b Latex white. RAUH or KORIBA. EUP—Hureae e.g. Sapium

6.2 Key B—Tribes and Main Genera of the Malvaceae

103

68b KORIBA, one solitary gland on the upper side and the distal end of the petiole: Hippomane, 1–2 glands at base of lamina: Gymnanthes. EUP—Hippomaneae e.g. Hippomane, Gymnanthes (AM) 64b Leaves without glands. 70 70a Leaf margin entire. (L. crassulescent f. s. spp.). 71 71a Some twigs with alternate, distichous phyllotaxy (or nearly so), (Fig. 6.9d). 72 72a NOZERAN: Anthostema. PREVOST?: Dichostemma. EUP—Euphorbieae e.g. Anthostema (AF, MA), Dichostemma (AF) 72b MANGENOT Fig. 6.9e. L. crassulescent. EUP—Euphorbieae Euphorbia tithymaloides (AM, ornam.) 71b All twigs with spiral or opposite phyllotaxy Fig. 6.9f. 73 73a (FAGERLIND). EUP—Hippomaneae e.g. Senefeldera (AM) 73b RAUH, KORIBA, or other model. (Plant spiny or succulent, f. s. spp.). L. opposite: Chamaesyce. EUP—Euphorbieae Synadenium (AM, AF, MA), Chamaesyce, Euphorbia 70b Leaves serrulate. 74 74a Branches orthotropic. L. spiny, glabrous. (RAUH). EUP—Pachystromateae Pachystroma (AM, Brasil) 74b Branches plagiotropic. PREVOST: Excoecaria, NOZERAN: Mabea EUP—Hippomaneae Excoecaria (AF, AS), Mabea (AM)

Fig. 6.9

6.2

Key B—Tribes and Main Genera of the Malvaceae

Subfamilial delimitation of Malvaceae according to [2] 1a Leaves all simple. 2 2a Some twigs with two-ranked leaf arrangement Fig. 6.10a. 3 3a AS. Leaves with peltate hairs Fig. 6.10b. 4 4a Pinnate venation. Peltate hairs only on the upper side. Branching monopodial plagiotropic. (ROUX, MASSART). Helicterioideae e.g. Durio (AS, cult.) 4b AM. (TROLL?). Peltate hairs on both sides of the lamina. Grewioideae e.g. Mollia (AM) 3b No peltate hairs (hairs simple or stellate). 5 5a Leaves distinctly asymmetrical Fig. 6.10c. (TROLL, ROUX). 6

104

6

Supplementary Keys

6a Leaves very hairy or scabrous. (GUs with apical meristem abortion, domatia, f. s. spp.). (TROLL: Grewia, Colona, Luehea, MASSART: Desplatsia). Grewioideae e.g. Luehea (AM), Desplatsia (AF), Colona (AS) Grewia (Paleo) 6b Leaves not hairy or scabrous (but generally hairy). (See 7a to 7c). 7a Leaf margin entire or with some large teeth. (TROLL). Dombeyoideae e.g. Scaphopetalum (AF), Pterospermum (AS). 7b Leaves toothed. Petiole distally pulvinate Fig. 6.10d. Trunk becoming oblique: erect form of a TROLL’s model). Byttnerioideae e.g. Guazuma (AM) 7c Leaves crenulate. Petiole distally pulvinate. (TROLL) V II secant. Grewioideae e.g. Schoutenia (AS) 5b Leaves symmetrical or bases faintly asymmetrical Fig. 6.10e. 8 8a Presence of GUs without apical meristem abortion. Fig. 6.10f. (Axillary flowering). (See 9a to 9c). 9 9a Leaf margin entire MASSART’s, NOZERAN’s or TROLL’s model. (Apex of lamina toothed, short petiole, f. s. spp.). Byttnerioideae MASS. or NOZ., e.g. Theobroma (AM, cult.), TROLL, e.g. Scaphopetalum (AF) Sterculioideae MASSART, e.g. Heritiera (AF, MA, AS), Helicterioideae, e.g. Helicteres (AS) 9b Leaves toothed or lobate with toothed margin. (TROLL).

Fig. 6.10

Helicterioideae-Grewioideae e.g. Helicteres (AS), Lueheopsis (AM) 9c Leaves palmately lobate. Stipule hood-like (Terminal flowering, MASSART). Sterculioideae, Triplochiton (AF, cult.) 8b GUs with apical meristem abortion. (Terminal flowering). 10 10a Trunk rhythmically ramified. (MASSART Fig. 6.11a, FAGERLIND Fig. 6.11b). 11 11a Leaf margin entire: Malvoideae, e.g. Matisia, Quararibea (AM) 11b Inflorescence oppositifoliate. Petiole not distally pulvinate. L. palmatilobate. Malvoideae: Fremontodendron (AM, California) 10b Branches regularly disposed on trunk (TROLL Fig. 6.11c, PETIT). 12 12a TROLL: e.g. Apeiba, PETIT, spiny twigs: Triumfetta. (Base of L. with domatia, f. s. spp.). Grewioideae e.g. Apeiba (AM), Grewia (Paleo), Triumfetta. 12b PETIT’s model. Leaves serrulate. Helicterioideae: e.g. Helicteres (AM) 2b All stems with spiral phyllotaxy (or monocaulous plant). 13 13a Climbing plant (AM: weakly prostrate plant with prickly twigs, AS: twining plant). Byttnerioideae, Byttneria (AM, AS) 13b Tree or shrub. 14 14a Petiole not distally pulvinate (or faintly enlarged or twisted). (See 15a to 15d) 15 15a RAUH’s model. (V II ending in a tooth: Dombeya, one large domatium at underside of leaf base: Hampea). Malvoideae: e.g. Hampea (AM), Hibiscus (mainly AS, ornam.) Dombeyoideae, Dombeya (AF, MA)

6.2 Key B—Tribes and Main Genera of the Malvaceae

105

15b SCARRONE Fig. 6.11d. Sterculioideae, e.g. Hildegardia (e.g. AF) Grewioideae, e.g. Heliocarpus (AM) 15c KORIBA Fig. 6.11e. Sterculioideae e.g. Firmiana (Paleo) Malvoideae, e.g. Ochroma (AM), Kydia (AS) Byttnerioideae, e.g. Kleinhovia, Eriolaena (AS) 15d PETIT Fig. 6.11f. (Leaves with glands). Malvoideae-Helicterioideae e.g. Gossypium (mainly AM, cult.), e.g. Ambroma (AS, AU) 14b Petiole distally pulvinate. 16 16a Branches plagiotropic by apposition. Axillary, not terminal, flowering. Sterculioideae e.g. Pterocymbium (AS, OC), Sterculia

Fig. 6.11 16b Branches not plagiotropic by apposition, or flowering terminal. 17 17a Branching monopodial Fig. 6.12a. 18 18a Branches erect, orthotropic growth (RAUH Fig. 6.12b). 19 19a L. hairy beneath, glaucous or ferrugineous aspect. (L. entire or lobate, heterophylly: the shape of L. variable, f. s. spp.). Sterculioideae e.g. Sterculia? (AM), Cola (AF) 19b Leaves (sub)glabrous, shiny. Bombacoideae e.g. Catostemma (AM). 18b Branches plagiotropic. (MASSART). (See 20a to 20c). 20a Leaves serrulate, V II ending in a tooth. Mansonia (AF, AS), Sterculioideae? 20b Leaves different. Short shoots bearing cordate leaves (Fig. 6.12c). e.g. Berrya (AS), Brownlowioideae. 20c Leaves not cordate, scaly hairs. e.g. Heritiera (AS), Sterculioideae? 17b GUs with apical meristem abortion (terminal flowering Fig. 6.12d). 21 21a (SCARRONE, leaves often broad, turning yellow before fall, L. entire: Brownlowia, Leaves serrulate: Clappertonia). Brownlowioideae e.g. Clappertonia (AF), Brownlowia (AS, OC) 21b (KORIBA). 22 22a AM. Leaves cordate or quite broad. Bombacoideae? Cavanillesia (AM) 22b AS. Leaves oval with cuneate base. Helicterioideae Neesia (AS) 1b Leaves palmately compound (some simple leaves also present f. s. spp.). 23 23a Branches plagiotropic by apposition (AUBRÉVILLE Fig. 6.12e). 24 24a (Buttressed tree, or stilt-roots). Sterculioideae, e.g. Heritiera, Sterculia (Paleo). 24b (Trunk swollen, often spiny). Bombacoideae e.g. Pseudobombax (AM), Bombax (AF, AS, cult.)

106

6

Supplementary Keys

23b Branches not plagiotropic by apposition. 25 25a Branches plagiotropic with spiral phyllotaxy. (Trunk spiny f. s. spp.). Branching of trunk and branches rhythmic (MASSART Fig. 6.12f). Leaflets serrulate: Ceiba pp. Bombacoideae e.g. Ceiba, Catostemma, Pachira (AM), Adansonia (AF, MA, AU), Bombax (AS) 25b Branches more or less erect. (i.e. RAUH’s model if trunk is monopodial and rhythmically branched). 26 26a AM. Bombacoideae, e.g. Eriotheca, Gyranthera, Pachira (AM). 26b AF. (Trunk with flattened buttresses, leaves with tertiary veinlets secant on the margin). Sterculioideae, e.g. Cola (AF)

Fig. 6.12

6.3

Key C—Families of the Order Sapindales + Picramniaceae

When tasting bark, caution!! Some Anacardiaceae are toxic or cause blisters. 1a Leaves stipulate. 2 2a Mainly AM. Leaves bipinnate Fig. 6.13a. Self-supporting form of a liana. SAPINDACEAE e.g. Paullinia (AM) 2b Leaves simply pinnate. (See 3a to 3c). 3a Bitter bark. Leaflets entire or toothed (Picrasma quassioides). Rhythmic growth. SIMAROUBACEAE Picrasma (AS, OC) 3b No bitter bark. (Basal leaflets resembling stipules in Garuga spp.). BURSERACEAE e.g. Canarium (e.g. AS), Garuga (AS, AU, OC) 3c Spines in stipular position. Rachis winged Fig. 6.13b. SIMAROUBACEAE Harrisonia (AS) 1b Leaves not stipulate (in some cases with small leaflets at the base of the rachis or with a pair of basal spines in stipular position, f. s. spp.). 4 4a Leaves with translucent dots (use a hand lens and observe against the light). (Crumpled leaves have an aromatic or peculiar smell). Several MELIACEAE have also translucent dots but these are very small. 5 5a Leaves compound. 6 6a Spiral leaf arrangement. RUTACEAE L. paripinnate Fig. 6.13c: e.g. Zanthoxylum L. imparipinnate Fig. 6.13d: e.g. Pilocarpus (AM), Murraya (AS, ornam.) L. trifoliolate Fig. 6.13e: e.g. Amyris, Monniera (AM), Vepris (AF), Toddalia (AS) L. compound palmate Fig. 6.13f: e.g. Casimiroa (AM-C), Vepris (Paleo). 6b Opposite leaf arrangement. RUTACEAE L. pinnate: e.g. Euodia, L. trifoliolate: e.g. Balfourodendron (FAGERLIND). L. palmate: e.g. Myllanthus, Balfourodendron, Myllanthus (AM), Euodia (Paleo)

6.3 Key C—Families of the Order Sapindales + Picramniaceae

107

5b Leaves simple. 7 7a Alternate leaf arrangement. 8 8a Bark or leaves aromatic. (Bitter bark). Petiole winged or distally swollen: e.g. Esenbeckia, Toddalia, not winged: e.g. Angostura. Spines: e.g. Atalantia, Citrus. RUTACEAE e.g. Angostura, Esenbeckia (AM), Citrus (AS, cult.), Toddalia (AF, MA, AS)

Fig. 6.13

8b No aromatic smell. Bitter bark. Underside of lamina with glands.

SIMAROUBACEAE e.g. Samadera (Quassia) indica (AS)

7b Opposite leaf arrangement (Fig. 6.14a). Petiole distally enlarged or pulvinate, or with stipuliform sheathing base. RUTACEAE e.g. Metrodorea (AM), Tetractomia (AS), Sarcomelicope (OC) 4b Leaves without translucent dots. 9 9a AS, OC. Veins II abruptly curved near the margin Fig. 6.14b. (Leaves or bark aromatic or with a resinous smell: Mangifereae, Semecarpeae). ANACARDIACEAE e.g. Astronium (AM), Pentaspadon (AS, OC) Anacardium (AM, cult.), Gluta (MA, AS), Mangifera (AS, cult.) 9b AS, NG, OC. Veins II not curved near the margin. 10 10a Leaves simple. (See 11a to 11c). 11a V II or V III secant at the margin: e.g. Ozoroa. Fimbrial vein: e.g. Holigarna (vesicant !). V. brochidodromous, translucent exudate in bark: e.g. Buchanania. ANACARDIACEAE e.g. Ozoroa (AF), Holigarna (AS), Buchanania (AS, AU) 11b AS, OC. Branches plagiotropic by apposition. V. camptodromous. MELIACEAE Vavaea (AS, OC) 11c Leaves entire, aromatic smell. V. brochidodromous. (Plant spiny). RUTACEAE Atalantia (AS) 10b Leaves compound (Fig. 6.14c). 12 12a Petiolules distally pulvinate, at least in terminal leaflet (Fig. 6.14d). (Essential oils or resins in many BURSERACEAE and RUTACEAE). 13 13a Bark or leaves resinous or petiolar base canaliculate (Fig. 6.14e). Venation brochidodromous. (Rhythmic growth: e.g. Crepidospermum, not rhythmic: e.g. Protium). BURSERACEAE e.g. Aucoumea (AF, cult.), Trattinickia (AM) Canarium (Paleo, ornam.), Dacryodes, Protium 13b AS. Bark and leaves not resinous. V. camptodromous (Fig. 6.14f). Petiole different. MELIACEAE Trichilia (e.g. AM), Walsura (AS) 12b Petiolules not distally pulvinate (s.t. only the terminal one in Tetragastris). 14 14a Petiolar base canaliculate or with a narrow raised edge. 15

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Fig. 6.14

15a V. brochidodromous (Fig. 6.15a). Rhythmic growth noticeable due to short internodes, or brittle monopodial branches, or resinous smell. Leaves imparipinnate. (Leaflets stipellate f. s. spp.). BURSERACEAE e.g. Bursera, Tetragastris (AM) 15b No resinous smell, bark aromatic f. s. spp. Quite supple twigs, L. paripinnate, V. camptodromous and growth not very rhythmic (e.g. Guarea, Trichilia). Leaves with a terminal leaflet, e.g. Trichilia. Rhythmic growth and L. bipinnate: Melia. MELIACEAE e.g. Guarea (AM, AF), Trichilia (AM, AF), Melia (Paleo, ornam.) 14b Petiolar base different, i.e. flat or convex (Fig. 6.15b). 16 16a Cut bark producing (aromatic) resin or crumpled leaves smelling of resin. 17 17a V II curved near the margin (Fig. 6.15c) or V. forming peripheral series of arches which almost touch the margin (marginal vein f. s. spp.). Rachis more or less winged: e.g. Schinus. ANACARDIACEAE e.g. Schinus (AM), Sorindeia (AF, MA). 17b Venation different (brochidodromous). 18 18a Branching sympodial. Petiolar base decurrent (Fig. 6.15d), rachis not canaliculate. ANACARDIACEAE e.g. Tapirira (AM), Sorindeia (AF, MA) 18b Branching monopodial with rhythmic growth, (s.t. short twigs): Commiphora). BURSERACEAE e.g. Tetragastris (AM), Commiphora (AF) 16b No smell of resin or of essential oils, but smell could be weakly aromatic in MELIACEAE and a few SAPINDACEAE. (Latex whitish or coloured f. s. spp.). 19 19a Latex whitish or coloured (caustic f. s. spp.). Latex-like resin in a few BURSERACEAE. (See 20a to 20c). 20a Latex sparse, milky white. Branches with growth weakly rhythmic (Fig. 6.15e) and bearing more or less spaced out leaves. Young branches stout and flexible, not rigid and brittle. MELIACEAE Trichilioideae: e.g. Aglaia, Chisocheton (AS, AU, OC) 20b Latex white or coloured, even brownish or turning black (Fig. 6.15f), (if necessary, wait to observe colour change) or venation as here. (Latex generally thick and sticky, toxic f. s. spp.). (Branching sympodial). Latex white-beige: e.g. Astronium, Rhus, Trichoscypha, or turning yellow: e.g. Thyrsodium, or black: e.g. Gluta, Loxopterygium. ANACARDIACEAE e.g. Astronium, Loxopterygium, Thyrsodium (AM), Trichoscypha (AF), Gluta (MA, AS), Rhus

6.3 Key C—Families of the Order Sapindales + Picramniaceae

109

Fig. 6.15

20c Latex white, in bark or in leaves. Young branches not flexible, but brittle and rigid. SAPINDACEAE e.g. Harpullia (AS, AU, OC) 19b Absence of latex. 21 21a Some leaves ending in a terminal leaflet (Fig. 6.16a). (Leaflets opposite: e.g. Simaba, Quassia (AM), alternate leaflets: e.g. Picramnia (AM), Simarouba (AM). 22 22a Venation as in (a), (b) or (c). ANACARDIACEAE (a) V II curved near the margin or secant (Fig. 6.16b), (at least at the apex of leaflets): e.g. Astronium (AM), Rhus. (b) Intramarginal vein (Fig. 6.16c): Spondias (AM, AS, cult.). (c) Small intercostal V II oriented ‘backwards’: e.g. Sorindeia (AF, MA), Thyrsodium (AM, AF). 22b Venation different. (See 23a to 23d). 23a Stipuliform spines: Harrisonia (East AF, AS). SIMAROUBACEAE 23b Base of petiolule with an abscission joint (Fig. 6.16d). PICRAMNIACEAE Alvaradoa, Picramnia (AM). 23c Leaflets glandulate. SIMAROUBACEAE e.g. Simaba (AM), Perriera (MA), Ailanthus (AS) 23d One of these characters different. 24 24a Bitter bark. (Axillary buds somewhat displaced from leaf axil f. s. spp. Venation not very visible). SIMAROUBACEAE e.g. Simarouba (AM), Odyendea (AF), Eurycoma (AS). 24b Bark not bitter. 25 25a Young branches brittle, breaking with a sharp cracking noise when bent. (See 26a to 26c). 26a Young internodes angular or grooved (Fig. 6.16e). If leaflets serrulate: Azadirachta (MELIACEAE). Leaflets alternate, e.g, Cupania, Matayba, Schleichera, Toechima, Toulicia, opposite, e.g. Cossinia, or L. trifoliolate: Allophylus. (Rachis winged f. s. spp.). SAPINDACEAE e.g. Toulicia (AM), Cossinia (MA, AU, OC), Schleichera (AS, cult). Lepisanthes (mainly AS), Toechima (NG,AU), Allophylus 26b Bark not aromatic. Growth weakly rhythmic. L. trifoliolate (Fig. 6.16f), generally serrulate: Allophylus SAPINDACEAE e.g. Allophylus 26c Bark aromatic or rhythmic growth. Leaflets entire. (Domatia: e.g. Trichilia). MELIACEAE Trichilia (e.g. AM), Sandoricum (AS, ornam.)

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Fig. 6.16

25b Young branches bending appreciably before breaking or young internodes cylindrical. Indumentum of scaly hairs: Aglaia, Lepidotrichilia, Trichilia. Leaflets alternate: e.g. Trichilia pp. Large leaves with stout petiole enlarged at base: Aphanamixis. Leaflets serrulate: Azadirachta. MELIACEAE e.g. Trichilia (AM), Aphanamixis (AS), Aglaia (AS, AU, OC) Azadirachta (Paleo, cult.) 21b Leaves not ending in a terminal leaflet (with a terminal pair of opposite leaflets (Fig. 6.17a), a terminal mucro or a bud of leaflets). (Leaflets opposite or alternate, leaves bifoliolate: a few SAPINDACEAE). 27 27a Foliar glands or red resin in bark. SIMAROUBACEAE Ailanthus (AS, AU) 27b No foliar glands, bark without red resin. 28 28a Bitter bark. 29 29a Underside of leaflets glaucous, with almost invisible venation. (Young twigs angular, leaflets alternate). SIMAROUBACEAE e.g. Simarouba (AM) 29b Leaflets different. Strong rhythmic growth. MELIACEAE e.g. Khaya (AF) 28b Bark not bitter and underside of leaflets different. 30 30a Leaves with indeterminate growth, ending in a terminal bud (tb) (Fig. 6.17b). MELIACEAE Trichilioideae: Guarea (AM, AF), Chisocheton (AS, OC) 30b Leaves different. 31 31a Young branches brittle and rigid. (See 32a to 32c). 32a Young internodes angular or grooved (Fig. 6.17c). Bark not aromatic. (Heterophylly: simple leaves in juvenile plant, f. s. spp.). Rachis winged: e.g. Harpullia (AS, AU, OC). Domatia: Arytera, Mischocarpus, Toechima (AS, AU, OC). L. bipinnate: Dilodendron (AM), Macphersonia (AF, MA). SAPINDACEAE Also: Melicoccus (AM), Blighia (AF, cult.), Nephelium (AS, cult.) numerous genera, better identified by their fruits. 32b Young internodes more or less cylindrical or with resinous smell. (V. brochidodromous (Fig. 6.17d), monopodial branching). BURSERACEAE e.g. Dacryodes (*Pachylobus) edulis (AF). 32c L. imparipinnate (i.e. leaflets opposite, (Fig. 6.17e). Growth weakly rhythmic (RAUH). ANACARDIACEAE e.g. Lannea (AF, AS) 31b Branches flexible, bending appreciably before breaking. 33 33a GUs with several scale-leaves separated by short internodes (Fig. 6.17f). (Bark aromatic). Bitter bark or garlic smell: e.g. Cedrela. MELIACEAE

6.4 Key D—Tribes and Main Genera of the Leguminosae

111

Fig. 6.17

Cedreloideae, e.g. Cedrela, Swietenia (AM) Entandrophragma (AF), Toona (AS, AU–N), Xylocarpus (Paleo mangroves) 33b Absence of strong rhythmic growth. MELIACEAE Trichilioideae, e.g. Aglaia (AS, AU, OC), Trichilia (e.g. AF)

6.4

Key D—Tribes and Main Genera of the Leguminosae

CSL—Caesalpinioideae, MIM—Mimosoideae, PAP—Papilionoideae. Systematics according to [2]. Note: “Serial buds” in leaf axils does not exclude the presence of some leaves with solitary buds. 1a Some twigs with two-ranked leaf arrangement (Fig. 6.18a). 2 2a Leaves simple or simply pinnate. 3 3a Leaves simple or with rachis ending in a terminal leaflet (Fig. 6.18b). 4 4a Leaflets opposite (at least the apical pair), (Fig. 6.18c). 5 5a AM. Buds solitary (Fig. 6.18d) or in a weak zigzag line Fig. 6.18e). Growth weakly rhythmic. (Rachis winged, leaves stipellate, red exudate, f.s.spp.). PAP—Swartziaeae Swartzia (AM) 5b Axillary buds solitary. Leaflets not stipellate. (Conspicuous scaly buds, translucent dots) PAP-Millettieae, e.g. Lonchocarpus (mainly AM) Cassieae (Dicorynia, AM) 4b Leaflets alternate or leaves simple 6 6a Liana or tree with L. bilobate or heart-shaped (Bauhinia). Tendrils: Bauhinia, hooked twigs, L. bilobate or tripliveined: Bandereia. Spiny stems: Spiny stems: Dalbergia, Machaerium spp. CSL—Dalbergieae Dalbergia, Machaerium (AM), Bandereia (AF) CSL—Cercideae (Griffonia, AF, Bauhinia) 6b Tree or shrub with leaves different. 7 7a Rachis ending in a leaflet or leaves simple with a pulvinate petiole. 8 8a AM. Leaves with translucent dots. 9 9a Serial buds arranged vertically (Fig. 6.18f). PAP—Sophoreae e.g. Myroxylon (AM). 9b AF, MA. Buds solitary. PAP—Swartzieae e.g. Cordyla (AF, MA).

Fig. 6.18

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Supplementary Keys

8b Absence of translucent dots. 10 10a Leaves simple. PAP—several tribes Red exudate in bark: Swartzieae (Swartzia, AM). Petiole pulvinate: Dalbergieae (Dalbergia ecastaphyllum, AM), Millettieae (Poecilanthe, AM), Swartzieae (Bocoa, AM), Sophoreae (Baphia, Paleo). Leaves serrulate, petiole not pulvinate, V. camptodromous: Phaseoleae, Flemingia (AS). 10b Leaves compound. Red exudate in bark: e.g. Machaerium, AM, Pterocarpus. Well-developed buds: e.g. Pterocarpus. Fimbrial vein: e.g. Machaerium, AM. Plant spiny: e.g. Dalbergia, Machaerium. PAP—Dalbergieae Leaflets subopposite. (GU in monopodial series). CSL—Cassieae e.g. Dicorynia (AM). 7b Leaves simple. Petiole not distally pulvinate. 11 11a Leaf margin entire. PAP—Dalbergieae e.g. Inocarpus (NG,OC) 11b Leaves serrulate. PAP—Swartzieae e.g. Lecointea, Zollernia (AM) 3b Leaves compound, with rachis not ending in a terminal leaflet (Fig. 6.19a). 12 12a AM. Rachis with cupuliform glands at the nodes of the rachis (Fig. 6.19b). (Serial buds present (Fig. 6.19c). MIM—Ingeae e.g. Inga (AM) 12b Rachis without cupuliform glands (s.t. with other kinds of glands or with glands disposed otherwise). 13 13a Plant climbing by means of tendrils (Fig. 6.19d) or spines (L. bilobate f. s. spp.). CSL—Cercideae Bauhinia 13b Plant different. 14 14a Stems with longitudinal ridges (Fig. 6.19e). Buds solitary. (Underside of leaflets glaucous). Gland at the base of the petiole: Chamaecrista, or on rachis: Senna). CSL—Cassieae e.g. Chamaecrista (AM), Senna, Cassia CSL—Caesalpinieae e.g. Poeppigia (AM) 14b Stems different (rarely more than one bud in leaf axils). 15 15a Leaflets not stipellate. (Intrapetiolar stipules, L. glandular, pp.). Leaves bifoliolate: e.g. Cynometra, Macrolobium, Peltogyne. Leaflets alternate (Fig. 6.19f): e.g. Crudia (AM), Kingiodendron (AS). L. with translucent dots: Colophospermum, Eurypetalum, Hymenaea, Peltogyne.

Fig. 6.19

6.4 Key D—Tribes and Main Genera of the Leguminosae

113

Serial buds in Hymenaea.

CSL—Detarieae/Amherstieae e.g. Brownea, Hymenaea, Macrolobium (AM) e.g. Anthonotha, Berlinia, Colophospermum, Eurypetalum, Gilbertiodendron (AF) Maniltoa (AS, OC), Amherstia (AS), Cynometra 15b Leaflets stipellate (Fig. 6.20a). 16 16a TROLL. Rachis ending in a short segment bearing no leaflets distally. Axillary bud a short condensed branch system. PAP—Robinieae e.g. Coursetia (AM) 16b Shrub. Leaves paripinnate. Branches decumbent, but extremities becoming erect. PAP—Robinieae e.g. Sesbania (AM, cult.) 2b Leaves bipinnate (Fig. 6.20b). (Serial buds). 17 17a Young internodes with marked longitudinal ridges and grooves. (One gland at the base of the petiole and, f. s. spp., at the point of leaflet insertion). MIM—Acacieae Acacia 17b Young internodes different. 18 18a Short shoots present (Fig. 6.20c). 19 19a Mostly AM. Rachis or petiole without glands. MIM—Ingeae e.g. Calliandra (AM,MA, AS, ornam.) 19b Mostly MA. Glands at the base of the petiole. MIM—Mimoseae e.g. Dichrostachys (MA, Paleo), Gagnebina (MA) 18b Short shoots absent. 20 20a Leaves with glands on the rachis or rachilla (Fig. 6.20d). 21 21a TROLL’s model but the branches turn to be erect. (Leaflets numerous, e.g. Cojoba). MIM—Mimoseae e.g. Newtonia (AF), Cojoba (AM) 21b Branches plagiotropic. (TROLL, Fig. 6.20e). MIM—Ingeae e.g. Enterolobium, Lysiloma (AM) 20b Leaves without glands. (See 22a to 22c). 22a Leaflets not thigmonastic. Rachis ending in a terminal pinna (Fig. 6.20f). (Axillary buds solitary). CSL—Caesalpinieae e.g. Caesalpinia (AM) 22b Leaflets not thigmonastic. No terminal pinnae. MIM—Mimoseae e.g. Zapoteca (AM)

Fig. 6.20

22c Leaflets thigmonastic.

MIM—Mimoseae e.g. Mimosa bimucronata (AM)

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Supplementary Keys

1b All stems with spiral leaf arrangement (Fig. 6.21a) or leaves opposite (or whorled) or branch GUs consisting of short modules with apical meristem abortion (Fig. 6.21b). (with only 1–2 well developed leaves). 23 23a Leaves opposite or whorled. Leaflets alternate to opposite, Venation camptodromous: Tipuana. PAP—Dalbergieae-Dipterygeae e.g. Platymiscium, Taralea, Tipuana (AM) 23b Alternate phyllotaxy. 24 24a Leaves simple or simply pinnate. 25 25a Leaves simple or existence of rachis with a terminal leaflet (Fig. 6.21c). (Several leaves should be observed). 26 26a Plant twining. (See 27a to 27c). 27a Some leaves with at least five leaflets. (Short shoots: Derris). PAP—Millettieae e.g. Millettia (Paleo), Derris 27b Leaves with no more than three leaflets. Leaflets stipellate (st). PAP—Phaseoleae e.g. Dioclea (mainly AM), Mucuna 27c Leaves simple. PAP—Sophoreae e.g. Bowringia (AF, MA) 26b Plant not twining. 28 28a Axillary buds solitary (Fig. 6.21d) or existence of short shoots. 29 29a AM, AF. Strong rhythmic growth noticeable due to the presence of short internodes. (Leaflets stipellate, short shoots: Dalbergia spp.). PAP—Dalbergieae e.g. Andira (AM, AF), Centrolobium, Vatairea (AM) Hymenolobium, Pterocarpus podocarpus (AM) 29b No strong rhythmic growth. 30 30a Leaves trifoliolate, stipellate (Fig. 6.21e) or with glands. (Plant spiny: Erythrina). PAP—Phaseoleae e.g. Butea (AS), Clitoria, Erythrina (ornam.) 30b Leaves neither trifoliolate, stipellate, without glands. 31 31a GUs without apical abortion (Fig. 6.21f). 32 32a Branches plagiotropic. 33 33a Bark without spermatic odour. PAP—Millettieae e.g. Millettia (AF), Lonchocarpus (mainly AM). 33b Spermatic odour in bark. PAP—Sophoreae e.g. Ateleia (AM)

Fig. 6.21

32b Branches erect or decumbent. 34 34a Branches decumbent (Fig. 6.22a). PAP—Robinieae e.g. Acosmium, Gliricida (AM)

6.4 Key D—Tribes and Main Genera of the Leguminosae

115

34b Branches erect.

PAP—Sophoreae e.g. Alexa (AM), Sophora 31b GUs with apical meristem abortion. (See 35a to 35d). 35a Leaflets alternate (Fig. 6.22b). PAP—Sophoreae e.g. Bowdichia (AM) 35b Phyllotaxy distichous in the branches of the young individuals. CSL—Cassieae e.g. Dicorynia (AM) 35c Leaflets opposite. PAP—Millettieae e.g. Poecilanthe (AM), Tephrosia 35d Leaf unifoliolate with articulate petiole. PAP—Millettieae e.g. Philenoptera (AF) 28b Serial buds and no short shoots. (Rhythmic growth noticeable due to the presence of short internodes). See (36a to 36d). 36a Serial buds arranged vertically (Fig. 6.22c). (Rachis grooved, young internodes angular. CSL—Sclerolobieae e.g. Campsiandra, Vouacapoua (AM) 36b Serial buds arranged vertically. Rachis not grooved. (Leaflets opposite). PAP—Sophoreae e.g. Clathrotropis, Diplotropis, (AM), Ormosia (AM, AS, AU) 36c Buds zigzagging. L. paripinnate. AP-Robinieae e.g. Sesbania (AM). 36d Buds collateral (Fig. 6.22d). L. trifoliolate. PAP—Millettieae e.g. Millettia (Paleo) 25b Leaves compound. Rachis not ending in a terminal leaflet (Fig. 6.22e) (ending in a mucro). 37 37a Leaflets opposite. 38 38a Plant twining (Fig. 6.22f). (Small liana). PAP—Abreae Abrus 38b Plant not twining. 39

Fig. 6.22

39a Serial buds (Fig. 6.23a). 40 40a Spines (short twigs): Gleditsia, subwhorled leaves disposed on very short twigs, venation secant: Haematoxylum. CSL—Caesalpinieae Haematoxylum (AM), Gleditsia (extratrop. AM, AS) 40b Characters different. (AM, serial buds, Strong rhythmic growth and occurrence of shorter internodes (Fig. 6.23b). 41 41a Young stems grooved. Pinnatifid stipules. (Some species housing ants, with hollow petioles: Tachigali). (SCARRONE, PETIT). CSL—Sclerolobieae e.g. Tachigali (AM)

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Supplementary Keys

41b Stems not grooved. (Red exudate in trunk). CSL—Dimorphandreae e.g. Mora (AM) 39b All axillary buds solitary (Fig. 6.23c). 42 42a Young internodes angular or with longitudinal ridges. (TROLL or CHAMPAGNAT). (Cupuliform, or nipple-like glands, underside of leaflets glaucous). CSL—Cassieae e.g. Cassia, Senna 42b Stems different, erect? Rachis without glands. PAP—Aeschynomeneae e.g. Amicia (AM) 37b Leaflets alternate or subopposite (Fig. 6.23d). AM. (Rachis winged, spirodistichous phyllotaxy: Coumarouna, or embossed venation: Dipteryx). PAP-Dipterygeae e.g. Coumarouna (*Dipteryx), (AM). 24b Leaves bipinnate (Phyllodes for some Australian Acacia). 43 43a Presence of buds disposed in two rows (Fig. 6.23e), (s.t., only two buds disposed more or less longitudinally). 44 44a Climbing or weakly prostrate plant. 45 45a Branching monopodial. (Spiny plant). If spiny: stem 5-sided with spines on the five sides. Leaves without glands and leaflets thigmonastic (see F5): Mimosa, L. with glands: Piptadenia, Mimosa myriadenia. Plant prostrate, not spiny, growing along streams: Entada. MIM—Mimoseae e.g. Piptadenia (AM), Mimosa, Entada 45b Branches distinctly sympodial (Fig. 6.23f). Leaves with glands. Leaflets not thigmonastic. Plant spiny. Stems angular or grooved, 10-sided (with spines on only five-sided). MIM—Acacieae Acacia

Fig. 6.23

44b Tree or self-supporting shrub. 46a Branches plagiotropic (TROLL, Fig. 6.24a). 47a Some GUs without apical meristem abortion. 48a Leaves with glands. (Young internodes angular).

46 47 48 MIM—Ingeae e.g. Samanea (AM) 48b Leaves without glands. MIM—Mimoseae e.g. Piptadenia (AM), Pseudoprosopis, Piptadeniastrum (AF) 47b GUs with apical meristem abortion (Fig. 6.24b). (See 49a to 49d). 49a Rachis grooved. Twigs angular with two decurrent ribs from petiole. Leaflets very numerous. (Spiny plant). MIM—Acacieae Acacia 49b Rachis grooved (Fig. 6.24c). Twigs not angular. Leaflets very numerous. Rachis furrowed with a basal gland. MIM—Mimoseae Anadenanthera (AM)

6.4 Key D—Tribes and Main Genera of the Leguminosae

117

49c Rachis convex in section. (Plant not spiny, leaflets not very numerous, pinnae stipellate). MIM—Ingeae e.g. Enterolobium (AM), Albizia (AF, AS) 49d Rachis convex in section (Fig. 6.24d). Developed axillary buds. (Short shoots modified into spines, leaflets not very numerous: Chloroleucon, leaflets few, but quite large: Zygia). MIM—Ingeae e.g. Chloroleucon, Zygia (AM) 46b Branches orthotropic. 50 50a Leaflets thigmonastic. (Spiny plant of open places, indument of thick hairs f. s. spp.). MIM—Mimoseae Mimosa 50b Leaflets not thigmonastic. 51 51a Existence of GU in monopodial series (Fig. 6.24e). (See 52a to 52d). 52a Leaves without glands. Leaflets alternate, and buds forming a zizagging series (Fig. 6.24f): Adenanthera. MIM—Mimoseae Tetrapleura, Adenanthera (AS, AU, OC) 52b Convex glands situated somewhat below the insertion of the pinnae, leaflets quite large and almost symmetrical. MIM—Ingeae Cedrelinga, Pseudosamanea (AM)

Fig. 6.24

52c Rachis with cupular glands, leaflets asymmetrical. MIM—Ingeae Leucaena, Piptadenia, Stryphnodendron (AM) 52d Rachis with sessile or stalked glands. Leaflets symmetrical. Short twigs modified into thorns. MIM—Mimoseae Dichrostachys (AF, MA) 51b Apical meristem abortions (Fig. 6.25a). (L. with glands). (See 53a to 53c). 53a Leaflets >5mm wide. (Short shoots f. s.spp. of Pithecellobium). MIM—Ingeae e.g. Pithecellobium, Zygia (AM) 53b Leaflets 100). MIM—Parkieae Pentaclethra (AF), Parkia

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Supplementary Keys

56b Leaflets large, opposite, not many (